UNA ENS RRA Fa OS PR e eS DA pl re 19 ME NEMA AIN IRON A AL NO de SNS SINN e ero absense aa Rowen Asia ARG Pta ve E a y N eae Red Tey I De i SER Te Hk \ De an ASIA in lan fe m EHEN SUN airy SER FERIEN NET ET ig baba fate de ow BERNER Pos SI ESEL ROTEN 5 rue hs Meat? y Lat dei cedo ; coins ain fi | Are : ps = a ny A phe) E 3 \ \ { i 3 ñ : A SD as) } N j my . oy Eon mS r y 7 ñ u y ar A 1 é 1 pod 7 . \ u | o o ‘ o 5 1 ' \ a 7 o ‘ . ; 7 o - R > nn . f o \ ' | N , Ñ : o 7 : ares i 7 ' 7 ' is 7 Pe in a { A) N O er a ‘ ‘ be a aut q Be I oe an 7 DS 1 QL | BAZ Nt Bonner zoologische Beitrage \keraries 5 Editor-in-Chief Michael Schmitt Bonn, Germany Editors Renate van den Elzen Bernhard A. Huber I n dex Gustav Peters Bradley Sinclair Dieter Stüning Volume 55 Bonn, ir AFFELD, Sven; WÄGELE, Heike; AVILA, Conxita; KEHRAUS, Stefan & KÖNIG, Gabriele M.: 181-190 Distribution of homarine in some Opisthobranchia (Gastropoda: Mollusca) Aupbisio, Paolo & DE BIASE, Alessio: 151-157 A new Meligethes of the M. pubescens species-group from South Africa (Coleoptera, Nitidulidae, Meligethinae) BARTSCH, Ilse: 47-59 The Freshwater Mite Porolohmannella violacea (Kramer, 1879) (Acari: Halacaridae), Description of Juveniles and Females and Notes on Development and Distribution DINAPOLI, Angela; TAMER, Ceyhun; FRANSSEN, Susanne; NADUVILEZHATH, Lisha & KLUSSMANN-KOLB, Annette: 191-202 Utility of H3-Genesequences for phylogenetic reconstruction — a case study of heterobranch Gastropoda DUBATOLOV, Vladimir « HOLLOWAY, Jeremy D.: 113-121 A new species of the Creatonotos transiens-group (Lepidoptera: Arctiidae) from Sulawesi, Indonesia FORSCHLER, Marc Imanuel & SIEBENROCK, Karl Heinz: 159-162 Morphological differentiation of mainland Citril Finches, Carduelis [atrine/la\ citrinella and insular Corsican (Citril) Finches, Cardvelis [citrinella) corsicanns GARCIA, Francisco J.; DOMINGUEZ, Marta; & TRONCOSO, Jesús S.: 203-222 Biogeographic considerations of the Opisthobranchia (Mollusca: Gastropoda) fauna from the Brazilian littoral and nearby areas GÖBBELER, Katrin & KLUSSMANN-KOLB, Annette: 223-229 Paddle cilia on the cephalic sensory organs (CSOs) of Opisthobranchia (Mollusca: Gastropoda) — genuine structures or artefacts? HANDELER, Katharina & WAGELE, Heike: 231-254 Preliminary study on molecular phylogeny of Sacoglossa and a compilation of their food organisms HAHN, Ingo: 95-100 Bioacoustic Characteristics and Population Numbers of Endemic Cinclodes oustaleti baeckstroemii (Aves: Furnariidae) Lónnberg, 1921 of Alejandro Selkirk Island, Chile HAHN, Ingo: 101-103 Biogeographical Isolation and Bioacoustics: the Juan Fernandez Firecrown, Sephanoides fernandensis (Aves: Trochilidae) (King, 1831), of Robinson Crusoe Island, Chile > HERRMANN, Hans-Werner; SCHMITZ, Andreas; HERRMANN, Patricia A. & BOHME, Wolfgang: 27-35 Amphibians and Reptiles of the Tchabal Mbabo Mtns, Adamaoua Plateau, Cameroon JENSEN, Kathe R.: 255-281 Biogeography of the Sacoglossa (Mollusca, Opisthobranchia) JÖRGER, Katharina M.; NEUSSER, Timea P. & SCHRÖDL, Michael: 283-290 Re-description of a female Pontohedyle brasilensis Rankin, 1979), a junior synonym of the Mediterranean P. wilaschemitchii (Kowalevsky, 1901) (Acochlidia, Gastropoda) KLEMP, Sverre: 61-71 Patterns of Geographic Variation in Body Measures and Plumage Colour of the Brimstone Canary Crithagra sulphurata MUNIAIN, Claudia; ARDILA, Néstor E. & CERVERA, Juan Lucas: 291-300 Plenrobranchaea inconspicua Bergh, 1897 (Opisthobranchia: Pleurobranchidae): Redescription and distribution from Argentina and Colombia NEUSSER, Timea P.; JORGER, Katharina & SCHRÖDL, Michael: 301-310 Exploring Cerebral Features in Acochlidia (Gastropoda: Opisthobranchia) PFEIFFER, Martin; SCHULTZ, Roland; RADCHENKO, Alexander; YAMANE, Seiki; 1-8 \VOYCIECHOWSKI, Michal; ULYKPAN, Aibek & SEIFERT, Bernhard: A Critical Checklist of the Ants of Mongolia (Hymenoptera: Formicidae) RHEINWALD, Götz: 3746 The Position of Trochiliphagns Carriker within the Ricinidae (Insecta: Phthiraptera) ROHWEDDER, Dirk; LAMPE, Karl-Heinz & SCHMIDT, Carola: 163-178 African Coleoptera Type Specimens collected by Thomas Wagner in the collection of the ZFMK SMOLIS, Adrian & SKARZYNSKI, Datiusz: 73-77 A New species of Psendachorntella Stach, 1949 (Collembola: Neanuridae) from Poland STAUBACH, Sid € KLUSSMANN-KOLB, Annette: 311-318 The cephalic sensory organs of Acteon tornatilis (Linnaeus, 1758) (Gastropoda Opisthobranchia) — cellular innervation patterns as a tool for homologisation VAN ROOSMALEN, Marc G. M.; FRENZ, Lothar; van HOOFT, Pim; DE IONGH, Hans H. & LEIRs, Herwig: 105-112 A New Species of Living Peccary (Mammalia: Tayassuidae) from the Brazilian Amazon VOIGLÄNDER, Karin: The Life Cycle of Lithobins mutabilis L. Koch, 1862 (Myriapoda: Chilopoda) WAGNER, Philipp & BOHME, Wolfgang: A new species of the genus Trape/ns Cuvier, 1816 (Squamata: Agamidae) from arid central Africa WAGNER, Philipp & BÖHME, Wolfgang: Herpetophauna Kakamegensis: The amphibians and reptiles of Kakamega Forest, western Kenya ZACHOS, Frank E.; OTTO, Marthe; HMWE, San San & HARTL, Günther B.: Populationsgenetische Untersuchungen zur Differenzierung des schleswig-holsteinischen Rehwildes (Capreolus capreolus Linnaeus, 1758; Artiodactyla: Cervidae) Buchbesprechungen / Book Reviews BRAUN, Monika & DIETERLEN, Fritz (Hrsg.): Die Säugetiere Baden-Württembergs (G. PETERS, Bonn) JUNKER, Thomas (2004): 158 88 Die zweite Darwinsche Revolution — Geschichte des Synthetischen Darwinismus in Deutschland 1924 bis 1950 (M. SCHMITT, Bonn) MERZ, B. (ed.) 2006: Phylogeny, Taxonomy, and Biology of Tephritoid flies (Diptera, Tephritoidea) (B. J. SINCLAIR, Ottawa) Publication dates: no. 1 30.01.2007, no. 2 28.08.2007, nos. 3/4 07.11.2007 Author Index AFFELD, Sven 181 ARDILA, Néstor E. 291 AUDISIO, Paolo 151 AVILA, Conxita 181 BARTSCH, Ilse 47-59 BÖHME, Wolfgang 27, 81, 123 CERVERA, Juan Lucas 291 DE BIASE, Alessio 151 DE IONGH, Hans H. 105 DINAPOLI, Angela 191 DOMÍNGUEZ, Marta 203 DuBaAroLov, Vladimir V. 113 FÖRSCHLER, Marc Immanuel 159 FRANSSEN, Susanne 191 FRENZ, Lothar 105 GARCIA, Francisco J. 203 GÖBBELER, Katrin 223 HAHN, Ingo 95, 101 HANDELER, Katharina 231 HARTL, Günther B. 89 HERRMANN, Hans-Werner 27 HERRMANN, Patricia A. 27 HMWE, San San 89 HOLLOWAY, Jeremy D. 113 JENSEN, Kathe R. 255 JÖRGER, Katharina M. 283, 301 KEHRAUS, Stefan 181 KLEMP, Sverre 61 KLUSSMANN-KKOLB, Annette 191, 223, 311 KONIG, Gabriele 181 LAMPE, Karl-Heinz 163 LEirs, Herwig 105 MUNIAIN, Claudia 291 NADUVILEZHATH, Lisha 191 NEUSSER, Timea P. 283, 301 OTTO, Marthe 89 PETERS, Gustav 158 PFEIFFER, Martin 1 RADCHENKO, Alexander 1 RHEINWALD, Götz 37 ROHWEDDER, Dirk 163 SCHMIDT, Carola 163 SCHMITT, Michael 88 SCHMITZ, Andreas 27 SCHRÖDL, Michael 283, 301 SCHULTZ, Roland 1 SEIFERT, Bernhard 1 SIEBENROCK, Karl Heinz 159 SINCLAIR, Bradley J. 104 SKARZYNSKI, Dartusz 73 SMOLIS, Adrian 73 STAUBACH, Sid 311 TAMER, Ceyhun 191 TRONCOSO, Jesús S. 203 ULYKPAN, Aibek 1 VAN HOOFT, Pim 105 VAN ROOSMALEN, Marc G.M. 105 VOIGTLÄNDER, Karin 9 WAGELE, Heike 181, 231 WAGNER, Philipp 81, 123 \VOYCIECHOWSKI, Michal 1 YAMANE, Seiki 1 ZACHOS, Frank E. 89 104 m HUNGS BZSZG museum KOENIG Herausgegeben vom BONNET | Forschungsmuseum 70010 oische er... 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Dr. Miguel Angel ALONZO-ZARAZAGA, Museo nacional, Ciencias Naturales, E-28006 Madrid: E-Mail: zarazaga(qWmncn.csis.es. Prof. Dr. Ulrike Aspöck, Naturhistorisches Museum, 2. Zoologische Abteilung (Insekten), Postfach 417, A-1014 Wien; E-Mail: ulrike.aspoeck(@nhm-wien.ac.at. Prof. Dr. Paolo AUDISIO, Universita di Roma “La Sapienza”, Dipl. Biol. Anim e dell’Uomo (Zool.), Viale dell’Universitä 32, 1-00185 Roma, Te.: +39 6-49914744; E-Mail: paolo.audisio@@uniromal.it. Prof. Dr. Aaron M. BAUER, Villanova University, Department of Biology, 800 Lancaster Avenue, Villanova, PA 19085-1699, USA, Tel: +1-610-519-4857, Fax: +1-610-519-7863; E-Mail: aaron.bauer(@villanova.edu. Dr. Júrgen HAFFER, Tommesweg 60, D-45149 Essen, Tel.: +49 201-710426; E-Mail: j.haffer@web.de. Dr. Jeremy D. HOLLOWAY, The Natural History Museum, Department of Entomology, Cromwell Road, London, SW7 5BD, U.K.: E-Mail: j.holloway(@nmh.ac.uk. Dr. Marion KOTRBA, Zoologische Staatssammlung, Diptera, Münchhausenstr. 21, D-81247 München, Tel.: +49 89-8107 147, Fax: +49 89-8107 300; E-Mail: marion.kotrba@zsm.mwn.de. Prof. Dr. Boris KRYSTUFEK, Slovenian Museum of Natural His- tory, P. O. Box 290, SI-1001 Ljubljana: E-Mail: boris.krystufek@uni-]j.si. Pro. Dr. Sven O. KULLANDER, Swedish Museum of Natural His- tory, Department of Vertrebrate Zoology, P. O. Box 50007, SE-104 05 Stockholm; E-Mail: sven.kullander@nrm.se. Prof. Dr. Steven PERRY, Rheinische Friedrich-Wilhelms-Univer- sitát, Institut fúr Zoologie, Poppelsdorfer Schloss, D-53115 Bonn, Tel: +49 228-73 3807; E-Mail: perry@uni-bonn.de. Dr. Wolfgang SCHAWALLER, Staatliches Museum für Naturkunde, Rosenstein 1, D-70191 Stuttgart, Germany, Tel.: +49 711-8936 221, Fax: +49 711-8936 100; E-Mail: schawaller.smns(@naturkundemuseum-bw.de. Dr. W. David Sissom, Dept. of Life, Earth and Environmental Sciences, W. Texas A. & M. University, WTAMU Box 60808, Canyon, Texas 79016, USA; E-Mail: dsissom@wtamu.edu. Dr. Miguel VENCES, Technische Universität Carolo-Wilhelmi- na, Zool. Inst., Abt. Evolutionsbiol., Mendelssohnstr. 4, D-38106 Braunschweig, Tel.: +49 531-391 3231, Fax: +49 531-391 3222; E-Mail: m.vences@tu-braunschweig.de. PD Dr. Heike WÄGELE, Rheinische Friedrich-Wilhelms-Univer- sität, Institut für Evolutionsbiologie und Ökologie, D-53121 Bonn, Tel.: +49 228 73 5159, Fax: +49 234-322 4114; E-Mail: hwaegele@evolution.uni-bonn.de. Dr. Erich WEBER, Eberhard-Karls-Universität, Zoologische Schausammlung, Sigwartstr. 3, D-72076 Tübingen, Germany; E-Mail: erich.weber@uni-tuebingen.de. | Bonner zoologische Beitrage | Band 55 (2006) Heft 1 Seiten 1-8 Bonn, Januar 2007 A Critical Checklist of the Ants of Mongolia (Hymenoptera: Formicidae) Martin PFEIFFER!), Roland ScHULTz2), Alexander RADCHENKO>), Seiki YAMANEY), Michal WOYCIECHOWSKP), Aibek ULYKPAN®) & Bernhard SEIFERT?) University of Ulm, Ulm, Germany ?)Ernst-Moritz-Arndt-University, Greifswald, Germany 3)Museum and Institut of Zoology PAS, Warsaw, Poland “Kagoshima University, Kagoshima, Japan 5)Jagiellonian University, Krakow, Poland 6)Mongolian National University, Mongolia "Staatliches Museum für Naturkunde Görlitz, Görlitz, Germany Abstract. Here we present a critical species list of the ants of Mongolia, that is based on current literature, previously unpublished taxonomic changes, the unpublished records of the zoological expeditions of Japanese-Mongolian and Ger- man-Mongolian teams in the years 1997, 1999 and 2003 and the results of a four year Polish-German-Mongolian coope- ration. We report on 68 species of 17 genera of ants that have been found within Mongolia: Camponotus, Cardiocondy- la, Cataglyphis, Crematogaster, Dolichoderus, Formica, Harpagoxenus, Lasius, Leptothorax, Messor, Myrmica, Plagio- lepis, Polyergus, Proformica, Tapinoma, Temnothorax and Tetramorium. Six species are new to Mongolia: Formica pres- silabris Nylander, 1846, Lasius gebaueri Seifert, 1992, Myrmica commarginata Ruzsky, 1905, Myrmica kamtschatica Kupyanskaya, 1986, Myrmica eidmanni Menozzi, 1930 and Myrmica taediosa (Bolton, 1995). Keywords. Asia, Mongolia, Formicidae, species list. 1. INTRODUCTION Mongolia occupies several natural zones where the Siber- ian taiga forest meets the Central Asian steppe and the Go- bi desert. Steep gradients of temperature and moisture show opposing trends and run contrary from North to South resulting in a vegetation sequence of forest, steppe, semi desert and desert that is considerably influ- enced by different soil conditions and altitudinal changes. This complex set of parameters, together with the large area of the country, creates a large variety of habitats and has a distinct influence on the biodiversity of the region (EMELJANOV & KERZHNER 1983; WALTER 1983; BRECKLE et al. 1994). Ants are a dominant part of the ground- dwelling Mongolian entomofauna, and form distinct com- munities in the different biomes and vegetation zones of this country (PFEIFFER et al. 2003). The ant fauna of Central Asia has been studied for more than a century (e.g., MOCSÁRY & SZÉPLIGETI 1901), how- ever, either these early expeditions did not occur on the territory of today’s Mongolia (RuzskY 1905; StITZ 1934; YASUMATSU 1940), or other authors had caste doubt on the validity of the determinations (e.g., DLUSSKY 1965 on Ruzsky 1915, and PISARSKI 1969a on FOREL 1904 and MocsaAry & SZÉPLIGETI 1901). Basic information on the Mongolian ant fauna has been gathered by the Hungari- an entomologist Dr. Z. KASZAB, whose collections from the 1960s were identified by DLUSsky (1965), DLUSSKY & PISARSKI (1970), PISARSKI (1969 a,b) and PISARSKI & KRZYSZTOFIAK (1981). DLUSSKy (1964, 1967, 1969), RAD- CHENKO (1994b,c, 1995a, 1997a, 2005) and SEIFERT (2000, 2003, 2004) provided additional information on the ant species composition of the Mongolian fauna and were fo- cused to the taxonomic editing of the species (for details see below). PFEIFFER et al. (2003, 2004) obtained the first ecological research data that revealed the biogeographi- cal patterns of the Mongolian ant fauna. Here we present a preliminary checklist of the ants of Mongolia based on a critical evaluation of older literature and on the results of our own expeditions. 2. MATERIALS AND METHODS Our study is based on altogether 2145 samples from 174 locations that were accumulated in Mongolia between 1997 and 2004 (Fig. 1). We collected ant specimens dur- ing several expeditions to the Gobi desert and the steppe and forest zones of this country. Additionally several hun- dred specimens of other collections were taxonomically evaluated by A. Radchenko and B. Seifert. i) Three German Mongolian expeditions were conducted by M. Pfeiffer and K. Ulykpan: the first together with L. Chimedregdzen from July to September 1997, the second in July/August 1999 and the third together with A. Ulyk- pan in July/August 2003. They collected 703 samples mostly from baits at 67 locations in 1997 and 1999, in- cluding 11 sites that have been sampled most intensively (see PFEIFFER et al. 2003), and about 200 samples of a larg- er investigation at altogether 37 locations in 2003. All specimens were collected from several North-South tran- sects between E95° and E118° longitude all over Mon- golia. M. Woyciechowski collected 262 samples from Myrmica nests within four years (1999, 2000, 2001, and 2002) in the forest steppe zone in the Hentii region (North Mongolia). Ants from nests were collected in all types of vegetation (MUHLENBERG et al. 2000) around and between three main locations stretched across ca 100 km along the 4gth parallel: Honin Nuga Research Station (N49%04*48”, E107°17’15”), Hot springs (N49°01’08”, El07*32*43”) and Ming River valley (N49%00*06”, E108°02°36”). The Martin PFEIFFER et al.: Checklist of the Ants of Mongolia Japanese-Mongolian expedition of Sk. Yamane and A. Ulykpan accumulated 2200 ant specimens from 233 sam- ples from colonies and general collection that were sam- pled in June and July 2003, in Bogd Han National Park, Hustai N.P., and Terelj N.P. (all Tuv aimag); in Zamyn uud (Dornogovi aimag) and in July 2004 in Bogd Han N.P.; in Honin Nuga, and in Han Hentii Mts. (Selenge aimag). R. Schultz sampled the western part of Mongolia in July and August 2003 on an expedition from Ulaanbaatar through Hangai Mts. to Hovd and the Mongolian Altai (Hovd aimag) and examined 255 nest samples collected from 47 sites. He added more data to our list by the eval- uation of 1) the collection of the University of Halle (77 samples from 7 locations), which was mainly due to the work of Mrs. Dr. A. Stubbe and 2) of the diploma thesis of A.-L. Lucau (2004), who collected 145 samples of the same three sites that had also been collected by M. Woy- ciechowski (Honin Nuga Research Station, Hot springs and Ming River valley) and was supervised by Prof. Dr. M. Muhlenberg, University of Göttingen. E 50 N Fig. 1. Map of Mongolia. Given are the positions of our sample graphical position of Mongolia (black) in the centre of Asia. North 100 km se! sites. The world map in the upper right corner shows the geo- Bonner zoologische Beitráge 55 (2006) 3 If not explicitly stated otherwise, the material that we have collected during these journeys was identified by Alexan- der Radchenko (genera Camponotus, Cataglyphis, Cre- matogaster, Messor, Myrmica, Plagiolepis, Proformica and Temnothorax), Bernhard Seifert (genera Cardiocondy- la, Formica, Lasius and Polyergus), Graham-W. Elmes (Myrmica) and the other authors. Collection details and localities for all species will be given in future publica- tions, in which we plan to map the distribution patterns of the main species. To compile the species list we compared our data with that in the literature (see Table 1 and references). These ref- erences were critically scrutinized by our taxonomic ex- perts (B. Seifert, A. Radchenko) and the validity of the nomenclature was checked by evaluating the latest taxo- nomic publications (BOLTON 1995, 2003; RADCHENKO 1994a,b, 1995b,c, 1997a,b; RADCHENKO et al. 2002; SEIFERT 1992, 2000, 2003, 2004). 3. RESULTS The resulting list of the Mongolian fauna comprises 68 species of ants of 17 genera (Table 1). After cross check- ing of literature we conclude that six species are new to the Mongolian fauna (R. Schultz, Sk. Yamane & M. Woy- ciechowski, unpubl. results): Formica pressilabris Nylander, 1846 Hangai mountain region, Arhangai aimag (province), Tset- serleg soum, ca. 12 km west of Tsetserleg, 1886 m, leg. R. Schultz, 02.08.2003, det. B. Seifert. Lasius gebaueri Seifert, 1992 Tuv aimag: Bogd Han N.P., 1550 m, leg. Sk. Yamane, 27.06.2003, det. B. Seifert. Mongolian Altay, Hovd aimag, ca. 8 km south of Hovd, ca. 20 km west of Hovd, 2021 m, leg. R. Schultz, 08.08.2003, det. B. Seifert. Myrmica commarginata Ruzsky, 1905 Hovd aimag, Durgun soum, Chonoharaihiin gol, Derris, 1154 m, leg. A. Stubbe, 27.08.2003, det. R. Schultz 8 A. Radchenko. Myrmica eidmanni Menozzi, 1930 Selenge aimag, western Han Hentii Mts, 3 km SW from Honin Nuga Research Station, near Eruu River (N49%03.81”, E107°16.14’), 930 m, 06.08.1999 and 8 km E from Honin Nuga Research Station, near Sharlan Riv- er (N48°57.70’, E107°04.26’), 970 m, 23.07.2000, both leg. M. Woyciechowski, det. G. W. Elmes & A. Rad- chenko. Selenge aimag, western Han Hentii Mts, Sharlan River, near Honin Nuga research station, 1000 m, leg. A.-L. Lu- cau, July 2001, det. A. Radchenko. Selenge aimag, western Han Hentii Mts, Sharlan River, near Honin Nuga research station, leg. Sk. Yamane, July 2004, det. Sk. Yamane. Myrmica kamtschatica Kupyanskaya, 1986 Selenge aimag, western Han Hentii Mts, 8 km E from Honin Nuga Research Station, near Sharlan River (N48°57.70’, E107°04.26°), 970 m, leg. M. Woyciechows- ki, 23.07.2000 and the same location on 6.07.2001, det. G. W. Elmes & A. Radchenko. Myrmica taediosa (Bolton, 1995) Selenge aimag, western Han Hentii Mts, 9 km SW from Honin Nuga Research Station, near Eruu River (N49°02.39°, E107°11.62°), 1045 m, leg. M. Woy- ciechowski, 28.07.2002, det. G. W. Elmes & A. Rad- chenko. Due to taxonomic changes and to our critical evaluation of the species lists that have been previously published, this first comprehensive list of the Mongolian ant fauna contains a changed species spectrum compared to the old- er literature. For example Lasius alienus (Förster, 1850) has been reported from Mongolia in four publications (DLUSSKY & PISARSKI 1970; PISARSKI 1969a,b; PISARSKI & KRZYSZTOFIAK 1981), however, regarding to SEIFERT (1992), this species is constricted to Europe, so the taxo- nomic position of these samples was doubtful. A recheck- ing of these specimens that had been collected by Kasz- ab and are housed in the Hungarian National Museum showed that they were most possibly specimens of Lasius cf. obscuratus, but definitively not of Lasius alienus (Sán- dor Csösz, Budapest, pers. comm. 2005). Lasius cf. ob- scuratus has been also sampled by PFEIFFER (2003) but due to the low number of specimens this determination is still uncertain and needs to be confirmed. Other records that have been previously published are al- so uncertain and may be a result of misidentifications (un- published results A. Radchenko): e.g., Camponotus her- culeanus herculeanus Linnaeus, 1758 (in DLUSSKY & Pis- ARSKI 1970), Cardiocondyla stambuloffi Forel, 1892 (in PISARSKI 1969b; PISARSKI & KRZYSZTOFIAK 1981, but see RADCHENKO 1995c and SEIFERT 2003), Myrmica bergi Ruzsky 1902 that was confirmed to be Myrmica divergens Karavaiev, 1931 (in PFEIFFER et. al. 2003, but see RAD- CHENKO et al. 2002), Myrmica saposhnikovi Ruzsky, 1903 (in PISARSKI 1969a,b, PISARSKI & KRZYSZTOFIAK 1981), that was proved to be M. pisarskii Radchenko, 1994 (see RADCHENKO 1994b, 1995b), and Myrmica schencki Viereck, 1903 (in PISARSKI & KRZYSZTOFIAK 1981), which 4 Martin PFEIFFER et al.: Checklist of the Ants of Mongolia Table 1. A critical list of the ant species that have been found in Mongolia, according to literature, to our expeditions and the ex- pertise of our taxonomic experts A. Radchenko and B. Seifert. a = STITZ (1934), b = HOLGERSEN (1943), c = DLusskY & PISARKI (1970), d = DLussky 1965, e = PISARSKI (1969a), f = PISARSKI (1969b), g = PISARSKI & KRZYSZTOFIAK (1981), h = RADCHENKO (1994b), y = RADCHENKO (1994c), k = RADCHENKO (1995a), | = BOLTON (1995), m = German-Mongolian expedition 1997; 1999 (collection M. Pfeiffer, unpublished), n = collection of Kawaguchi, o = Radchenko (1997a), p= Dubatolov (1998), q = collection of M. Woyciechwski (1999-2004), r = collection of R. Schultz (leg. A.-L. Lucau 2001, 2003), s= collection of R. Schultz (leg. University of Halle, Germany), t = Imai et al. (2003), u= Pfeiffer et al. (2003), v = German-Mongolian expedition 2003 (M. Pfeif- fer), w = German-Mongolian expedition 2003 (R. Schultz), x = SEIFERT 2003, y = Japanese-Mongolian expedition 2003 (Sk. Ya- mane), z = Japanese-Mongolian expedition 2004 (Sk. Yamane), | = SEIFERT (2004), 2 = RADCHENKO (2005), 3 = Personal collec- tion A. Ulykpan. Valid scientific names were obtained from BOLTON 1995, 2003; RADCHENKO 1994 a,b, 1995b,c, 1997 a,b; RAD- CHENKO et al. 2002 and SEIFERT 2000, 2003, 2004. Scientific name References Camponotus japonicus Mayr, 1866 b,¢,¢,8,6,8,7;2,3 Camponotus herculeanus sachalinensis Forel, 1904 CEL £,W,y,Z,3 Camponotus saxatilis Ruzsky, 1895 m,0,W,Z, Camponotus turkestanus André, 1882 f,g,S,U,V Cardiocondyla koshewnikovi Ruzsky, 1902 U,W,X Cataglyphis aenescens (Nylander, 1849) c,e,f,g,S,U, V,wW Crematogaster subdentata Mayr, 1877 g,u Dolichoderus sibiricus Emery, 1889 2 Formica aquilonia Yarrow, 1955 e,n,v,y, Z Formica candida Smith, 1878 c,d,e,f,g,n,r,s,u,v,w,y,z,1,3, Formica clara Forel, 1886 c,f,u,v,y Formica clarissima Emery, 1925 S,U,W Formica cunicularia Latreille, 1798 CLOS Formica exsecta Nylander, 1846 d,m,w,y,Z Formica forsslundi Lohmander, 1949 c,e,f,g,w, 3 Formica japonica Motschoulsky, 1866 t Formica kozlovi Dlussky, 1965 d,e,f,g,m,w,y,3 Formica lemani Bondroit, 1917 c,d,f,g,m,w,y,z,3 Formica lugubris Zetterstedt, 1838 f,w,y Z, Formica manchu Wheeler, 1929 c,d,e,f,g,l,m,n,s,w,y,z,3 Formica pisarskii Dlussky, 1964 d,e,f,g,v,w,y,3 Formica pratensis Retzius, 1783 el Formica pressilabris Nylander, 1846 Ww Formica sanguinea Latreille, 1798 c,d,e,f,g,s,u,y,z,3 Formica truncorum Fabricius, 1804 RUN Re Formica uralensis Ruzsky, 1895 c,d,e,f,g,.u,w,y,3 Harpagoxenus zaisanicus Pisarski, 1963! e,f,g,3 Lasius distinguendus (Emery, 1916) e,L3 Lasius flavus (Fabricius, 1781) b Lasius gebaueri Seifert, 1992 w,y,z, 3 Lasius niger (Linnaeus, 1758) b,c, fg Lasius przewalskii Ruzsky, 1915 8,W,y,2,3 Leptothorax acervorum (Fabricius, 1793) e,f,g,w,3 Leptothorax muscorum (Nylander, 1846) c,e,f,g,y,3 Messor aciculatus (Smith, 1874) e,f,u,y Messor excursionis Ruzsky, 1905 g Myrmica angulinodis Ruzsky, 1905 c,e,f,g,r,w,q,y,z, 3 Myrmica arnoldii Dlussky, 1963 Ce 2 TG y7.5 Myrmica commarginata Ruzsky, 1905 S Myrmica divergens Karavaiev, 1931 C,€,2,W,Q,y,Z,3 Myrmica eidmanni Menozzi, 1930 Q,1,Z Bonner zoologische Beiträge 55 (2006) 5 Myrmica forcipata Karavaiev, 1931 Myrmica kamtschatica Kupyanskaya, 1986 Myrmica kasczenkoi Ruzsky, 1905 Myrmica koreana Elmes, Radchenko & Kim 2001 Myrmica pisarskii Radchenko, 1994 Myrmica rubra (Linnaeus, 1758) Myrmica ruginodis Nylander, 1846 Myrmica sulcinodis Nylander, 1846 Myrmica taediosa Bolton, 1995 Plagiolepis manczshurica Ruzsky, 1905 Polyergus nigerrimus Marikovsky, 1963 Proformica buddhaensis Ruzsky, 1915 Proformica coriacea Kuznetsov-Ugamsky, 1927 Proformica jacoti (Wheeler, 1923) Proformica kaszabi Dlussky, 1969 Proformica mongolica (Emery, 1901) Tapinoma orthocephalum Stitz, 19342 Tapinoma sinense Emery, 1925 Temnothorax kaszabi (Pisarski, 1969) Temnothorax melleus (Forel, 1904) Temnothorax mongolicus (Pisarski, 1969) Temnothorax nassonowi (Ruzsky, 1895) Temnothorax serviculus (Ruzsky, 1902) Tetramorium armatum Santschi, 1927 Tetramorium concaviceps Bursakov, 1984 Tetramorium inerme Mayr, 1877 Tetramorium tsushimae Emery, 1925 e,f,g,1,w,9,3 q c,e,f,g,v,w,y,3 u,q h,s,u,w,q,y,3 b,k k,q €,1,1,0,3 q e,f,g,u p.u f,g,u f g,w cc g,u u f,u c,e,f,g,u,w,y ! This species is on the red list of Mongolia (http://www.redlist.org). 2 This species has been reported from South Mongolia, but it is unclear whether this place is now in the Peoples Republic of China. in fact is M. koreana Elmes, Radchenko & Kim 2001. Oth- er mistakes seem to be most probably 7. caespitum Lin- naeus, 1758 (in DLUSSky & PISARSKI 1970; PISARSKI 1969b), that may be Tetramorium tsushimae Emery, 1925 and also Tetramorium ferox Ruzsky, 1903 (in PISARSKI & KRZYSZTOFIAK 1981). Similarly, DLusskY & PISARSKI (1970) and PisarskI (1969b) reported about Formica "polyctena Forster, 1850 to occur in Mongolia’s forest steppe, however, this seems to be a misidentification of specimens of Formica aquilonia Yarrow, 1955 (B. Seifert, pers. obs.). Because of the cold winter the occur- rence of F. polyctena within Mongolia should be impos- sible. We excluded all suspicious records from our species list. Due to the failure to access type material of Formica sub- pilosa ruzskyi Dlussky 1965, only indirect assessment of the status of this taxon is possible. Most certainly this tax- on is conspecific with F. clarissima Emery, 1925 because any material known from Mongolia and Tibet seems to belong to one and the same species according to structur- al characters. Furthermore there is no indication that those pigmentation characters proposed by the DLussky (1965) for the differentiation of the Formica subpilosa subspecies pamirica Dlussky 1965, clarissima Emery 1925 (to which Dlussky applied the unnecessary replacement name ruzskyi Dlussky 1965) or litoralis Kuznetzov-Ugamsky 1926 could have any practical value. However, structur- al characters such as body morphometrics or setae counts computed in a discriminant analysis allow the separation of four Asian allospecies Formica subpilosa Ruzsky 1902, FEF. clarissima Emery 1925, F! litoralis Kuznetzov-Ugam- sky 1926 and F! pamirica Dlussky 1965 (B. Seifert, un- publ. data). According to the material investigated by us only F clarissima Emery 1925 could be confirmed for Mongolia. The possible occurrence of FE subpilosa in semideserts of S Mongolia, as extension of the population from Chinese Gobi desert, should be checked during fur- ther field studies. Similarly Cataglyphis aenescens roickingeri and C. aenescens tankrei have been syn- onymized with C. aenescens (Nylander, 1846) (RAD- CHENKO 1997b). 6 Martin PFEIFFER et al.: Checklist of the Ants of Mongolia Table 2. Valid names of Mongolian ants and their former names or junior synonymies that have been used in the older literature. Valid name Former name Cataglyphis aenescens (Nylander, 1846) Formica manchu Wheeler, 1929 F dlusskyi Bolton, 1995 Formica candida Smith, 1878 Temnothorax kaszabi (Pisarski, 1969) Temnothorax melleus (Forel, 1904) Temnothorax mongolicus (Pisarski, 1969) Temnothorax nassonovi (Ruzsky, 1895) Temnothorax serviculus (Ruzsky, 1902) Tetramorium tsushimae Emery, 1925 Cataglyphis aenescens roickingeri For C. aenescens tankrei For F. longiceps Dlussky, 1964 F" picea Nylander, 1846, sensu DLusskY 1967; sensu DLUSSKY & PISARSKI 1971, and other authors Fl transkaucasica Nassonov, 1889, sensu COLLINGWOOD 1979, and other authors Leptothorax kaszabi Pisarsk1, 1969 Leptothorax melleus Forel, 1904 Leptothorax melleus csikii Pisarski, 1969 Leptothorax mongolicus Pisarski, 1969 Leptothorax serviculus mongolicus Pisarski, 1969 Leptothorax nassanovi Ruzsky, 1895 Leptothorax serviculus Ruzsky, 1902 Tetramorium annectens Pisarski, 1969 Tetramorium jacoti Wheeler, 1927 In several cases the names of the species have been changed since that time when DLussky and PISARSKI iden- tified the ant species from KASZAB’s rich collection of the Mongolian fauna. For example Formica manchu Wheel- er, 1929 was formerly named F longiceps Dlussky, 1964 or E dlusskyi Bolton, 1995 (see SEIFERT 2000) or 7. tsushi- mae Emery, 1925 formerly considered as Terramorium an- nectens Pisarski, 1969 or Tetramorium jacoti Wheeler, 1927 (see BOLTON 1995). In the case of Formica candi- da Smith, 1878 this ant taxon was divided into two dif- ferent species with separate zoogeography (SEIFERT 2004): the western “Black Bog Ant” redescribed as Formi- ca picea Nylander, 1846, and Fl candida, which is found in East Siberia from the eastern Altaı mountains up to the Russian Far East, in Tibet, Mongolia, and North Korea. With these redescriptions the older name Formica tran- skauscasica Nassonov, 1889 that has been used for F! can- dida, e.g., in PFEIFFER et al. (2003), is invalid. Similarly some species of the genus Leptothorax have been trans- ferred to the genus Temnothorax (e.g., Temnothorax mon- golicus (Pisarski, 1969) or Temnothorax nassonovi (Ruzsky, 1895) (BOLTON 2003). We excluded all syn- onymies (see Table 2) from the list. 4. DISCUSSION For the first time we present a critical, tentative species list of the Mongolian Formicidae that includes all previ- ously recorded taxa. We also added six new species, F pressilabris, L. gebaueri, M. commarginata, M. eidman- ni, M. kamtschatica and M. taediosa to the ant species lıst of Mongolia. Up to now the genus Formica provides the highest number of species (18) in this list, while Myrmi- ca is represented by 14, Lasius, Proformica and Temnotho- rax by five species, each. The large number of Formica and Myrmica ants, that dominate the northern parts of Mongolia, may be a hint towards the higher productivity of these regions (forest, steppe) compared to the semi deserts and deserts in the southern country. However, the study of the Mongolian Formicidae is still going on, and we are expecting that more species will be added to the list within the next years (e.g., social parasitic ants), be- cause of new collections and/or changes in the taxono- mic system. Acknowledgements. We thank Prof. Dr. Kaman Ulykpan, Mon- golian National University, Ulaanbaatar, for his kind coopera- tion and help during the organization of our expeditions and al- so for the collection of some of the ant specimens. We are in- debt to Dorjiin Oldokh, Odna Odonchimeg, and to the late Dr. Losol Chimedregzen for their help during field work. We thank Ana-Lena Lucau, for the collection data of some Mongolian ant species. Prof. Dr. M. Woyciechowski’s and Ana-Lena Lucau’s data were collected during their stay in the Honin Nuga Research Station guided by Prof. Dr. Michael Múhlenberg, Georg-August- University, Góttingen, Germany and thanks to the financial sup- port of Georg-August-University of Gottingen. This Station was founded thanks to the cooperation of the Georg-August-Univer- sity of Gottingen and the National University of Mongolia, Ulaanbaatar. We are very grateful for this support of our study. We owe special gratitude for collection data to Assoc. Prof. Dr. Sadao Kawaguch1, Faculty of Agriculture, Kyushu University, Japan, and to Dr. Annegret Stubbe, Martin-Luther-University, Halle. We thank especially Sandor Csósz of the Hungarian Na- tional Museum for sending an actual, tentative identification of the specimens formerly described as Lasius alienus and Dr. G.W. Elmes for his help during the determination of Myrmica species. We appreciate an anonymous referee for his helpful comments on a former version of the paper. In 1999 the German-Mongo- lian expedition (M.P.) was generously funded by the “Arthur- von-Gwinner-Stiftung für naturwissenschaftliche Forschungsrel- sen”. Bonner zoologische Beiträge 55 (2006) 7 REFERENCES Botton, B. 1995. 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Contributions to the hymenopterous fau- na of Inner Mongolia and North China. Transactions of the Sapporo Natural History Society 16: 90-95. Authors’ addresses: Martin PFEIFFER (corresponding au- thor), Department of Experimental Ecology, University of Ulm, Albert-Einstein Allee 11, D-89069 Ulm, Germany; E-Mail: martin.pfeiffer@uni-ulm.de; Roland SCHULTZ, Zo- ological Institute & Museum, Ernst-Moritz-Arndt-Univer- sity, Johann-Sebastian-Bach-Str. 11-12, D-17487 Greifs- wald, Germany; Alexander RADCHENKO, Museum and In- stitute of Zoology, Polish Academy of Sciences, 64, Wilcza str., 00-679, Warsaw, Poland; Seiki YAMANE, De- partment of Earth and Environmental Sciences, Faculty of Science, Kagoshima University, Korimoto 1, Kagoshi- ma 890-0065, Japan; Michal WOYCIECHOWSKI, Institute of Environmental Sciences, Jagiellonian University, Gronos- tajowa 7, 30-387 Krakow, Poland; Aibek ULYkPAN, Mon- golian National University, Department of Ecology, Ulaanbaatar 46, P.O. Box 377, Mongolia; Bernhard SEIFERT, Staatliches Museum für Naturkunde Görlitz, Am Museum 1, D-02826 Görlitz, Germany. Received: 06.06.2005 Revised: 17.06.2005 Accepted: 01.07.2005 Corresponding editor: D. Stüning Bonner zoologische Beiträge Band 55 (2006) Heft 1 Seiten 9-25 Bonn, Januar 2007 The Life Cycle of Lithobius mutabilis L. Koch, 1862 (Myriapoda: Chilopoda) Karin VOIGTLÄNDER Görlitz, Germany Abstract. The post-embryonic development and life cycle of Lithobius mutabilis L. Koch, 1862 were studied. Data for stage analyses were obtained by laboratory breeding and continuous observations of field-collected specimens in capti- vity over more than five years. Developmental stages are described with respect to the following characters: number of legs (anamorphic stages only), head length and width, length and width of tergite 3, body length, biomass, number of co- xal pores, ocelli, and antennal articles. All characters were measured on living individuals under CO)-anaesthesia. Infor- mation concerning oviposition and egg development, onset of sexual dimorphism and maturity, stage duration and moul- ting activity, life span and mortality as well as observations on feeding behaviour are provided. The results are compa- red with those of other investigations on L. mutabilis and other lithobiid species. Keywords. Post-embryonic development, morphological characters, bionomy, Lithobiidae. 1. INTRODUCTION Centipedes are essential components of the predatory arthropod fauna. Because of their considerable suitabili- ty as indicators of ecological site conditions, Lithobiidae attract utmost attention. The use of species as biological indicators is based on the knowledge of its ecofaunistical behaviour, phenology and bionomical strategy. Studies on centipede bionomics are much rarer than those on milli- pedes. Life cycles and stage morphology are known from only 40 centipede species, among them some 20 Litho- biomorpha, whereas about 150 millipede species have been investigated. For practical reasons, attention should be paid firstly on common and frequent species, like Litho- bius mutabilis in Central Europe, which plays an impor- tant role as a dominant epigeic predator especially in woodlands. Further, this species has the advantage that its life cycle has already been the subject of studies by AL- BERT (1983a, b) and FRUND (1983) in western Germany (Solling, Steigerwald), using field-collected individuals as well as laboratory breeding. The availability of data on the life cycle of L. mutabilis from other localities allows pos- sible similarities and differences in pattern to be investi- gated. Comparisons are also made with the post-embry- onic development of other Lithobius species. Our knowledge on post-embryonic development of myr- iapods was outlined by VERHOFFF (1905, 1925) with the concepts of “anamorphosis” (moultings connected with in- creases of body segments and legs), “epimorphosis” (moultings cause only differentiations in certain charac- ters but keep constant numbers of body segments and legs) and “hemianamorphosis” (both developmental pathways in succession). All Lithobiomorpha develop by hemi- anamorphosis: a juvenile hatches with a small number of segments and legs and develops by a series of moults. At each moult there is an increase of segments and legs un- til a defined number is reached (anamorphic development). Further moults only lead to a growth in body size and a modification of various structures without increase of seg- ments (epimorphic development). This study is mainly focussed on the post-embryonic de- velopment (breeding biology, development, attainment of maturity and life span) of Lithobius mutabilis L. Koch, 1862. This has been done by laboratory breeding and cul- tures over several years. Description of stages, of varia- tion in some morphological characters (e.g., number of an- tennal articles, ocelli, coxal pores) and growth analyses occupies a large part of the present study. 2. MATERIALS AND METHODS 2.1. Field collections A total of 184 individuals of L. mutabilis were collected by hand in a deciduous woodland in the Neiße Valley near Görlitz through all seasons in the course of seven years: (1992) 1993-1997 (1998). The animals were transferred to the laboratory for measurement and further rearing. To ascertain the stage of freshly captured adults, it was some- times necessary to observe two or more moults to over- come the wide overlap of characters between stages. 10 Karin VOIGTLANDER: Life Cycle of Lithobius mutabilis 2.2. Laboratory rearing For rearing, I used closed plastic vessels measuring 6 x 12x 6 cm with a ground-layer of gypsum (no soil or leaf litter). The vessels were kept in a climate chamber at a con- stant temperature of 16 °C, which is the preferred tem- perature under natural conditions (BAUER 1955; PFLEIDER- ER-GRUBER 1986). They were examined every second day (youngest juveniles every day) in order to record all moults and egg-laying or to remove dead or sick individuals. At the same time the animals were fed (with collembolans, pieces of earthworms, enchytraeids, 7ubifex, and, addition- ally, green leaf pieces of indoor plants), vessels were cleaned and the gypsum layer was moistened. Normally, two to three field collected or laboratory-bred specimens that were individually distinguishable were kept in one vessel. Seventy-seven juveniles hatched under laboratory condi- tions. They were immediately isolated and transferred to other vessels, in order to prevent them from being eaten by the parents. 2.3. Measurements The following results are based on the analyses of all the material, regardless of whether it was laboratory reared or more or less freshly captured. Body-length, length and width of head and tergite 3, bio- mass (total numbers of body measurements see Table 1); numbers of coxal pores, antennal articles and ocelli were measured using a stereo-microscope with 12.8-40 x mag- nification (at um-precision) and an analytical balance (pre- cision 0.1 mg), respectively. Dead individuals were not weighed. All individuals were measured alive under CO>-anaesthe- sia. To check all characters, 3 to 4 gas treatments (each for some seconds only) were necessary. The CO,-method has two important advantages: firstly, it is not damaging, i.e., it is possible to follow the development of one indi- vidual by measuring it after each moult; secondly, meas- urements of the relaxed animal can be obtained in con- trast to measurements in alcohol, where body parts are contracted or curled up. However, using living material has the disadvantage of restricting measurements to “coarse” characters due to the short anaesthesia period. 3. RESULTS 3.1. Analysis of post-embryonic developmental stages Designation of stages. In the centipede literature there is still disagreement about counting and nomenclature of the stages defined by the moults. VERHOEFF (1905, 1925) called the 1st juvenile stage a “foetus”, the 2nd a “larva prima”, “larva secunda”, etc. Some authors (e.g., PFLUGFELDER 1932; DOHLE 1969, 1986) emphasized that the post-embryonic stages of centipedes are not true lar- vae. They neither have distinct larval characters nor do they live under ecological conditions different from those of the adults. Nevertheless, most of the authors continue Table 1. Total numbers of measurements taken at different sta- ges (BL — body length, HL — head length, HW — head width, T3L - length of tergite 3, T3W — width of tergite 3, BM — bio- mass). To be read in conjunction with Figure 1. Stage BL HL HW T3L T3W BM I 9 8 9 = I] 9 7 7 = Im 34 37 36 34 36 = IV 3d. 3 3 31 a V ee 3 SEUA 32 11 VI (1) 35 35 35 35 35 26 VIO) + m 18 53 18 17 9 19 20 20 20 18 14 VII) 9 14 16 16 16 16 13 9 i7 ir. u 17 15 IX(49 39 16 10 10 10 10 9 9 g 9 g 9 9 9 X (5) gs 21 30 30 28 28 25 9 14 15 15 14 14 13 XI(6) 3 40 44 43 43 43 41 9 19 24 24 24 24 23 XI(D) 8 37 4-46 44 45 43 o 36 3%. 32: 3 31 28 XII(8) Y 25 28 29 29 29 27 9 i? 2. 3 . 36 21 20 XIV(9) 3 14 14 14 14 14 14 o LB. 1 e $3 13 12 XV (10) 9 g 9 9 9 9 8 9 i ne E 48 13 13 XVI(11) a so 5 so 5 5 4 9 33 5 3 3 XVI (12) & 3 4 4 4 4 3 9 4 4 4 4 4 4 XVIII (13) 9 2 2 y $ 2 2 Q f° 4 4 4 4 XIX (14) 9 RA q E 3 2 Q 1 1 1 1 1 1 XX) oS 1 1 1 1 1 1 Sum 497 540 3555 543 548 405 Bonner zoologische Beiträge 55 (2006) 11 Fig. la. body length [mm] Fig. Ib. biomass [mg] II IV VI VIE X XI XIV XVEXVIIE XX I IV VI VIE XX XI XIV XVEXVIN XX stage stage Fig. Ic. head length [mm] Fig. 1d. head width [mm] [597 I IV VI Vili XX NIE XIV XVIXVIM XX iI IV VI VH X NTE XIV XAVIXVII XX stage stage Fig. le. tergit3 length [mm] Fig. If. tergit3 width [mm] ll IV VI VOL X XIE XIV XVEXVHE XX 11 IV VI VU XX XI XIV XVIXVII XX stage stage Fig. 1. Averages of body and biomass measurements of L. mutabilis. For numbers of measured specimens see Table 1; grey cir- cles: males, black circles: juveniles and females. 12 Karin VOIGTLÁNDER: Life Cycle of Lithobius mutabilis Table 2. Comparison of different characters in epimorphic stages of L. mutabilis in the investigations from Neiße Valley (NV, present investigation), Steigerwald (St, FRUND 1982) and Solling (So, ALBERT 1982). Stage Sum of coxal pores of the leg Antennal articles Number of Head-width in mm Body length pairs 12-15 ocelli in mm NV St NV St So NV St NV St So NV So VIA) min. 18 24 22 3 0.57 0.58 0.66 4.13 mode 8 8 24 25 5 5 0.69 0.70 0.75 5.63 8.20 max. 26 26 32 6 0.83 0.74 0.88 7.09 VII) Fj Qj min. 8 8 8 24 26 24 4 5 0.67 0.78 0.76 5.28 mode 12 16 11 28,29 28 6 6 0.82 0.83 0.91 6:73). 49:95 max. 16 16 16 34 30 33 8 7 0.98 0.89 1.10 8.82 VII) min. 14 16 13 26 29 30 5 6 0.59 0.89 0.86 7.00 mode 18 20 18 31 31 9 8 0.92 0.94 1.03 7.94 11.37 max. 20 24 23 36 36 39 11 10 1.27 1:03 12 9.74 IX(4) min. 18 18 19 31 35 33 5 9 0.85 0.92 1.06 8.18 mode 22 22 23 34,35 38 9 10 1.00 1,06 1.18 8.96 13.11 max. 22 26 28 39 43 39 10 12 1.17 1.16 1.34 10.00 X(5) min. 24 22 23 31 36 36 7 9 1.00 1.07 9.50 mode 28 28 27 40 41 9 12 1.18 1.20 1.34 11.04 14.88 max. 30 36 33 44 45 45 13 15 14 1.32 14.00 o Q O XI(6) min. 26 28 28 33 28 40 10 li 0.96 1.23 9.73 mode 32 34 32 37 39 43 12 14 1:31. - 185 1.47 13.01 16.36 max. 38 38 36 44 44 48 13 17 1.56 1.47 15.50 XIK7) min. 30 30 28 35 30 40 9 13 1:07 1.35 11.14 mode 36 37 34 38 3840 44 13 16 1.43 1.48 1.56 14.15 17.38 max. 42 44 39 43 45 48 18 20 1,63. 1.57 [75 XIII(8) min, 32 36 32 36 33 41 9 14 1.37 1.54 12.08 mode 38 42 36 40 39 43 13 18 1.58 1.61 1.65 14.88 18.40 max. 44 44 42 43 46 46 19 21 1.73 1.74 18.70 XIV(9) min. 34 38 37 45 33 40 12 19 1.46 1.64 13.63 mode 40 46 40 48 39 44 14 21 1:69 1,72 176 15.91 19:65 max. 46 48 47 49 47 48 20 23 1.94 1.83 22:50 to use the original terms of VERHOEFF (ANDERSSON 1976 ff.; FRÜND 1983; Daas et al. 1996; SERRA & MIQUEL 1996; Tur 2002) and to call the anamorphic “larval” stages LO —L4, and the epimorphic “post larval” stages PL 1, PL 2, Pl 3 etc. In the present study, the stages are designated (fol- lowing DOHLE 1986) by Roman numerals from first to last stage (I-XX). Additionally, the epimorphic stages are numbered in parentheses with Arabic numerals [(1— 15)]. A special problem was caused by ALBERT (1982) using the classification of stages created by VERHOEFF (1905) with- out consideration that VERHEOFF’s “agentilis” consists of two different stages (BRÖLEMANN 1930; JoLy 1966; LEWIS 1981). This mistake was corrected by ALBERT herself (AL- BERT 1983b). ! The number most frequantly is underlined.. Determination and number of stages. The anamorphic period of the post-embryonic development of L. mutabilis comprises five stages, each terminated by a moult. Easi- ly discernible morphological characters distinguish each stage. Stage I. Seven pairs of legs and one pair of unbristled limb buds (“Beinanlagen”). The buds of legs 9 can be seen as small lateral bumps at the end of the body. The number of ocelli and antennal articles ıs 2 and 7, respectively, in all individuals investigated (Figs 3, 4). Stage II. Eight pairs of legs, but legs 8 are incompletely developed and non-functional. All individuals have 11 an- tennal articles and 2 ocelli. coxal pores coxal pores eae NW UD I 00 coxal pores - NW ek mA 10 SO coxal pores = NY WwW E yn A — Fig. Bonner zoologische Beiträge 55 (2006) 13 —R3 12" leg Es A 9 3 Shey hae BAe | : UN TA. E v VI Vil VII IX x XI XII XIE XIV XV XVI XVII XVIIE XIX XX (1) Q) (3) (4) (5) (6) (7) (8) (9) (10) any Ad) (3) (AM) (HS) stage 13" leg N 8 9 Ñ 7 NS L, E le] aN gue q gue Q DK nen VI Vil Vul IX x XI XII XIE XIV XV XVI XVII XVII XIX XX (1) (2) (3) (4) (5) (6) (7) (8) (9) (10) an (13. (3) (AM) (HS) stage \> = Y QA) E 0°. 00 y ssp V VI Vil Vill IX Xx XI XII XIII XIV XV XVI XVI XVII XIX XX (1) (2) (3) (4) (5) (6) (7) (8) (9) CO an (2) (13) (AN) (5 stage 100% mE 7 hos aoe VI Vil VHI IX X XI XII XI XIV XV XVI XVI XVII XIX XX (1) (2) (3) (4) (5) (6) (7) (8) (9) (10) (1) (12) (13) (dA) (1) stage 2. Variation in numbers of coxal pores in different stages of L. mutabilis. 14 Karin VOIGTLÄNDER: Life Cycle of Lithobius mutabilis 100% Bo number of ocelli I W ill IV Y vi vu Vill IX x (1) (2) (3) (4) (3) stage n=12 n=14 n=12 m 2 n=6 20 n=2 - 18 — 16 14 = S nz 10 8 6 2 4 El 2 NI NIL NIU NIN XV XVI XVIL XVII XIX XX (6) (7) (8) (9) (10) (il) (12) (13) (14) (15) Fig. 3. Variation in numbers of ocelli in different stages of L. mutabilis, n: number of ocular fields counted. Stage III. Eight fully developed pairs of legs and two pairs of limb buds; 14 antennal articles and 2 ocelli are char- acteristic for this stage. Very rarely 3 ocelli occur. Stage IV. Ten pairs of legs and two pairs of limb buds. The number of antennal articles varies between 17, 18 and 191, the number of ocelli between 3 and 4. Stage V (last anamorphic stage). Twelve pairs of legs and three pairs of limb buds, mostly with 23 antennal articles (16-23). The number of ocelli varies between 3, 4 and 5. The first epimorphic stage VI (1) and all following stages have 15 functional pairs of legs. Therefore, other morpho- logical characters (parameters of size as well as numbers of coxal pores, ocelli and antennal articles) must be used for determination of stages. The variability of these char- acters at each stage was determined for a better charac- terization of each stage. The direct observation of all hatchings and the investiga- tion of the development of different morphological char- acters during the life cycle of L. mutabilis in captivity led to the discovery of 15 epimorphic stages. Growth analyses. The growth analyses show more or less sigmoid curves (Fig. 1) for all parameters, especially for body length, head length and width, and width of tergite 3. Up to stage V (the last anamorphic stage), there is a rel- atively small increase between each moult. With the next stage VI (1), relatively constant greater increments appear (linear curves) up to approximately stage XIV (9), where the curves become flatter again. The differences between various stages are highly signif- icant for all measurements for males and females, where- as the differences between males and females of the same stage are very rarely significant (Mann-Whitney-U test). Number of coxal pores. The last anamorphic stage V of L. mutabilis has one coxal pore on its last leg (leg 12; Fig. 2). The following epimorphic stages have pores at the cox- ae Of legs 12-15. Each moult can lead to an increase of 1 or 0 coxal pores per leg. On leg 12 there is never an in- crease of pores during the moult from stage V to stage VI (1). From 273 observed epimorphic moults only 6 (2%) occurred without any addition of pores at one of the four pairs of coxae. Figure 2 shows the combination of coxal pores in different stages and sexes of L. mutabilis. Females have more coxal pores than males (Table 2). Fig- ure 2 shows that the differences between the sexes are due Bonner zoologische Beiträge 55 (2006) 15 100% 50 1 45 4 n=59 40 4 n=24 PR: 35 - n=67 A uk wi 30 4 i A J n=56 n=64 number of antennal articles n=17 n=16 27 1 n=15 n=13 n=11 - - . m - # ; + 40) r 30 number of antennal articles r 10 T T el ae T PT q A _ T 1 T 1 Il ii IV Y vi Vil Vill IN x NI Nt NIN xy AVI XVI. NVI NIX XX (1) (2) (3) (4) (5) (7) (8) (9) (10) am) (12) (13) (14) (15) stage Fig. 4. Variation in antennal articles on each side in different stages of L. mutabilis, n: number of antennae counted. to the lower number of coxal pores especially of leg 15 of males. In each stage there are certain preferred arrange- ments of increase. Overlaps of the preferred variants in males were between VIII (3) and IX (4), ın females al- ready between VII (2) and VIII (3). However, such over- laps apply to two stages only. In 16 % the left and right sides were different, but at most in one pore and on one leg only. Number of ocelli. The first three stages have two ocelli on each side ofthe head. From stage IV onwards, the num- ber of ocelli increases with each moult (Fig. 3). Variations begin with the (anamorphic) stage IV and become greater in later epimorphic stages. Within one stage, they can amount to as much as 12 ocelli. Thus, the overlap between different stages is very high. In both sexes, stage XVIII (13) with 24 ocelli has the highest observed number. The single male in the highest stage [XX (15)] had only 19 ocelli. No significant differences exist between males and females. Differences between the two sides of the head were rarely observed. Number of antennal articles. The number of antennal ar- ticles increases during post-embryonic development from 7to a maximum of 49 (Fig. 4). In the first three stages there is no variation in number. All studied antennae show 7, 11 or 14 articles, respectively. In stage IV, most indi- viduals have antennae with 19 articles (with little varia- tion). Beyond stage V, the number of antennal articles varıes very widely within one stage. Also the left-right variability is very high (more than 50 %). In field-sam- pled individuals, it is often impossible to know whether a lower number results from an incomplete regeneration after loss of some articles. In the first epimorphic stage VI (1), the usual numbers are 20 - 24 - 25. The increase continues up to stage X (5), af- ter which it ceases (Fig. 4, Tab. 2). There is no significant difference in the number of antennal articles between males and females in any stage. 3.2. Reproduction biology and life cycle Mating behaviour, oviposition and egg development. Mating behaviour and spinning of webs with sper- matophores by males, as described by KLINGEL (1960) and DEMANGE (1956) for the species L. forficatus and L. piceus gracilitarsus Brölemann, 1898, were not observed for L. mutabilis in this investigation. Of 57 adult females, 26 (both from the field and from lab- oratory cultures, investigated between 1993 and 1997) laid 16 Karin VOIGTLANDER: Life Cycle of Lithobius mutabilis n=42 500 450 up to 788 stages duration in d n=33 n=3] =36 n=32 N tee I Il Hl IV Vv VI (1) 2) X (4) X (5) X1 (6) VIL ( XII (7) XI (8) XIV (9) XV (10) XVI (11) XVII (12) XIX (14) VIII (3) XVII (13) Fig. 5. Average stage duration, minimum and maximum of L. mutabilis [in days]. —t— 1997 —=— 1998 Fig. 6. Moulting activity of L. mutabilis ın laboratory under constant conditions during the years 1993 to 1998 in % of the total number each month. Bonner zoologische Beiträge 55 (2006) 17 % 100 60 40 20 IX ( X (5) XI (6) Fig. 7. eggs in captivity, some of them in several successive stages and in several years. Altogether 35 egg laying “periods” were observed by females of stages XI (6) to XVII (12). Egg numbers vary between | and 35, on average 11 eggs per female per egg laying period. The eggs are laid singly. There is no maternal care of the brood in L. mutabilis as in other lithobiid species. The on- ly protection for the eggs is a thin cover of fine soil par- ticles, which is applied to the eggs by the female. In the rearing boxes without soil, this behaviour is retained and fine gypsum dust is used by the females. The egg laying itself and the following processes (BROCHER 1930 and SERRA & MIQUEL 1996 described it in detail for L. forfi- catus and L. pilicornis) could not be observed directly in this study. Only the covered eggs and females carrying eggs with their gonopods could be confirmed. The eggs of L. mutabilis, as in other lithobiid species, are spherical and whitish. The mean diameter is 0.89 mm. The appearance of the youngest stages in nature (Neiße Val- ley) suggested that oviposition is not connected with a def- inite time in the year. This was also established by labo- ratory cultures of L. mutabilis in the present study. From 1/3 of laid eggs, young L. mutabilis were hatched. Though all eggs were kept under the same constant conditions (16 °C), they needed quite different times for their develop- ment. The time between oviposition and hatching ranged between 21 and 284 days (9.5 months). Most individuals hatched after 3-4 months. Onset of sexual dimorphism and maturity. The sexes of L. mutabilis can be distinguished for the first time (with Mortality rate of epimorphic stages of L. mutabilis in % of the total number in these sta 3) So > = = a e > a = > = = = au ~ = = — > = = E A x z > x = = = ¡de = PS x = > 5 A > es. go 7% some uncertainty) in stage VII (2), definitely in stage VIII (3). In the laboratory, first egg layings were observed in stage XI (6). Thus XI (6) can be seen as the first ma- ture stage for females. Duration of stages and moulting activity. The data for the duration of the different stages in the postembryonic development of L. mutabilis come from 150 specimens, both laboratory-reared as well as freshly captured. Some 490 moults were observed, 2—11 per individual. Observa- tion of moults every two days was sufficient to ensure ac- curacy. If moults were not observed directly, at least the * remains of exuviae were found. If they were completely eaten by moulted individuals, they could be seen in the faeces. The mean stage duration increases with stage number (Fig. 5), but with higher variability in older epimorphic stages caused by some “slow” or “fast” individuals. It varies from only a few days to about 500 days. An extreme case was found in stage XII (7). Here, the shortest and longest du- ration was 10 and 788 days, respectively. The mean du- ration of stages XVII (12) to XIV (14) ıs based on only 1-4 animals and is therefore not representative. However, it was not usual that a specimen had exclusive- ly long (or short) durations of all the stages throughout its life. Therefore, it is possible to make general conclu- sions about the most important data based upon the aver- age of stage duration (Figs 5, 8). In L. mutabilis the first epimorphic stage 1s reached after 4.5 months and sexual maturity is attained at the earliest after 1.5 years, defini- tively after 2 years. The maximal life expectancy is about 18 Karin VOIGTLÄNDER: Life Cycle of Lithobius mutabilis spigomuu +7 sıpgopmun sadisspi9 | ponuyuos 11 panuyuos Ost fs II] panuyuos Si 006 zip if ap el A E IX (L) TIX + CO IX E 056 lt (6) AIX (6) AIX + (8) IX (0) IX 0001 AA AAA — (6) AIX + (8) MIX 1 OSO! u SE 1 0011 059 (A) IX — (EI) MAX A li eh E (ee (OL) AX — (2) IX (L) 1X OST 11 4d: (L) TIX + (0) 1X (8) TIX + (2) LIX 0071 LI 1 Ss Ses | (2) TIX +(9) IX ; "| ı 5 | 7 : N 5 9 oe | | ® | a] 3 Fig. 8. Duration of developmental stages of different Lithobiomorpha (according to ANDERSSON 1990) and £. mutabilis (this in- vestigation). For L. pilicornis, SERRA & MIQUEL (1996) give only data until stage V. Bonner zoologische Beiträge 55 (2006) 19 5 years. On average, L. mutabilis lives 3-3.5 years. The moulting activity of the epimorphic stages of L. mu- tabilis during the years 1993-1998 can be seen in Figure 6. On average, 27 individuals per month were available. Under constant culture conditions, moults in Z. mutabilis take place throughout the entire year without any peaks at a specific time. Mortality. The mortality rate of 213 individuals of the epi- morphic stages VI (1) to XX (15) was established. Fig- ure 7 shows the number of dead individuals as a percent- age of the total number in these stages. The values are not fully correct, because freshly collected and “laboratory” individuals were analysed together. It is possible that fresh individuals were more viable. Nevertheless, the mortali- ty of the youngest epimorphic stages until XI (4) was very low. The transition from stage X (5) to XI (6), the first mature stage, was only reached by 50% of the individu- als. In stage XI (6) the mortality was very low, but then it increased very rapidly. Most of the specimens died in stages XII (7) to XV (10). Only 11 individuals reached stage XVI (11) and only one the highest stage XX (15). 3.3. Observation and discussion on feeding behaviour Some observations on the feeding behaviour were under- taken in the laboratory. They are compared with results of PFLEIDERER-GRUBER (1986) and ALBERT (1983a). These authors studied feeding behaviour more intensive- ly in various centipede species, including Z. mutabilis, ın regard to prey capture, prey spectrum and preference, in- fluence of different food resources on growth, and others. Animal food. In the laboratory cultures, I offered differ- ent animal foods separately to L. mutabilis and other species at different time periods. Small pieces of earth- worms and living 7ubifex were given, as well as aphid lar- vae, small Diptera larvae and also Collembola. The choice of prey depended on the size relationship between pred- ator and prey. Small collembolans (Folsomia spec.) were only attacked by juveniles of L. mutabilis and smaller species of the subgenera Monotarsobius and Sigibius [L. austriacus (Verhoeff, 1937), L. microps]. They noticed the prey only when their antennae came into direct contact with it. The observed feeding process is in correspondence with the description by RILLING (1968). Minced meat was refused, contrary to results from PFLEI- DERER-GRUBER (1986), who found it to be eagerly taken after 14 days of food withdrawal. Particularly bad results were experienced with ox heart. It led to a mass occur- rence of mites (Heterocerconids) in the rearing vessels. Their nymphs attacked the Lithobius species so massive- ly, that they could not feed or were unable to move. Cannibalism. Lithobius mutabilis was never observed to feed on members of the same species during the entire in- vestigation period of more than 5 years, provided that the individuals had approximately the same size and were not injured. Once, a £. forficatus was observed attacking a de- fenceless specimen of the same species during moulting. The specimen bit at first in the “neck”, fed then on soft- er interior substances before eating pieces of the integu- ment. Plant food. In my investigations, I always offered green leaf pieces of indoor plants in addition to other food. In most cases, these plant pieces were clearly nibbled off. According to PFLEIDERER-GRUBER (1986), all Chilopoda are pure carnivores and never eat plant material, detritus or soil, in contrast to other literature (e.g. DOBRORUKA 1961). PFLEIDERER-GRUBER assumed that such particles, which were only found as gut contents, were accidental- ly picked up with other food, or 1t could be still undigest- ed gut contents of herbivorous prey. ALBERT (1983a) al- so writes that L. mutabilis does not seem to be able to feed on litter, even in times of food shortage. My investigation refutes these doubts, at least in the case of L. mutabilis, which is able to bite off plant parts as food in small amounts. Therefore, ıt can be assumed that this species also can take in vegetable food under natural con- ditions. 4. DISCUSSION 4.1. Character analyses Growth analysis. The growth of body parameters shows sigmoid curves corresponding to the two types of devel- opment: after a slow beginning during the first five anamorphic stages, linear growth rises more rapidly after the onset of epimorphic development. Then, stages XIV (9) to XX (15) seem to grow more slowly than the younger stages. However, this may be a result of the low number of individuals studied. An approximately linear growth of the body parameters of epimorphic stages is also found for many other Lithobius-species (ALBERT 1982; ANDER- SSON 1976-1984; Kos 1997; VOIGTLANDER 2000). It seems to be typical for all Lithobiomorpha. In millipedes, maturity often causes a cessation of growth because all reserves are used for the development of eggs and sperm (e.g., VOIGTLANDER 1987, 1996). This could not be established for Z. mutabilis in the present in- 20 Karin VOIGTLANDER: Life Cycle of Lithobius mutabilis Table 3. Lithobiomorpha with investigated life cycles (or at least some aspects of it). * = examples for detailed descriptions of immature stages Taxon Arenobius manegitus (Chamberlin, 1911) Bothropolys asperatus (L. Koch, 1878) Esastigmatobius longitarsis Verhoeff, 1934 Ethopolys xanti (Wood, 1863) Eupolybothrus nudicornis (Gervais, 1837) [Syn. E. elongatus (Newport, 1849)] Eupolybothrus grossipes (C.L. Koch, 1847) Lamyctinus coeculus (Brölemann, 1889) Lamyctes emarginatus (Newport, 1844) [Syn. L. fulvicornis Meinert, 1868)] Lithobius atkinsoni Bollman, 1887 Lithobius austriacus (Verhoeff, 1937) Lithobius borealis Meinert, 1868 Lithobius b. burzelandicus Verhoeft, 1931 Lithobius calcaratus C. L. Koch, 1844 Lithobius canaliculatus Murakami, 1963 Lithobius crassipes L. Koch, 1862 Lithobius curtipes C. L. Koch, 1847 Lithobius dentatus C. L. Koch, 1844 Lithobius ervthrocephalus C. L. Koch, 1847 Lithobius forficatus (Linné, 1758) Lithobius hirsutipes Eason, 1989 Lithobius lapidicola Memert, 1872 Lithobius macilentus L. Koch, 1862 Lithobius melanops Newport, 1845 Lithobius microps Meinert, 1868 Lithobius mutabilis L. Koch, 1882 Lithobius nihamensis Murakamı, 1960 Lithobius obscurus azoreae Eason & Ashmole, 1992 Lithobius pachypedatus Takakuwa, 1938 Lithobius peregrinus Latzel, 1880 Lithobius pilicornis Newport, 1844 Lithobus schawalleri Eason 1989 Lithobius tenebrosus fennoscandicus Lohmander, 1948 Lithobius punctulatus C. L. Koch, 1847 [Syn. £. validus Meinert, 1872] Lithobius variegatus Leach, 1814 Nothembius insulae Chamberlin, 1916 Oabius pylorus Chamberlin, 1916 Paobius boreus Chamberlin, 1916 Tidabius tivius (Chamberlin, 1909) Author(s) CHAMBERLIN (1917)* MURAKAMI (1958) MURAKAMI (1960d) CHAMBERLIN (1925b)* Daas et al. (1996) EASON (1970)* ANDERSSON (1979, 1990) ANDERSSON (1979, 1984b, 1990), ZULKA (1991) CHAMBERLIN (1925a)* VOIGTLÄNDER 2000 ANDERSSON (1979, 1980a, 1990) MATIC & STENTZER (1978) ANDERSSON (1979, 1982a, 1990) MURAKAMI (1963) ANDERSSON (1979, 1981, 1990), WIGNARAJAH (1968) ALBERT (1982, 1983b), ANDERSSON (1979, 1983, 1990) VOIGTLÄNDER 2000 ANDERSSON (1978b, 1979, 1990), VOIGTLÄNDER (2000) CHAMBERLIN (1925a)*, VERHOEFF (1925), BROCHER (1930), LEwIs (1965), WIGNARAJAH (1968), ANDERSSON (1976, 1979, 1990) EASON (1989) * ANDERSSON (1980a) ANDERSSON (1990) ANDERSSON (1979, 1980b, 1990) ANDERSSON (1979, 1982b, 1990), VOIGTLÄNDER (2000) ALBERT (1982, 1983b), FRÜND (1983) MURAKAMI (1961) EASON & ASHMOLE (1992) * MURAKAMI (1960a, b, c) Barber & Eason (1986) * SERRA & MIQUEL (1996) EASON (1989) * ANDERSSON (1984a, 1990) Kos (1997) ROBERTS (1957), EASON (1964)*, LEWIS (1965) CHAMBERLIN (1916)* CHAMBERLIN (1916)* CHAMBERLIN (1916)* CHAMBERLIN (1913)* Bonner zoologische Beiträge 55 (2006) 21 vestigation. In spite of the early maturity, at stage XI (6), this species still grows continuously in all measured pa- rameters. Corresponding results were published by FRÜND (1983) (head width) and ALBERT (1982) (body length, head width) for L. mutabilis. Coxal pores. In all Lithobiomorpha, the first coxal porus is developed on the leg 12 in stage V already, whereas the coxal pores of legs 13-15 appear in stage VI (1) at first. The present investigation corroborates this for L. muta- bilis (Fig. 2). In contrast to other species (e.g., L. erythro- cephalus, L. melanops, L. crassipes, L. curtipes), where two pores may be observed on leg 12 of stage VI (1) (AN- DERSSON 1978b, 1980b, 1981, 1983), for £. mutabilis on- ly one coxal pore was found in all individuals of stage VI (1). As a general rule for Lithobiomorpha shown in many in- vestigations (Tab. 3), no more than one coxal pore per leg is added during a moult. For this reason, the number of pores of legs 13-15 does not exceed the number of the epimorphic stage; rarely it is the same at least. Only on leg 12, one more coxal pore is possible, because one pore is already developed in stage V. Therefore, the number of coxal pores is a very good character for separating the dif- ferent stages of L. mutabilis, just as in other Lithobius- species. For some species, e.g., L. calcaratus, L. crassipes, L. cur- tipes (ANDERSSON 1981, 1982a, 1983), L. piceus and L. tricuspis (TOBIAS 1969), it was established that coxal pores are more numerous in females than in males. This 1s al- so valid for L. mutabilis. Neiße Valley- and Steigerwald-populations show approx- imately the same numbers of coxal pores summarized from the last four legs in minimum, maximum and “mean” (Tab. 2). The “mean” values differ only by 1-2 (3) pores. The differences between minimum values and maximum values, respectively, are a little higher (up to 7 pores). In both studies, females had significantly more coxal pores than males. If a comparison between the coxal pores of each leg is made, a very close similarity between the populations can also be seen (Fig. 3 herein and fig. 2 in FRUND 1983). Not only are the numbers of pores on each leg the same in comparable stages, but also the distribution of the num- bers. In both studies, legs 13 and 14 reach the most (7) coxal pores. In the present investigation, the number of coxal pores increases to 8 or 9 in later stages, not found by FRUND (1983). If ALBERT’s (1982) PL1 consists of two different stages, then the number of coxal pores is similar to FRÜND’s and the studies presented here. Ocelli. In the investigations at Steigerwald (FRÚND 1983), variations in the number of ocelli begin in stage VI (2) (Tab. 2), whereas in the present study they start dur- ing anamorphic development in stage IV (Fig. 3). How- ever, in stage VI (1) the usual number was 5, as in the Steigerwald-population. The increase of the number of ocelli was more regular in the investigations by FRÜND (1983) than in the presented study, where some succes- sive stages show the same number of ocelli on average. The increase of ocelli during post-embryonic development of L. mutabilis shows more or less the same pattern as in other Lithobius-species. With the exception of £. forfica- tus, L. variegatus and L. pilicornis, all species investigat- ed up to now have two ocelli in the first three anamor- phic stages. Then a more or less linear increase can be ob- served. However, in higher stages, a greater variability in increase of ocelli number occurs. Differences between the sexes do not exist in all Zithobius species, and irregular- ities between the left and right side are rare. Antennal articles. In the present investigation, a gradual increase in the mean numbers of antennal articles is on- ly seen until stage X (5). However, in the population from Steigerwald (FRUND 1983) this cessation in increase oc- curs later [XI (6)]. The numbers reached in the different stages are mostly smaller in the present investigation and some differ very much from those observed by FRUND (1983) (Table 2). The variations within stages are larger, as 1s the overlap between different stages. Altogether, the growth of antennae in the population from Steigerwald is more regular than in that of the Neiße Valley-population. The numbers of antennal articles of specimens from Solling (ALBERT 1982) show also high variation, especial- ly in the younger stages (Tab. 2). In comparison with other Lithobiomorpha (ANDERSSON 1976-1984), there are no differences for the first three anamorphic stages. In later stages of Lithobius, the in- crease takes place in two different ways. In some species (L. melanops, L. calcaratus, L. tenebrosus fennoscandius), the increase continues until the last stage almost in linear fashion. In others (L. microps, L. ervthrocephalus, L. for- ficatus), the growth of antennae is almost completed at a specific stage [IX (4), X (5), XI (6) or XII (7)]. To this group belongs L. mutabilis [stagnation from X (5)]. Characteristically for all Lithobius-species (including L. mutabilis) 1s the absence of differences in the number of antennal articles between males and females. The exten- sive variation in the number of antennal articles within a stage in all species is also typical, but may partly result from undetected regeneration. Therefore, the use of this character for stage determination 1s restricted. hn m 4.2. Number of stages Like all Lithobiomorpha, L. mutabilis develops by hemi- anamorphosis with five anamorphic and a various num- ber of epimorphic stages. VERHOFFF (1905) described on- ly five epimorphic stages for L. mutabilis and called them “status agentilis” (realized by BRÓLEMANN [1930] and JOLy [1966] as “agentilis I” and “agentilis IT”), “status im- maturus”, “status praematurus”, “status pseudomatururs” and “status maturus”. ALBERT (1982) and FRUND (1983) distinguish nine epimorphic stages. How can differences in the number of observed stages come about? ALBERT (1982) and FRÜND (1983) almost ex- clusively investigated animals that were collected in the field. Only a few individuals were reared for testing in the laboratory. It is impossible to determine the exact epimor- phic stage of field animals, because the range of variation of all characters used for stage determination is very large and overlaps between stages. Only direct observation of moulting in the laboratory makes it possible to determine the correct number of stages. Additionally, most individ- uals die in stages XIII (7) to XV (10), even under favourable conditions in the laboratory (Fig. 6). Thus, 1t is conceivable that, in the field, death happens even ear- lier. FRUND (1983) sampled most individuals at stages X (5) to XII (7). In the Neiße Valley more than half of all sampled individuals were found to be stages XI (6) and XII (7). Later stages were sampled very rarely (20%) in the Neiße Valley. 4.3. Life cycle Oviposition. Oviposition in L. mutabilis is not connect- ed with a certain time of year. This was established by lab- oratory cultures in the present study as well as by inves- tigations of ALBERT (1979, 1983b). According to LEWIS (1965) and WIGNARAJAH & PHILLIPSON (1977), lithobiids are able to lay eggs during the whole year (L. variegatus), but some species (e.g., L. forficatus) have a main egg-lay- ing period in spring and a smaller one in autumn. This dif- ferent behaviour of the species and the small number of eggs laid during each oviposition period might be inter- preted in correlation to habitat choice. A long egg-layıng period guarantees a breeding success, even if some of the eggs are destroyed through unfavourable conditions, such as dryness. The behaviour of £. variegatus (LEWIS 1965) and L. mutabilis may indicate that they are less resistant to climatic extremes than L. forficatus. Both species live in woodland, whereas L. forficatus is more euryoecious and often prefers dry habitats. A female £. mutabilis can lay eggs in several periods (each of these periods can reach up to two months), each sepa- Karin VOIGTLÄNDER: Life Cycle of Lithobius mutabilis rated by one moult at least. In L. forficatus — as well as in some diplopods — the development of the eggs and moults is subject to mutual hormonal mechanisms of reg- ulation (RICHTER 1967; SCHEFFEL 1969; LEUBERT & SCHEF- FEL 1984; VOIGTLÄNDER 1987). The same mechanisms seem to control the life cycle of L. mutabilis. Egg development. The present investigation shows a large variation in the duration of egg development. This agrees with other studies on L. mutabilis. ALBERT (1983b) ascer- tained individually variable developmental times between 27-269 days (15 °C) and Tur (2002) found time periods of approximately 150-200 days under natural conditions. This different “hatching period” in combination with a long egg-laying period characterises the life strategy of Lithobius in general, not only of L. mutabilis. It guaran- tees the survival of at least some individuals. Onset of maturity and insemination time(s). In agree- ment with the present results, ALBERT (1983b) also defined stage XI (6) as first mature stage of L. mutabilis. FRUND (1983) deduced from investigations of the structure of the secondary sex characters that X (5) is not yet adult, but stages XII (7) to XIV (9) definitely are. To determine the first stage of maturity in males, it is nec- essary to examine the testes, i.e. whether they contain sperm. It 1s not possible to prove this in breeding cultures. Each female was kept with one or two males, but it is not certain whether their eggs were fertilized by the males present, because females are able to conserve sperm in their seminal receptacles for up to two years (personal ob- servations on a lone female). The same phenomenon was found in Scutigeromorpha (MURAKAMI 1956, DOHLE 1969). ToBIAS (1969) observed egg laying in Lithobius seven months after isolation of the females and VERHO- EFF (1915) after approximately one year. Personal obser- vations support the period of one year for L. forficatus. Individual development. The high individual variabili- ty of stage duration (Fig. 5) means that new developmen- tal periods (epimorphic development, maturity) begin at very different times in each individual. Moulting activi- ty and stage duration depend to a large degree on the en- vironmental conditions. Mostly food shortage and low temperatures inhibit development (ROBERTS 1957; JOLY 1977; ANDERSSON 1978a). The loss of extremities can stimulate moulting (VERHOEFF 1915; SCHEFFEL 1980). Under constant temperature conditions and optimal feed- ing in the present study, it was expected that all individ- uals would have a relatively similar stage duration. But, on the contrary, they varied very widely especially in high- er epimorphic stages. This is not only a phenomenon of Bonner zoologische Beiträge 55 (2006) 23 L. mutabilis, but occurs in other species too (ANDERSSON 1990). However, the individual differences in stage duration in the studies of ANDERSSON, ALBERT, and myself cannot be caused by hunger, since all individuals were well fed in all investigations. Further, the different constant culture temperatures (ANDERSSON: 20 °C, ALBERT: 15 °C, pres- ent study: 16 °C) could have had an influence. But An- DERSSON investigated all species under the same conditions and also found large differences between the species ad- ditionally to the individual differences. Thus, it seems that environmental conditions have only a secondary influence on intra- and interspecific variability of life span. To explain the interspecific differences ANDERSSON (1990) suggested the possibility of a “built-in clock” by inspection of moults in different months. The smaller ($i- gibius Monotarsobius) species, L. microps and L. cras- sipes, show many fewer moults during the winter. The larger (Lithobius s.str.) species, L. forficatus and L. ery- throcephalus, show no regular differences. Up to now, there is no explanation for the difference in moulting pat- tern between large and small species. The investigations have shown that L. mutabilis moults throughout the en- tire year under constant temperature conditions. It can thus be assumed that L. mutabilis does not have a “built-in clock”. This corresponds with the other larger Lithobius- species (ANDERSSON 1990). Population development. The most important life histo- ry data for L. mutabilis from Neiße Valley are in concor- dance with those found for the Solling-populations by AL- BERT (1983b). The author gave 0.5-1 year till attainment of stage VI (1) (4.5 months ın the present paper). Matu- rity is reached earliest in stage X (5) after 1.5—2 years ac- cording to both investigations. The total life span in L. mu- tabilis is suggested by ALBERT’s abundances data to be be- tween 4 and 7 years, which 1s a little longer than in the present study by laboratory observations (3-5 years). Figure 8 shows the developmental time of different species (according to ANDERSSON 1990; SERRA & MIQUEL 1996) in comparison to L. mutabilis (present paper). The short- est stage of L. mutabilis was stage I with only 5 to 12 days, which is much longer than in all other investigated species. The first epimorphic stage was reached after 4 months, ap- proximately the same time as in L. forficatus, but longer than in other species. L. mutabilis has reached stage XI (6) after 510 days only, whereas other species reach this stage as early as after 255-390 days (Fig. 8). This means a very retarded development of L. mutabilis. This species has the highest number of stages as well as the greatest age of all species investigated in the laboratory. Howev- er, it seems that L. mutabilis dies earlier in the field than in captivity. 4.4. Conclusions The results of the character analyses (growth, biomass, number of coxal pores, antennal articles and ocelli) in the geographically widely separated populations of L. muta- bilis Steigerwald, Solling (western Germany) and Neiße Valley (eastern Germany) show very high correspondence. The number of stages recognised in this laboratory study was much higher than in the other investigations, perhaps due to methodological problems of stage analyses on old- er field-collected individuals. However, the life cycles of the different populations are very similar. This suggests that in L. mutabilis the study of an isolated case can be generalised and this may also be true for other species. Acknowledgements. | thank Prof. Dr. W. Dunger/Görlitz, Prof. M. Zapparoli/Viterbo, and Dr. J. G. E. Lewis/Taunton for criti- cal reviews and comments on the manuscript and especially the latter for his help correcting the English text. Prof. Dr. W. Xy- lander provided institutional support. 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Leipzig: 1-725. [1915: 83.-85. Liefer- ung: 313-394; 1925: 100. Lieferung: 539-666]. VOIGTLÄNDER, K. 1987. Untersuchungen zur Bionomie von En- antiulus nanus (Latzel, 1884) und Allajulus occultus C. L. Koch, 1847 (Diplopoda, Julidae). Abhandlungen und Berich- te des Naturkundemuseums Görlitz 60 (10): 1-116. VOIGTLÄNDER, K. 1996. The life cycle of Cylindroiulus lates- triatus (Curtis, 1845). Pp. 501-508 in: GEOFFROY, J.-J., MAU- RIES, J.-J. & NGUYEN Duy-JAQUEMIN, M. (eds.) Acta Myri- apodologica. Mémoires du Musce national d’Histoire naturelle 169. VOIGTLANDER, K. 2000. Vergleichende Untersuchungen zur Postembryonalentwicklung von Lithobius-Arten (Chilopoda, Lithobiidae). Mitteilungen der deutschen Gesellschaft ftir all- gemeine und angewandte Entomologie 12: 535-540. WIGNARAJAH, S. 1968. Energy dynamics of centipedes popula- tions (Lithobiomorpha — £. crassipes and L. forficatus in woodland ecosystems). Ph. D. Thesis, University of Durham.* WIGNARAJAH, S. & PHILLIPSON, J. 1977. Numbers and Biomass of Centipedes (Lithobiomorpha: Chilopoda) in a Betula-Al- sus Woodland in N.E. England. Oecologia 31: 55—66. ZuLKA, K. P. 1991. Uberflutung als ökologischer Faktor: Verteilung, Phanologie und Anpassungen der Diplopoda, Lithobiomorpha und Isopoda in den Flußauen der March. The- sis, Universitat Wien. Author’s address: Dr. Karin VOIGTLANDER: State Muse- um of Natural History Górlitz, PF 30 01 54, 02806 Gór- litz, Germany; E-Mail: Karin. Voigtlaender@smng.smwk.sachsen.de Received: 02.2005 Revision: 28.04.2005 Accepted: 29.04.2005 Corresponding editor: B. A. Huber JS , . > 7 = ] ' . \ : : ] o : & ae. vee 7 : u en 7 Ñ . A 7 | 7 5 y e ne u u nn 5 A o o o u u u o o o o . Bonner zoologische Beiträge | Band 55 (2006) Heft | Seiten 27-35 Bonn, Januar 2007 Amphibians and Reptiles of the Tchabal Mbabo Mountains, Adamaoua Plateau, Cameroon Hans-Werner HERRMANN!), Andreas SCHMITZ), Patricia A. HERRMANN?) & Wolfgang BOHME?) ‘Center for Reproduction of Endangered Species (C.R.E.S.), San Diego 2)Museum of Natural History, Geneva, Switzerland 3)Messa, Yaounde, Cameroon WZoologisches Forschungsmuseum Alexander Koenig, Bonn, Germany Abstract. Tchabal Mbabo is a remote mountain range in Central Cameroon which covers 79,000 ha; only three amphi- bian and no reptile species are listed for this area. We conducted two expeditions, in 1998 and 2000, to survey the her- petofauna. We used opportunistic visual encounter surveys and driftfence/pitfall trap arrays for sampling. Our findings increased the number of amphibian and reptile species known for the area to 15 each. We discovered one recently des- cribed new frog species (Cardioglossa alsco).and found an additional six species of anurans and one lizard species which could not be assigned to any described species. The deceptive endemism of amphibians for this area is remarkable. We report a range extension of 400 km and montane gallery forest as a novel habitat for the skink Lacertaspis chriswildi. The importance of the montane forests on the northern slopes as well as the gallery forests on the southern slopes of the mountain range are highlighted. Keywords. Amphibia, Reptilia, Tchabal Mbabo Mtns, Adamaoua Plateau, Cameroon, faunistics, conservation. 1. INTRODUCTION Although herpetological work in Cameroon has been re- current during the last half century (PERRET 1959, 1960, 1961, 1966; AmieT 1971; BOHME 1975; AmIET 1978; JOGER 1982; BOHME & SCHNEIDER 1987; EUSKIRCHEN et al. 1999, 2000; LeBreton 1999; SCHMITZ et al. 1999, 2000; HERRMANN et al. 2000, 2005a, 2005b) large areas remain unsampled. Such an area 1s the Tchabal Mbabo Mtn range on the western border of the Adamoua Plateau (Fig. 1). The Tchabal Mbabo Mtns represent the last segment in the chain of mountains known as “Dorsale Camerounaise”, which extends northeasterly from the island of Bioko con- tinuing inland with Mt. Cameroon, Mt. Kupe, the Manen- gouba Mtns, Mt. Lefo and Mt. Oku. Tchabal Mbabo is a broad horseshoe-shaped mountain massif with a west-eastern orientation and altitudes of up to 2,400 m. Below 1,600 m the massif is connected to oth- er mountains. The area above 1,600 m covers 79,000 ha with 4,900 ha of relict montane forest along the steep northern slopes (Fig. 2a and 2b) adjacent to the rim which represents the northernmost occurrence of this forest type in Cameroon (THOMAS & THOMAS 1996). The southern slopes are extensive grasslands with interspersed gallery Location of the Tchabal Mbabo Mtns within Cameroon. Fig. 1. 28 Hans-Werner HERRMANN et al.: Amphibians and Reptiles of Tchabal Mbabo Fig. 2. Different Tchabal Mbabo habitats: (a) northern slope near Fungoi with patches of cloud forest (ca. 2,000m elevation), (b) cloud forest along the rim (ca. 2,000 m elevation), (c) savanna and gallery forest on the southern slopes between Fungoi and Pin- ku peak (ca. 2,100 m elevation), (d) dry forest along Mayo Selbe (ca. 800 m elevation). forests along creeks (Fig. 2c). The grasslands are managed by Fulanı cattle herdsmen and burned every year. Yearly precipitation at the southern base of the mountains is ap- proximately 1,700 mm (THOMAS & THOMAS 1996). The dry season extends from November to February, precip- itation measures < 50 mm/month; the peak rainy season extends from June to October with precipitation up to 300 mm/month. The hottest months are April and May with average daily temperatures of up to 30 °C. Annual tem- peratures on the plateau average 18 °C with a daily am- plitude of 13-15 °C (THOMAS & THOMAS 1996). We mounted data loggers on tree trunks (two meters height) in forests in the vicinity of Fungoi, at an eleva- tion of circa 2,100 m, which recorded air temperature and relative humidity at one hour intervals. The climatic da- ta for a gallery forest on the south flank is presented in Figure 3. During a data collecting period of 906 days com- prising February 2000 to July 2002, the temperature ranged from 8.6 to 26.7 °C (x = 15.4 °C). The relative hu- midity ranged from 13 to 100 % (x = 81 %). Data was al- so collected on the north flank of the mountain at an ele- vation of circa 2,100 m from February 2000 to May 2001 over a period of 465 days. Here the temperature ranged from 10.2 to 34.9 °C (x = 16.2 °C) and the relative hu- midity ranged from <5 to 100 % (x = 65 %). These data indicate the northern slope to be considerably warmer and drier. The presence of a well developed montane forest ecosys- tem of the rare northern type and the geographical isola- tion of the area have led to a number of hypotheses pos- tulating undescribed herpetological species, especially in the montane forest patches of the Tchabal Mbabo (BOHME & KLAVER 1981; BOHME & SCHNEIDER 1987). Jean-Louis AMIET (Nyons, pers. comm. 2003) surveyed the area around Hama Aoudi im May 1970 and December 1972. In 1978, Amiet lists an undescribed species of Bu- fo (B. sp. 2, “espece orophile du Tchabal Mbabo”) and an undescribed species of Phrynobatrachus (P. sp. 5, “espece orophile endémique du Tchabal Mbabo”) in his “Liste pro- visoire des amphibiens anoures du Cameroun”. THOMAS & THOMAS (1996) report on a forthcoming herpetologi- Bonner zoologische Beiträge 55 (2006) 29 30 25 2 20 = =| £ o 2 EE 10 5 : FMAMJJAS ON DJFMAMJS JASONDJFMAMJJ February 2000 - July 2002 100 80 se = = 60 £ 3 Tos o = E 40 = 20 FMAMJJASONDJFMAMJ JAS ON DJIFM AMS J Fig. 3. Temperature in °C (a) and relative humidity in % (b) over a period of 30 months in a gallery forest in the vicinity of Fun- goi on the south flank of Tchabal Mabao at an elevation of approximately 2,100 m. 30 Hans-Werner HERRMANN et al.: Amphibians and Reptiles of Tchabal Mbabo cal survey by Dwight Lawson, who apparently visited the area in August 1997 (LAWSON 2000). In the literature very few amphibians and reptiles have been reported from Tch- abal Mbabo: Astvlosternus rheophilus tchabalensis Ami- et, 1977, Leptopelis nordequatorialis Perret, 1966 (in BÓHME & SCHNEIDER 1987) and Crotaphatrema tchabalm- baboensis Lawson, 2000 (see Lawson 2000). In January / February 2000 we (H.-W.H., A.S. & P.A.H.) surveyed the herpetofauna of Tchabal Mbabo for 14 days. This period is at the end of the dry season and was cho- sen by need of vehicle accessibility. Accordingly, we ex- perienced no rain which may have influenced the num- ber of species found. Additionally, in November 1998 a two week herpetological sampling expedition was con- ducted by two native field assistants at the beginning of the dry season. Here we present a systematic list of the combined findings of these two expeditions. 2. RESULTS 2.1. Survey sites We surveyed the following localities (vegetation charac- terization follows LETOUZEY 1968 and THOMAS & THOMAS 1996). Altitude was measured with a Global Positioning System (GPS) unit and verified with a conventional al- timeter. 1. Sambolabo, 7°04°94”N, 11°59°06”E, 1,032 m eleva- tion, southern slope, Sudano-guineean savanna with Daniellia oliveri and Lophira lanceolata, very degrad- ed and managed as permanent pasture. bo . Mayo Kelele, 7°10°78”N, 12°01’68”E, 1,675 m ele- vation, southern slope, submontane wooded Hyparrhe- nia savanna, gallery forests with Syzygium guineense. ios) . 5 km NEE of Fungoi, 7°15’11”N, 12°03’58”E, 2,060 m elevation, on crest of southern slope, montane grass- land with Sporobolus indicus and gallery forest domi- nated by Syzygium guineense, Ilex mitis and Svmpho- nia globulifera, northern slope with montane forest and scrub. 4. Hunter’s Hut, 7°18’81”N, 12°02’83”E, 1,282 m ele- vation, northern slope, savanna with Lophira alata and Daniella oliveri, gallery forest diverse with many se- mi-deciduous species. 5. Mayo Selbe (Fig. 2d), 7°21’88”N, 12%02”58”E, 790 m elevation, for landscape and vegetation see Hunter’s Hut. 6. Dodeo, 7°28’03”N, 12°04’10”E, 681 m elevation, sa- vanna with /soberlinia doka, including gallery forests along flat flood plains, often with wetland prairies. 7. Sanganadje, no coordinate data, 1,000 m elevation, ca. 15 km SSE of Galim, for landscape and vegetation see Hunter’s Hut. 2.2. Survey methods Opportunistic visual encounter surveys (CRUMP & SCOTT 1994) were the predominant survey method we used. We sampled during day and night time. Additionally we in- stalled three Y-shaped drift fence/pitfall trap arrays with 5 m long segments (CORN 1994) in the vicinity of study site 3 near Fungoi. Two arrays were situated in a gallery forest (southern slope), one close to a creek. The third ar- ray was on a steep slope in the montane forest of the north- ern slope of the ridge. We worked the drift fences for the full study period oftwo weeks in January / February 2000, with two controls of pit falls every day (morning and evening), without trappıng success. 2.3. Species account Nomenclature in the subsequent systematic list of amphib- ian and reptile species follows Frost (2002) for amphib- 1ans and UETZ et al. (2004) for reptiles. All voucher spec- imens are deposited in the Zoologisches Forschungsmu- seum Alexander Koenig, Bonn (ZFMK) or in the collec- tion of the senior author (HWH) in Cameroon. The latter specimens will be transferred to the herpetological collec- tion of the National Museum of Natural History, Smith- sonian Institution, Washington, D.C. (USNM). Anura Bufonidae Bufo maculatus Hallowell, 1854 Locality: 10 km E of Sambolabo, in village, 1 specimen, ZFMK 68945 (15.X1.-7.X11.1998). Fig. 4. Bufo sp. (cf. Bufo sp. 2 sensu AMIET, 1978). Bonner zoologische Beiträge 55 (2006) 3] Bufo regularis Reuss, 1834 Localities: Sambolabo, in village, 2 specimens, ZFMK 75630 (‘, 24.1.2000), ZFMK 75730 (7.11.2000); Dodeo, in village, 1 specimen, ZFMK 75631 (‘, 2.11.2000). Bufo sp. (cf. Bufo sp. 2 sensu AMIET 1978) (Fig. 4) Locality: 5 km NEE of Fungoi, in creek in gallery forest on southern slope, 1 specimen, ZFMK 75769 (subadult, 30.1.2000). Remarks: The specimen possesses smooth paratoid glands which are medially constricted. Arthroleptidae Arthroleptinae Arthroleptis sp. (Fig. 5) Locality: Hunter’s Hut, near small creek in gallery forest, 1 specimen, ZFMK 75704 (31.1.2000). Cardioglossa alsco Herrmann, Herrmann, Schmitz & Böhme, 2004 (HERRMANN et al. 2004) Locality: 5 km NEE of Fungoi, in creek in gallery forest on southern slope, 73 specimens, ZFMK 75713-25, 77677-736 (11 &, 13 9, 49 subadults, 28.-29.1.2000). Fig. 5. Arthroleptis sp. from small creek on northern slope at ca. 1,200 m elevation. Astylosterninae Astylosternus Werner, 1898 Remarks: All Astylosternus were strongly polymorphic and difficult to assign to any described taxon. We identi- fied A. rheophilus tchabalensis based on morphology. Based on a 600 bp long segment of the 16S rRNA gene we identified two of the other polymorphic Astylosternus as conspecific with the latter species. Astylosternus rheophilus tchabalensis AMIET, 1977 Locality: 5 km NEE of Fungoi, in creek in gallery forest on southern slope, 5 specimens, ZFMK 75770-4 (30.1.-5.11.2000). Remarks: Our specimens show a remarkable polymor- phism and differ from topotypic A. r. rheophilus of Mt Lefo (Fig. 6a and 6b). Astylosternus sp. (Fig. 6c and 6d) Locality: Mayo Selbe, in creek in gallery forest, 4 speci- mens, ZFMK 75775-8 (2./3.11.2000). Astylosternus sp. indet. Locality: Dodeo, in creek in gallery forest, 15 tadpoles, ZFMK 75632-42 (5.11.2000). Leptodactylodon sp. Locality: 5 km NEE of Fungoi, in creek in gallery forest on southern slope, 1 juvenile specimen, ZFMK 75703 (4.11.2000). Ranidae Raninae Amnirana longipes (Perret, 1960) Locality: Sanganadje, 2 specimens, ZFMK 68946-7 (15.XI.—7.X11.1998). Ptychadena sp. Locality: Sanganadje, 1 (15.XI.—7.X11.1998). specimen, ZFMK 68948 Phrynobatrachus steindachneri Nieden, 1910 (Fig. 7a) Locality: 5 km NEE of Fungoi, in creek in gallery forest on southern slope, 8 specimens, ZFMK 75705-12 (28.1.2000). Phrynobatrachus spp. (Fig. 7b) Locality: 5 km NEE of Fungoi, in creek in gallery forest on southern slope, 64 specimens, ZFMK 75643-702, 75726-9 (28.1./4.11.2000). Remarks: Juvenile to subadult specimens with some prob- ably being P. steindachneri. Others could not be identi- fied. Hyperoliidae Hyperolius riggenbachi Nieden, 1910 Locality: Sanganadje, $ specimens, ZFMK 68949-56 (5 3, 3 9, 15.X1.-7.X11.1998). Squamata Gekkonidae Hemidactylus brooki Gray, 1845 Locality: Dodeo, 1 specimen, ZFMK 75749 (9, 2.11.2000). Agamidae Agama agama (Linnaeus, 1758) Localities: Sambolabo, 4 specimens, ZFMK 75731-34 (2 9, 2 9, 7.11.2000); Mayo Kelele, 4 specimens, ZFMK 75736-39 (2 9, 2 9, 7.11.2000). Agama doriae benuensis Monard, 1951 Locality: Dodeo, 1 specimen, ZFMK 75748 (9, 2.11.2000). 35) Hans-Werner HERRMANN et al.: Amphibians and Reptiles of Tchabal Mbabo Fig. 6. (a) Astylosternus rheophilus rheophilus from Mt Leto, (b) Astvlosternus cf. rheophilus tchabalensis from near Fungoi at an elevation of ca, 2,000 m, (c) and (d) Astvlosternus sp. from Mayo Selbe at ca. 800 m elevation. Chamaeleonidae Chamaeleo gracilis gracilis Hallowell, 1842 Locality: 15 km E of Sambolabo, 1 specimen, ZFMK 68942 (9, 15.X1.-7.X11.1998). Chamaeleo senegalensis Daudin, 1802 Locality: Dodeo, 2 specimens, ZFMK 73092, 75747 (2 9, 2.11.2000), Mayo Kelele, 1 specimen, HWH 1214 (9, no date). Chamaeleo wiedersheimi wiedersheimi Nieden, 1910 Localities: Mayo Kelele, 1 specimen, ZFMK 68943 (9, 15.X1.-7.X11.1998), 4 specimens, ZFMK 75740-43 (3 >, 1 9, 7.11.2000), 4 specimens, HWH 1210-13 (3 3, 1 9, no date); 5 km NEE of Fungoi, in gallery forest on southern slope, 3 specimens, ZFMK 75744-46 (1 9, 2 9, 25.1.2000). Scincidae Lacertaspis chriswildi (Böhme & Schmitz, 1996) Locality: 5 km NEE of Fungoi, in gallery forest on south- ern slope, | specimen, ZFMK 75735 (29.1.2000). Trachylepis sp. Locality: 3 km NEE of Mayo Kelele, on path in grassland, | juvenile specimen (25.1.2000, Fig. 8). Remarks: This specimen persisted only as photograph and tissue voucher. We follow MAUSFELD et al. (2002) but pre- fer to use the genus name 7rachylepis instead of Euprepis (sensu BAUER 2003). This specimen resembles juvenile 7: perrotetii (Dumeril & Bibron, 1839) but has a blue tail and shows a clear genetic differentiation from the latter species (MAUSFELD-LAFDHIYA et al. 2004). A blue tail in juveniles is otherwise only known in 7. quinquetaeniata (Lichten- stein, 1823) which, however, has a distinctly striped body. Varanidae Varanus niloticus (Linnaeus, 1758) Locality: Sambolabo, | specimen (head and skin), ZFMK 68930 (15.X1.-7.X11.1998). Boidae Python sebae (Gmelin, 1789) Locality: Mayo Kelele, 1 specimen (Fig. 9, no date), head of a specimen caught by villagers. Bonner zoologische Beitráge 55 (2006) 33 Fig. 7. (a) Phrynobatrachus cf. steindachneri from gallery forest near Fungoi at an elevation of ca. 2,000 m, (b) P. sp. from the same locality. Fig. 9. Head of adult Python sebae from hunters at Mayo Ke- lele. Colubridae Philothamnus angolensis Bocage, 1882 Localities: Sangandje, 11 specimens, ZFMK 68931-41 (15.X1.-7.X11.1998); Mayo Kelele, 1 specimen (head), HWH 1218 (no date); 5 km NEE of Fungoi, in gallery for- est on southern slope, 1 specimen, ZFMK 75751 (subadult, 2.11.2000). Elapidae Dendroaspis jamesoni jamesoni (Traill, 1843) Locality: Dodeo, 1 specimen, ZFMK 75750 (subadult, 2.11.2000). Naja nigricollis Reinhardt, 1843 Locality: Mayo Kelele, 1 specimen (head), HWH 1219 (no date). Viperidae Bitis arietans (Merrem, 1820) Locality: 10 km E of Sambolabo, 1 specimen (head), ZFMK 68944 (15.X1.-7.X11.1998). Causus rhombeatus (Lichtenstein, 1823) Locality: Mayo Kelele, 3 specimens (heads), HWH 1215- 17 (no date). 3. DISCUSSION A total of 30 herpetological species (one caecilian, 14 anu- rans, nine lizard and six snake species) are recorded from the Tchabal Mbabo Mtns. The number of recorded am- phibian species corresponds with comparable surveys in Western and Central Africa as listed in RÓDEL & AGYEI (2003). However, numbers from thoroughly surveyed sa- vanna/forest areas in West Africa suggest amphibian species counts of at least twice of those presented here. All species, except Lacertaspis chriswildi, are known as either savanna or farmbush/forest edge species and are not restricted to forests. Astylosternus, Cardioglossa or Lep- 34 Hans-Werner HERRMANN et al.: Amphibians and Reptiles of Tchabal Mbabo todactylodon are predominately found in forests with ex- ception of those species adapted to high elevation grass- lands and gallery forests. Lacertaspis chriswildi ıs known from only two specimens, the holotype which originated from the high elevation montane forest at Mt Kupe (BÖHME & SCHMITZ 1996) and one further specimen from the Takamanda lowland rain- forest, both in Southwestern Cameroon. Our finding rep- resents the first record of this species from a gallery for- est within high elevation grassland. It also extends the range of the species approximately 400 km to the north- east. This suggests its occurrence in other areas along the “Dorsale Camerounaise”. The high number of taxa which could not be assigned to any described species (1.e., Bufo sp., Arthroleptis sp., Asty- losternus sp., Leptodactylodon sp., Ptychadena sp. and Phrynobatrachus spp.) as well as the discovery of Asty- losternus rheophilus tchabalensis, Crotaphatrema tcha- balmbaboensis (LAWSON, 2000) and Cardioglossa alsco (HERRMANN et al. 2004) are indicative of both the paucı- ty of knowledge on the Tchabal Mbabo herpetofauna in general and the uniqueness of the anuran fauna with its endemism in particular. Future surveys to the Tchabal Mbabo Mtns are warrant- ed in order to clarify the taxonomic status of specimens collected during our surveys and to gain comprehension on the extant herpetofauna. The discovery of hitherto un- described species is likely. Due to the area’s remoteness and to the traditional lifestyle of the Fulanı herdsmen, no immediate threat to the herpetofauna of the area was ob- served. Nonetheless, efforts to preserve the unique mon- tane forests on the northern slopes as well as the semi-de- ciduous and gallery forests are warranted to ensure their welfare in the future. The montane forests on the north- facing slopes are relatively secure due to their extreme to- pography. The gallery forests on the southern slopes how- ever, are accessible to the Fulanı cattle and are increas- ingly degraded by husbandry practices. Acknowledgements. We are grateful to the Cameroon Ministry of Scientific and Technical Research (MINREST) for issuing re- search permits and the Ministry of Environment and Forestry (MINEF) issuing collecting and export permits. H. Nobis of ALSCO (Germany) provided financial support. Additionally, H.- W. H. was supported by the Cologne Zoo, Germany. P. Jaquet, Much, provided transportation and helped with other logistics. Ngolepie George and Alowbede Johnson undertook the first ex- pedition in 1998. I. Itoua and C. 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Revue Suis- se de Zoologie 103: 767-774. BÖHME, W. & SCHNEIDER, B. 1987. Zur Herpetofaunistik Kame- runs (III) mit Beschreibung einer neuen Cardioglossa (Anu- ra: Arthroleptidae). Bonner zoologische Beiträge 38: 241— 263. CORN, P. S. 1994. Straight-line drift fences and pitfall traps. Pp. 109-117 in: HEYER, W. R., DONNELLY, M. A., MCDIARMID, R. W., HAYEK, L.-A. C. & FOSTER, M. S. (eds.) Measuring and Monitoring Biological Diversity. Standard Methods for Amphibians. Smithsonian Institution Press, Washington, D.C. Crump, M. L. & SCOTT JR., N. J. 1994. Visual encounter sur- veys. Pp. 84-92 in: HEYER, W. R., DONNELLY, M. A., Mc- DIARMID, R. W., HAYEK, L.-A. C. & FOSTER, M. S. (eds.) Measuring and Monitoring Biological Diversity. Standard Methods for Amphibians. Smithsonian Institution Press, Washington, D.C. EUSKIRCHEN, O., SCHMITZ, A. & BOHME, W. 1999. Zur Herpe- tofauna einer montanen Regenwaldregion in SW-Kamerun (Mt. Kupe und Bakossi-Bergland) — II. Arthroleptidae, Ra- nidae und Phrynobatrachidae. herpetofauna, Weinstadt 21 (122): 25-34. EUSKIRCHEN, O., SCHMITZ, A. & BÖHME, W. 2000. Zur Herpeto- fauna einer montanen Regenwaldregion ın SW-Kamerun (Mt. Kupe und Bakossi-Bergland) — IV. Chamaeleonidae, biogeographische Diskussion und Schutzmaßnahmen. her- petofauna, Weinstadt 22 (125): 21-34. Frost, D. R. 2002. Amphibian Species of the World: an online reference. V2.21 (15 July 2002). Electronic database avail- able at http://research.amnh.org/herpetology/amphibia/in- dex.html HERRMANN, H.-W., BÓHME, W. & HERRMANN, P. 2000. The ALSCO Cameroon expedition 1998: The sampling of a mountain rainforest. Proceedings of the 4th International Symposium of Isolated Vertebrate Communities in the Trop- ics, Bonner zoologische Monographien 46: 95-103. HERRMANN, H.-W., HERRMANN, P. A., SCHMITZ, A. & BÖHME, W. 2004. A new frog species of the genus Cardioglossa from the Tchabal Mbabo Mtns, Cameroon. Herpetozoa 17: 119-125. HERRMANN, H.-W., BÖHME, W., HERRMANN, P. A., PLATH, M., SCHMITZ, A. & SOLBACH, M. 2005a. African biodiversity hotspots: The amphibians of Mt Nlonako, Cameroon. Sala- mandra 41: 61-81. Bonner zoologische Beiträge 55 (2006) 35 HERRMANN, H.-W., BÖHME, W., EUSKIRCHEN, O., HERRMANN, P. A& SCHMITZ, A. 2005b. African biodiversity hotspots: The reptiles of Mt Nlonako, Cameroon. Revue suisse de Zoolo- gie 112: 1045-1069. JOGER, U. 1982. Zur Herpetofaunistik Kameruns (II). Bonner zoologische Beiträge 33: 313-342. Lawson, D. P. 2000. A new caecilian from Cameroon, Africa (Amphibia: Gymnophiona: Solecomorphidae). Herpeto- logica 56: 77-80. LEBRETON, M. 1999. A Working Checklist of the Herpetofauna of Cameroon.- Netherlands Committee for IUCN, Amster- dam. LETOUZEY, R. 1968. Etude phytogéographique du Cameroun. Paris. MAUSFELD, P., SCHMITZ, A., BÖHME, W., MISOF, B., VRCIBRADIC, D. & RocHA, C. F. D. 2002. Phylogenetic affinities of Mabuya atlantica SCHMIDT, 1945, endemic to the Atlantic Ocean archipelago of Fernando de Noronha (Brazil): neces- sity of partitioning the genus Mabuya FITZINGER, 1826 (Scin- cidae: Lygosominae). Zoologischer Anzeiger 241: 281-293. MAUSFELD-LAFDHIYA, P., SCHMITZ, A., INEICH, I. & CHIRIO, L. 2004. Genetic variation in two African Euprepis species (Reptilia; Scincidae), based on maximum-likelihood and Bayesian analyses: taxonomic and biogeographic conclu- sions. Bonner zoologische Beitráge 52: 159-177. PERRET, J.-L. 1959. Études herpétologiques africaines. Bulletin de la Société Neucháteloise des Sciences Naturelles 82: 247-253. PERRET, J.-L. 1960. Études herpétologiques africaines II. Bulle- tin de la Société Neucháteloise des Sciences Naturelles 83: 93-100. PERRET, J.-L. 1961. Études herpétologiques africaines III. Bul- letin de la Société Neucháteloise des Sciences Naturelles 84: 130-138. PERRET, J.-L. 1966. Les Amphibiens du Cameroun. Zoologische Jahrbücher. Abteilung für Systematik, Ökologie und Geo- graphie der Tiere 8: 289-464. RÓDEL, M.-O. & AGYEI, A. C. 2003. Amphibians of the Togo- Volta highlands, eastern Ghana. Salamandra 39: 207-234. SCHMITZ, A., EUSKIRCHEN, O. & BÖHME, W. 1999. Zur Herpe- tofauna einer montanen Regenwaldregion in SW-Kamerun (Mt. Kupe und Bakossi-Bergland) — I. Einleitung, Bufoni- dae und Hyperolidae. herpetofauna, Weinstadt 21 (121): 5-17. SCHMITZ, A., EUSKIRCHEN, O. & BOHME, W. 2000. Zur Herpe- tofauna einer montanen Regenwaldregion in SW-Kamerun (Mt. Kupe und Bakossi-Bergland) — II. Einige bemerkens- werte Vertreter der Familien Lacertidae, Scincidae, Varani- dae, Elapidae und Viperidae. herpetofauna, Weinstadt 22 (124): 16-27. THOMAS, D. & THOMAS, J. 1996. Tchabal Mbabo Botanical Sur- vey. Report for WWF CPO Yaounde, Cameroon. UETZ, P., ETZOLD, T. & CHENNA, R. 2004. The EMBL reptile da- tabase. An online information resource on reptile taxonomy with focus on the species level. Heidelberg. http://www.embl-heidelberg.de/-uetz/livingreptiles.html Authors’ addresses: Hans-Werner HERRMANN (corres- ponding author), Center for Reproduction of Endangered Species (C.R.E.S.), Zoological Society of San Diego, P.O. Box 120551, San Diego, CA 92112-055, USA; Present ad- dress: School of Natural Resources & Genomic Analysis and Technology Core, University of Arizona, 1041 East Lowell Biological Sciences West, 246b, Tucson, AZ 85721, USA; E-Mail: hwh@email.arızona.edu. Andreas SCHMITZ, Department of Herpetology & Ichthy- ology, Museum of Natural History, C. P. 6434, 1211 Gene- va 6, Switzerland; Patricia A. HERRMANN, P.O. Box 3055, Messa, Yaounde, Cameroon & Wolfgang BÖHME, Zoolo- gisches Forschungsmuseum Alexander Koenig, Adenauer- allee 160, 53113 Bonn, Germany. Received: 28.04.2005 Accepted: 07.07.2005 Corresponding editor: R. van den Elzen Bonner zoologische Beitráge Band 55 (2006) Heft 1 Seiten 37-46 Bonn, Januar 2007 The Position of Trochiliphagus Carriker within the Ricinidae (Insecta: Phthiraptera) Goetz RHEINWALD St. Katharinen, Germany Abstract. Two forms of the Ricinidae (Mallophaga, Phthiraptera) that live on Trochilidae have independently developed piercing mouthparts. The one (Trochiloecetes) is obviously an ancient inhabitant of hummingbirds, whereas the other has settled these birds rather recently. CARRIKER (1960) established a new genus “Trothiliphagus” for the second one. It is demonstrated that these lice belong to Ricinus de Geer, 1778, a genus with normal biting mouthparts which is widespre- ad on Passeriformes. Trochiliphagus Carriker, 1960 ıs placed as a junior synonym of Ricinus and all species described till now should be named Ricinus jimenezi. Within Ricinus these hummingbird-dwelling lice are best placed in the doli- chocephalus species-group. Obviously mouthparts have a high plasticity and are not good for diagnostic characters. Keywords. Ricinus jimenezi, piercing mouthparts, secondary settlement. 1. INTRODUCTION CLay (1949) detected that among lice with biting mouth- parts (= Mallophaga) the members of the genus Trochiloe- cetes Paine & Mann, 1913 that live on hummingbirds have developed piercing mouthparts independent of the Anoplu- ra. She described details of the mouthparts that emerged from the hypopharynx and the ligular sclerites. Accord- ing to CLAY the mandibles are reduced. Rather incidentally CLAY mentions that on hummingbirds there are also members of the genus Ricinus de Geer, 1778 that have also lost the typical mouthparts of Mallophaga, though the changes are not as far-reaching as in Trochiloe- cetes. She continues: “The Ricinus species on the Trochili are typical of the genus apart from the modified mouth- parts, and this modification, therefore, must have taken place in the Ricinus species of the Trochili after the mor- phological characters of the genus Ricinus had been es- tablished; Trochiloecetes, closely related to Ricinus and restricted to the Trochili, must be a later derivate from a Ricinus-like ancestor.” CARRIKER (1960) revised this group of lice, but created many problems. For example, he established a new fami- ly (Trochiliphagidae) for the two genera parasitizing hum- mingbirds. However, he designated Trochiloecetes as genotype. Thus the correct family spelling should be Trochiloecetidae (s. EICHLER 1963). CARRIKER, too, trans- ferred the Ricinus-like mallophaga (s. CLAY) to a new genus Trochiliphagus. CARRIKER (1960) believed that Ricinus is the progenitor for both, Trochiloecetes and Trochiliphagus. He ignored that Ricinus itself digests blood and not feathers and con- cluded that both on hummingbirds living genera developed the sucking apparatus because of lack of suitable feath- ers. CARRIKER was not aware that the piercing mouth parts of Trochiloecetes and Trochiliphagus have developed in- dependently. In his mind the change from biting to pierc- ing mouthparts is so essential that it justifies the separa- tion into a new family. In his monograph EICHLER (1963) presents a system for “Mallophaga” within which the “superfamilia” Lae- mobothrioidea includes two “interfamilias”: Laemobo- thriformia and Riciniformia. Within the latter he defines two families, Ricinidae (with the genus Ricinus) and Trochiloecetidae (with the genera: Trochiloecetes and Trochiliphagus). More recently, PALMA (1996) in the Zo- ological Catalogue of Australia placed all three mentioned genera in the family Ricinidae, as did PRICE et al. (1993) in the last World Checklist. I will follow this decision here. Members of Ricinus have biting mouthparts and feed on blood of species of Passeriformes s. lat., whereas those of Trochiloecetes and Trochiliphagus have sucking mouth- parts and feed on blood of hummingbirds (Trochilidae). An earlier paper (RHEINWALD 1968) contained a revision of the Old-world species of Ricinus and established four species-groups. NELSON (1972) revised the New-world species and established eight species-groups, of which three are identical with the old-world groupings. There- fore six species-groups are recognized with altogether 54 species. 38 Gótz RHEINWALD: Position of Trochiliphagus (Insecta: Phtiraptera, Ricinidae) Fig. 1. Head and prothorax of females a: Trochiloecetes ochoterenai?, b: “Trochiliphagus”, c: Ricinus brevicapitatis (from NEL- son 1972, plate 2), d: R. dolichocephalus (from RHEINWALD 1968, Fig. 13). Left from dorsal, right from ventral. Scale bar 0.1 mm. Bonner zoologische Beitráge 55 (2006) 39 CARRIKER (1960) identified at least ten species of Trochiliphagus based on minimal size differences and minute changes in shape of the clypeus and prothorax, these differences are often an artefact due to preparation. Generally he says (1960: 331): “There is a generic con- formity in the general shape of the head and in the absence of darker markings, following the type of Ricinus micro- cephalus! Kellogg, except that the head is longer and usu- ally narrower in the temples.” Nearly all specimens of Trochiliphagus which are deposit- ed in US collections were collected by M. A. CARRIKER. Most of these specimens from a large number of hosts were loaned and studied by Dr. J. Willis-Oniki (Rio Claro, Brasilia) and me in Bonn. Here I noticed the enormous similarity between Trochiliphagus and Ricinus. It is the aim of this paper to clarify the position of Trochiliphagus within the Ricinidae as far as this is pos- sible with conventional methods. 2. MATERIALS AND METHODS Drawings were made and measurements taken from the specimens listed in the Appendix. Usually lice are killed and conserved in 70 % ethanol. For mounting on slides they first are hydrated, then treated with weak KOH to re- solve the non-chitinized parts of the inside and then em- bedded in resin. The resolution of the inner parts is nec- essary; otherwise details of the chitinized skeleton will not be visible. But that means that all characters we know are features of the chitinized skeleton, upon which all species descriptions are based exclusively. For a detailed study of the mouthparts of Trochiloecetes and Ricinus we possi- bly also need histological sections. In my revision of Ricinus I figured head and prothorax of every species as well as the terminal segments of the fe- male in dorsal and ventral views. Additionally mandibles, labium and cardo-stipes, gular sclerite and the sternites of pro- and metathorax as well as the male genitalia were pre- sented. NELSON (1972) also followed this format. There- fore I will present here for comparison just these parts in the same view. Additionally claws and limbs of the third leg of a female are measured and drawn. 3. RESULTS 3.1. Head and prothorax In Figure 1 the head and prothorax of Trochiloecetes ochoterenai?, “Trochiliphagus”, Ricinus brevicapitatis Carriker (from NELSON 1972, Table 2) and R. dolicho- 1) Which according to my experience (s. RHEINWALD 1968) belongs to R. Jringillae. cephalus (Scopoli) are presented in the same arrangement. Ricinus brevicapitatis is the typical species parasitizing Cotingidae — a bird family close to the base of the Passer- iformes. If the classification in EICHLER (1963) is accept- ed, then Trochiliphagus and Trochiloecetes should be most similar. However, as Figure | shows, the highest similar- ity exists between R. dolichocephalus and “Trochilipha- gus”. A special feature of “Trochiliphagus” and some species of the dolichocephalus-group 1s that the maxillary palpi exceed far beyond the sides of the head, whereas in all other species the palpi do not reach the border of the head. Additionally, the dorsal lamina of the occiput is well developed. 3.2. Mandibles (Fig. 2) The only clearly visible feature of the mandible is the ar- ticulation. If we follow the sclerites from the articulation further we find in Trochiloecetes the structure which CAR- RIKER (1960) called the “collar”, but it is most likely the remaining portion of the mandible. The mandible of CAR- RIKER appears to be part of the basal limb of the palpus. In (b) Trochiliphagus”) homologizing the “mandibles” is highly interpretable. Section series would be necessary to clarify these structures. 3. Gular plate, prosternite and metasternite (Fig. 3) When we compare again the same forms, Trochiliphagus differs significantly from R. brevicapitatis and Trochloe- cetes especially in the form of the gular plate and the pro- sternite. On the other hand, the agreement between “Trochiliphagus” and R. dolichocephalus is great, even in details in the form of the gular plate and the form of the sclerites in the pro- and metasternite. In RHEINWALD (1968), all species of the dolichocephalus-group (Figs 1321, each no. d) generally show the same type as “Trochiliphagus”, while the species of the rubeculae- group (Figs 22—33, each no. d) differ to some degree. 3.4. Labium and cardo-stipes (Fig. 4) As the figures show, the labium is very uniform in all Ricinidae and therefore contributes little to the systemat- ics of the group. But obviously the cardo-stipes in Trochiloecetes and “Trochiliphagus” assists considerably the stiletto in its function (see also Fig. 1). The plates be- hind the labium (cardo-stipes) in Trochiloecetes (a) have a rectangular form and in that differ considerably from Ricinus. As Figure | shows, the ligular sclerite of Trochiloecetes is connected directly with the stiletto and 40 Götz RHEINWALD: Position of Trochiliphagus (Insecta: Phtiraptera, Ricinidae) Fig. 2. Mouthparts of a: Trochiloecetes ochoterenai?, b: “Trochiliphagus”, and c: R. dolichocephalus (from RHEINWALD 1968, Fig. 13). Scale bar 0.1 mm. + Fig. 3. Gular plate, prosternite and metasternite of a: Trochiloecetes ochoterenai?, b: “Trochiliphagus”, e: Ricinus brevicapitatis (after NELSON 1972, combination from plate 2 and 3) and d: R. dolichocephalus (from RHEINWALD 1968, Fig. 13). Scale bar 0.1 mm. Fig. 4. Labium and cardo-stipes (= maxillar plate according to NELSON 1972) of a: Trochiloecetes ochoterenai?, b: “Trochilipha- gus”, and c: R. dolichocephalus (from RHEINWALD 1968, Fig. 13). In b) an additional intermediate plate. Scale bar 0.1 mm. Bonner zoologische Beitráge 55 (2006) 4] at its protruding edges a muscle appears to be attached which stretches well forward (dashed line); obviously the stiletto is bored into the skin by the ligular sclerite. The cardo-stipes is not enlarged, nor sclerotized. In “Trochiliphagus” (b) the ligular sclerite appears not to be connected with the stiletto, but the plate between the cardo-stipes 1s lengthened into a long, well sclerotized plate. The cardo-stipes in general has the form that is found in all Ricinus species (see NELSON 1972; RHEINWALD 1968) but it is enlarged and the inner margin is thickened. Ob- viously the short stiletto of “Trochiliphagus” 1s moved by these plates and bored into the skin of the host. Though it is very difficult to see the single parts of the stiletto in both forms with this preparation method, it seems to be rather clear that the stilettos are moved absolutely differ- ently, which underlines the meaning that the two stilettos have developed convergently. 3.5. Female postabdomen (Fig. 5) Trochiloecetes differs most; lacking pigmented sternal plates, having well pigmented pleurites to segment VI on- ly and a very small anal tergite (cf. RHEINWALD 1968). Arrangement and relative length of setae differs in many respects from that of Ricinus species. Ricinus brevicapi- tatis (c) has, like all Ricinus species, pigmented sternites, pleurites up to segment VIII and a fringe of small setae at the well-developed anal tergite. “Trochiliphagus” and R. dolichocephalus are nearly identical, both having the anal tergite surpassing the margin of segment XI and by that the tip of the abdomen, and in the arrangement and the relative length of the tergal and sternal setae. 3.6. Male genitalia (Fig. 6) In the three species of Trochiloecetes the general form is rather simple. The basal plate (cf. RHEINWALD 1968, Fig. lg) is not pointed, the parameres are short and rounded and the genital sclerite is a simple, more or less rectan- gular structure without significant ornamentation. In Rici- nus arctuatus (Kellog & Mann) (from Tyrannidae, plate 5 of NELSON 1972) and in other species-groups of Rici- nus we find many peculiarities in the form of the meso- somal plate, the length of the parameres, the form and or- namentation of the genital sclerite. We find high agree- ment between “Trochiliphagus” and R. dolichocephalus. This is especially true in the form of the mesosomal plate and the length of the parameres; in all species of the dolichocephalus-group and in “Trochiliphagus” the preputial-sac is ornamented with fine structures which are missing in other species-groups. 3.7. Tarsus (Fig. 7) In my revision of the genus Ricinus (RHEINWALD 1968) I did not investigate the tarsus. When studying more care- fully members of the genus Trochiloecetes | noticed that they have huge claws on the last tarsal limb. Thereupon I compared the tarsi of Trochiloecetes, “Trochiliphagus” and several species-groups of Ricinus. Some general remarks can be made: Obviously within a species the tarsı are morphologically identical, which means that besides individual differences which derive from mounting (as position, angle of the limbs to one an- other, wrinkles, shrinkages) and bad recognition deriving from the smallness of the object, a series of elements are species-specific. That is especially true for the form of the claws, the form of the outer sclerite at the second tarsal limb, position and size of the two “stonelets”(the homol- ogy of which is not clear to me). The form of the inner (second) tarsal limb is species-specific too, even when it is in most cases hard to see and by the position of the out- er limb (the angle between outer limb and tibia) as a con- sequence of mounting is often heavily deformed. As far as I have material, the species within a species- group of Ricinus (fringillae- and rubeculae-group) appear to correspond in the proportions of the tarsal limbs, the proportions of the claw, the form of the outer sclerite at the tarsal limbs and the tibia as well as in the length of the four large setae at the tip of the inner tibia. That means that members of Ricinus and Trochiloecetes are well dis- tinguishable by their tarsi. But within the genus Ricinus there are also group-specific differences. On the one hand these are the lengths of the claw and the second tarsal limb relative to the total length. On the other hand the form of the outer sclerites of the second tarsal limb and the tibia and the position of the “stonelets” are group-specific. In connection with the questions in this paper it is of main interest whether “7rochiliphagus”’ shows similarities to Trochiloecetes or to Ricinus. Table 1 shows that Trochiloe- cetes is separated from all other forms as well by the rel- ative length of the claw as by the relative length of the tarsus. The form of the lateral sclerites of the second tarsal limb and tibia are also clearly different. The tarsus of Rici- nus rubeculae (Schrank) appears also autapomorphic. It has the relatively smallest claw, the shortest tarsus, and the form of the lateral sclerite differs distinctly. In opposite to Ricinus rubeculae, “Trochiliphagus” shows much conformity in the form of the sclerites with R. fringillae and R. dolichocephalus. While the relative length of the claw agrees well with R. fringillae and R. dolichocephalus, the relative length of the tarsus differs and appears to be mid-way between Trochloecetes and 42 Götz RHEINWALD: Position of Trochiliphagus (Insecta: Phtiraptera, Ricinidae) © e Me rs, vorm a OMAN Fig. 5. Abdominal segments and the terminal plates of a: Trochiloecetes ochoterenai?, b: “Trochiliphagus”, c: Ricinus brevica- pitatis (after NELSON 1972, plate 3) and d: R. dolichocephalus (from RHEINWALD 1968, Fig. 13). Left from dorsal, right from ven- tral. Scale bar 0.1 mm. Fig. 6. Male genitalia a: Trochiloecetes rumpununi, b: T. insularis, e: T. ochoterenai, d: “Trochiliphagus”, e: Ricinus dolichoce- phalus (from RHEINWALD 1968, Fig. 13). Scale bar 0.1 mm. Bonner zoologische Beitráge 55 (2006) 43 Table 1. Length of the second tarsal limb and claw, and its relative length in relation to the total length of the two members of Ri- cinidae living on hummingbirds and of some Ricinus species. Trochiloecetes “Trochiliphagus” R. rubeculae R. fringillae R. dolichocephalus total length [mm] 2.18 3:39 3.32 3.02 4.70 length of tarsus [mm] 0.338 0.322 0.178 0.203 0.305 length of the claw [mm] 0.159 0.076 0.038 0.063 0.079 relative length of tarsus 0:153 0.096 0.056 0.067 0.065 relative length of claw 0.073 0.023 0.011 0.021 0.016 Fig. 7. Claws, two tarsal limbs and tip of tibia of a: Trochiloecetes ochoterenai?, b: Ricinus rubeculus, e: "Trochiliphagus”, d: R. dolichocephalus, and e: R. fringillae. Scale bar 0.1 mm. 44 Götz RHEINWALD: Position of Trochiliphagus (Insecta: Phtiraptera, Ricinidae) Ricinus. One may well suppose that similar to the mouth- parts, the tarsı have adapted to the same host-group — the hummingbirds — possibly convergently. 4. DISCUSSION 4.1. Position of “Trochiliphagus” within the Ricinidae For an outgroup comparison members of the family Lae- mobothriidae seem to be optimal since together with Ricinidae it forms the subclass/superfamily Laemoboth- riformia. Laemiobothrion maximum (Scopoli) recently was investigated by PEREZ et al. (1995) with SEM. But comparable studies in Trochiloecetes and Ricinus do not exist. SYMMONS (1952) studied the Mallophagan head and also presented figures and descriptions of Laemobothri- on and description of Ricinus. But polarities can not be derived from this. Larger series of Laemobothrion maxi- mum were examined, but I was unable to homologize the different parts of the head to derive evolutionary trends. Therefore the question of the polarity between Trochiloe- cetes and Ricinus and within Ricinus remains unsolved. Mallophaga as a rule live strictly host-specific, a fact that led to the rule of NITZscH-KELLOGG. Hummingbirds and songbirds, according to the accepted classifications (e.g., SIBLEY & MONROE 1990) belong to different orders (resp. superorders) that are not closely related. Therefore one should expect that the many species that live on Passeriformes and till now are combined in the genus Rici- nus should be differentiated significantly in their morphol- ogy from those living on Trochiliformes (Trochiloecetes, Trochiliphagus). As demonstrated in the figures this ıs true for Trochiloecetes. Not knowing the polarity, these cha- racters can be understood as autapomorphic for Trochiloe- celes. In the opposite “Trochiliphagus” and Ricinus can not be separated from each other. As CLAY (1949) observed, these animals are part of the genus Ricinus. The cladistic analy- sis of MARSHALL (2003), based on 147 morphological characters, supports this idea. Within Ricinus all specimens of “Trochiliphagus” are best placed within the dolicho- cephalus species-group. If somebody wants to split up the genus Ricinus then these separations have to occur along the species-groups that were described by NELSON (1972) and RHEINWALD (1968) earlier. The differentiations of these species-groups relative to the dolichocephalus-group are so essential, that the small deviations that accompa- nied the development of the stiletto and the reduction of the mandibles in “Trochiliphagus” appear rather insigni- ficant. Herewith I place the genus Trochliphagus Carriker, 1960 as a junior synonym of Ricinus De Geer, 1778. As much as I have seen in the different collections from a variety of hosts there are no consistent morphological differences between these populations. All appear to belong to one morpho-species. This species should be named Ricinus jimenezi Carriker, 1903, though CARRIKER (1960) made 7. lazulus Carriker 1960 the type-species of Trochiliphagus. There exists an older name: Physostomum lineatum Osborn, 1896 from Trochilus colubris (= Archilochus colubris). According to CARRIKER (1960) the three specimens on which the de- scription was based no longer exist. The description makes clear that this is not a Trochiloecetes. If the host is cor- rect this should be a “Trochiliphagus”, though the pierc- ing mouthparts are not mentioned. Under these circum- stances the name “lineatus” in the family Ricinidae should be suppressed. An application was directed to the Inter- national Commission for Nomenclature. 4.2. Primary or secondary? According to MARSHALL (2003) Laemobotriidae are not close relatives to Ricinidae and therefore not their ances- tors. The whole amblycerean louse-family Ricinidae is re- stricted to Trochilidae and Passeriformes. The differences between Trochiloecetes and Ricinus fit well in the usual ideas of the relations between Trochilidae as part of the Apodiformes and the Passeriformes. Since the possession of mandibles is symplesiomorphic within the Amblycera, the piercing mouth parts of Trochiloecetes ıs an autapo- morphy. If this relation is correct, then Ricinus jimenezi must have colonized the Trochilidae secondarily. 4.3. Origin of the secondary colonization The host-shift onto Trochilidae must have been from a songbird of the families Oriolidae, Meliphagidae, Nec- tariniidae, Rhipidurinae, Muscicapidae which are the host- families of the dolichocephalus species-group. But these bird families all occur exclusively in Africa or Asia. How could it happen that a Ricinus from the African-Asian realm colonized a hummingbird in South America? Col- onization from an American family of Passerines would be much more likely. But then Ricinus jimenezi must show relations to species-groups of Ricinus established by NEL- SON (1972). I could not detect any agreement there. Bonner zoologische Beiträge 55 (2006) 45 4.4 Are the Trochilidae a monophyletic taxon? I do not believe that this is a relevant question, though the- oretically it might be that some hummingbirds (the hosts of Trochiloecetes) are swift-relatives while others (the hosts of Ricinus jimenezi) are songbirds. But CARRIKER (1960) said that he had collected both forms from the same host. 4.5. The development of piercing mouthparts Hummingbirds have an extremely thick skin. Since the size of the parasites relative to the size of the host 1s lim- ited, blood-feeding Ricinidae could not grow bigger as they are today. On the smaller hummingbirds these para- sites are accordingly smaller. CARRIKER (1960) noted for Trochiloecetes a maximal body length of 2.54 mm (see Table 1), while according to his information females of “Trochiliphagus” reach 2.57 to 3.8 mm. With this body size the length of the mandibles is not sufficient to open the skin. If therefore a member of Ricinidae were to sur- vive on hummingbirds it had to develop piercing mouth- parts. This should be a plausible explanation for the con- vergent occurrence of two members of Ricinidae with piercing mouthparts on hummingbirds. For Trochloecetes this explanation may be evident. Be- cause these are the original parasites, they should have evolved on hummingbirds since their separation from swifts and could gradually evolve a stiletto in addition to the mandibles; as soon as this stiletto was completely func- tional the mandibles could be reduced. For the secondary settler, Ricinus jimenezi, great obsta- cles must have arisen. When it moved from songbirds to hummingbirds it had mandibles and certainly no stiletto because it did not need it for living on songbirds. But with its mandibles it could not pierce the skin of a humming- bird. Therefore it should have starved and a secondary in- festation would not have happened. If it would have found thinner parts of skin where it may have survived some time why should it then evolve a stiletto? Even when the membership of “Trochiliphagus” to Rici- nus and here to the dolichocephalus species-group seems to be evident, it is absolutely unclear how we can imag- ine the procedure of this secondary colonisation. The ques- tion of the offspring of Ricinus jimenezi and therefore its relations to Ricinus and perhaps the reconstruction of the history of its development might be a fascinating case for the application of modern enzymatic or DNA-techniques. The biggest problem here may be the procurement of re- levant Mallophaga which in no case are abundant and in many cases are known to science just by one specimen. Normally the eggs of Ricinus at the bases of the feathers of neck and anterior breast are so conspicuous that with some experience it is easy to detect those birds where it is worth while searching for adults. 4.6. Plasticity of the mouthparts The occurrence of biting and piercing mouthparts within the genus Ricinus throws new light on the question what value mouthparts have as characters for phylogenetic dis- cussions. It is meanwhile common sense that Ischnocera and Anoplura are sister-groups (BARKER et al. 2003; Hop- KINS 1949; WEBB 1946). In the Anoplura sucking mouth- parts developed for the first time in the Phthiraptera. In the other large group of the Phthiraptera, the Amblycera, a stiletto evolved twice, both within the Ricinidae. On the one hand in Trochiloecetes, which obviously is the cha- racteristic Ricinidae of the Trochilidae, and on the other hand within Ricinus when one of its members secondar- ily colonised Trochilidae. That means that piercing mouthparts evolved at least three times within Phthi- raptera. But if such fundamental alterations are in principle pos- sible, what general value has mouthparts as a phylogenet- ic feature? We have examples from other animal classes, such as birds, in which alterations in the mouthparts with- in small taxonomic units are of similar dimensions. As a consequence in future we should leave out features of the mouthparts in our data sets for the analysis of phylogeny. REFERENCES BARKER, S. C., WHITING, M. JOHNSON, K. P. & MURRELL, A. 2003. Phylogeny of the Lice (Insecta, Phthiraptera) inferred from small subunit rRNA. Zoologica Scripta 32: 407-414. CARRIKER, M. A. 1960. Studies in Neotropical Mallophaga, XVII: A new family (Trochiliphagidae) and a new genus of the lice of hummingbirds. Proceedings of the United States National Museum (Smithsonian Institution, Washington, DC) 112: 307-342. CLay, T. 1949. Piercing mouth-parts in the biting lice (Mallo- phaga). Nature (London) 164: 617. EICHLER, W. 1963. Mallophaga in: Dr. H. G. Bronns Klassen und Ordnungen des Tierreichs. 5. Bd. (Arthropoda), II. Abt. (In- secta), 7. Buch, b) (Phthiraptera). 290 S. Akademische Ver- lagsgesellschaft Geest & Portig, Leipzig. Hopkins, G. H. E. 1949. The host-associations of the lice of mammals. Proceedings of the Zoological Society, London 119: 387-604. MARSHALL, I. K. 2003. A morphological phylogeny for four fam- ilies of amblyceran lice (Phthiraptera: Amblycera: Meno- ponidae, Boopiidae, Laemobothriidae, Ricinidae). Zoological Journal of the Linnean Society 138: 39-82. NELSON, B. C. 1972. A revision of the New World species of Rici- nus (Mallophaga) occuring on Passeriformes (Aves). Univer- sity of California Publications in Entomology 68: 1-175. 46 Götz RHEINWALD: Position of Trochiliphagus (Insecta: Phtiraptera, Ricinidae) PALMA, R. E. 1996. Ricinidae. Pp. 145-146 in: WELLS, A. (ed.) Zoological Catalogue of Australia, Vol. 26. Perez, J. M., GRANADOS, J. E. & Ruiz, I. 1995. The morpholo- gy of Laemobothrion (Laemobothrion) maximum (Phthi- raptera: Laemobothriidae). Parasitología 37: 45-51. PRICE, R. D., HELLENTHAL, R. A., PALMA, R. L., JOHNSON, K. P. & CLAYTON, D. H. 2003. The chewing lice: world checklist and biological overview. Illinois Natural History Survey. Spe- cial Publication 24. RHEINWALD, G. 1968. Die Mallophagengattung Ricinus DE GEER, 1778. Revision der außeramerikanischen Arten. Mitteilungen des Hamburger Zoologischen Museums und Instituts 65: 181-326. SIBLEY, C. G. & MONROE, B. L. 1990. Distribution and taxono- my of birds of the world. Yale University Press, New Haven & London. SYMMONS, S. 1952. Comparative anatomy of the Mallophagan head. Transactions of the Zoological Society of London XXVII, 4: 349-436. APPENDIX Investigated material The Natural History Museum, London Trochiliphagus hirsutus Carriker O” Paratype Trochiliphagus hirsutus Carriker Q Trochiloecetes rupununi Carriker © Trochiloecetes rupununi Carriker 2,0 Trochiloecetes insularis Carriker o Paratype Trochiloecetes insularis Carriker 3 larvae Trochiloecetes insularis Carriker Q, | larva Ricinus dolichocephalus (Scopoli) 3 Q Neoparatype Personal material Trochiloecetes ochroterenal? 19,10 1971, leg. Poley Trochiloecetes spec. 19 1971, leg. Poley Ricinus rubeculae (Schrank) 19,30 Ricinus fringillae De Geer 19 WEBB, J. E. 1946. Spiracle structure as a guide to the phyloge- netic relationships of the Anoplura (biting and sucking lice), with notes on the affinities of the mammalian hosts. Proceed- ings of the Zoological Society of London 116: 49-119. Author’s address: Goetz RHEINWALD, Schónblick 10, D-53562, St. Katharinen, Germany; E-Mail: goetz.rheinwald@t-online.de. Received: 02.2005 Revised: 06.10.2005 Accepted: 09.11.2005 Corresponding editor: B. J. Sinclair Glaucis hirsuta, Arıpo Valley, Trinidad, 14.1V.1960, TRVL. 4505, Brit. Mus. 1963-351 Glaucis hirsuta, Arıpo Valley, Trinidad, Aitken coll. Brit. Mus. 1963-351 Phaethornis superciliosus (AMH 36), Belize, Aguacate, 4.V.1979, C. Lyal and A. Hutson Phaethornis superciliosus, 42984, Mosqueiro Ferry, Benevides, Brazil, 25.X.1968, T. Aitken, ML.17, Brit. Mus. 1970-726 Glaucis hirsuta, Arıpo Valley, Trinidad, 5.VH. 1960, TRVL. 4622, Brit. Mus. 1963-351 Glaucis hirsuta, Aripo Valley, Trinidad, 5.VIL 1960, TRVL. 4622, Brit. Mus. 1963-351 Glaucis hirsuta, Aripo Valley, Trinidad, 17.111. 1961, T. Clay, No. 194, Brit. Mus. 1963-351 Oriolus. o. oriolus, N.E. Poland, VIII 1935, 4190, Brit. Mus. 1951-171 Selasphorus rufus, Nationalpark Desierto de los Leones, 30 km w. von Mexico City, VIII Eugenes fulgens, Nationalpark Desierto de los Leones, 30 km w. von Mexico City, VIII Luscinia svecica cyanecula, Modracek, Pakvice, CSR, 4.1V.1955, leg. F. Balat Fringilla coelebs, Obergurgl, Ötztal, Österreich, 1.1X.1968, leg. A. Aichhorn Bonner zoologische Beiträge | Band 55 (2006) Heft 1 Seiten 47-59 Bonn, Januar 2007 The Freshwater Mite Porolohmannella violacea (Kramer, 1879) (Acari: Halacaridae), Description of Juveniles and Females and Notes on Development and Distribution Ilse BARTSCH Hamburg, Germany Abstract. The description of female Porolohmannella violacea (Kramer, 1879) is supplemented and the development of external characters from larva to adult described. Beside an increase in size and number of setae from instar to instar, si- milar to that in other halacarid species, there 1s a marked difference in the growth in length of the palp and leg segments. Larva and protonymphs have epimeral pores. Data on geographical distribution are summarized. According to present records, P. violacea 1s restricted to the northern hemisphere, nonetheless, it is expected to be found in the south, too. Keywords. Freshwater halacarid, Porolohmannella, development, distribution. 1. INTRODUCTION Halacarid mites are primarily marine but a few species live almost exclusively in continental freshwater. KRAMER (1879) was the first to describe a halacarid from freshwa- ter, Leptognathus violacea, a rather large species charac- terized by a long, slightly curved rostrum and palps, in all strikingly similar to the marine genus Leptognathus Hodge, 1863, and a violet to pink integument, hence the name violacea (cf. KRAMER 1879). The species, described from Thuringia, Germany, turned out to be wide-spread in Europe, as documented by the large numbers of cita- tions (cf. K. Viets 1956; K. O. Viets 1978, 1987). The name Leptognathus proved to be pre-occupied and LOHMANN (1901) proposed a new name, Trouessartella Lohmann, 1901. This was pre-occupied, too, and there- upon TROUESSART (1901) introduced Lohmannella Troues- sart, 1901. In the following decades the number of both marine and freshwater halacarid species increased. Karl VieTs (1927, 1933) separated the freshwater species, all with external genital acetabula, from the marine species with internal acetabula, and distinguished between the families Hal- acaridae Murray, 1877 and Porohalacaridae Viets, 1933, the former including the marine taxa, the latter the fresh- water taxa. Accordingly, K. VieTS (1927, 1933) distin- guished between a marine genus Lohmannella, in the Lohmannellinae Viets, 1927, and a freshwater genus, for which he introduced the name Porolohmannella Viets, 1933, with the type species P. violacea (Kramer), and the subfamily Porolohmannellinae Viets, 1933. The division into Halacaridae and Porohalacaridae, sole- ly based on the position of the genital acetabula, external or internal, was abandoned by NEwELL (1947), BARTSCH (1973) and PETROVA (1974, 1981). In several species of the marine genera Acarothrix Bartsch, 1990, Halacarel- lus Viets, 1927, Halacaroides Bartsch, 1981, Isobactrus Newell, 1947, and Thalassarachna Packard, 1871 males have external but females internal acetabula (BARTSCH 2004, and in press). External genital acetabula are 1on-per- meable areas with osmoregulatory function (BARTSCH 1973; ALBERTI 1977, 1979). In marine halacarids one to three pairs of acetabula are present, in freshwater forms often more than three. The genital acetabula may be large, e.g., in the genera Halacarus Gosse, 1855 and Thalas- sarachna, or small, as in most species of Copidognathus Trouessart, 1888 and Simognathus Trouessart, 1889. Ina few marine species, unusual enlargement is documented. Many, though not all halacarıd mites living ın freshwater have external genital acetabula. Species of freshwater genera differ from marine ones both in the position of the genital acetabula and in their integu- ment. The plates are thin, they show no coarse sculptur- ing or raised areolae, no rosette pores; they often are rather uniformly ornamented, with a reticulate or foveate pattern, and have tiny evenly spread canaliculi cross the integu- mental layers. The position of the genital acetabula as well as the structure of the integument and ornamentation of the plates certainly are influenced by physical and chem- ical environmental features, as in other meiofaunal taxa, e.g., In ostracods (MEISCH 2000). A8 Ilse BARTSCH: The freshwater mite Porolohmannela violacea (Kramer, 1879) Figs 1-8. Porolohmannella violacea, larva, 1, ıdıosoma, dorsal; 2, idiosoma, ventral; 3, gnathosoma, ventral; 4, gnathosoma, late- ral; 5, leg I, medial; 6, leg II, medial; 7, leg MI, medial; 8, tarsus III, lateral (medial parambulacral seta and claw omitted, dorso- medial fossary seta in broken line). (ep, epimeral pore; Ifs, dorsolateral fossary seta; mfs, dorsomedial fossary seta) Scale = 50 um. Independent of the position of the genital acetabula, Porolohmannella and Lohmannella proved to differ in a number of other characters, in the number of setae on the epimeral plates and genitoanal plate, the number and arrangement of setae on the palps and gnathosoma, the shape of the pharyngeal plate, the position of the soleni- dion on tarsus II, and absence or presence of a carpite (cf. BARTSCH 1989). The state of these characters is expected to be independent of environmental parameters. Recently, in a small pond near Hamburg, Germany, nu- merous females and juveniles of Porolohmannella vio- lacea were found. The development of external charac- ters, from instar to instar, has been studied in several ma- rine species (BARTSCH 1998, 2003) but to date in no rep- resentative of a freshwater halacarid genus. The larva, nymphs and female of P violacea are now described, the character development outlined and compared with ma- rine halacarids. 2. MATERIAL AND METHODS The material studied was collected in June, July and Au- gust 2006 in northern Germany, north of Hamburg, in a shallow pond adjacent to a helocrene swamp area. The pond is approximately 10x20 m (width x length), on the northern border there are small bushes, mainly willows Bonner zoologische Beiträge 55 (2006) 49 (Salix sp.), rarely more than 3 m ın height, along the south- ern border most trees have been removed. In the shallow water there is a mass occurrence of the moss Fontinalis howellii Renauld & Cardot. Obvious macrofaunal elements in the pond are the snails Lymnaea stagnalis (Linnaeus, 1758) and Planorbarius corneus (Linnaeus, 1758). Amongst the meiofauna living between the moss mites were very abundant, viz., the ha- lacarids Porohalacarus alpinus Thor, 1910 and Porolohmannella violacea and the oribatid mite genus Hy- drozetes Berlese, 1902; Hydrachnidia and the halacarid mites Soldanellonyx monardi Walter, 1919 and Lobohal- acarus weberi (Viets & Romijn, 1924) were rare. Other meiofaunal taxa were rotatorians, turbellarians, nematodes, oligochaetes, tardigrades, and larvae of insects (cerato- pogonids and chironomids); copepods or ostracods were almost lacking. In January, February and March 2006 the mean monthly temperature was below | °C (DEUTSCHER WETTERDIENST 2006). June and July 2006 were characterized by high tem- perature and intense, long-term sunshine. July, with sev- eral days beyond 30 °C, had higher temperature and more sunshine than ever registered since regular documentation of meteorological data (which started a century ago). The sunshine period, with up to 16 hours a day, was 50 % be- yond long-term mean July values. In both June and July rain was considerably below average. August was ex- tremely wet, with two to four times more rain than usual (DEUTSCHER WETTERDIENST 2006). Porolohmannella violacea was extracted from moss sam- ples. Ten individuals of each instar (larva, protonymph, deutonymph, female) were studied in detail. Abbreviations used in the descriptions are: AD, anterior dorsal plate; AE, anterior epimeral plate; ds-1 to ds-6, first to sixth pair of dorsal setae of idiosoma, numbered from anterior back- ward; El to EIV, epimeral plate(s) I to IV; GA, genitoanal plate; GO, genital opening; GP, genital plate; OC, ocular plate(s); P-2 to P-4, second to fourth palpal segment; pas, parambulacral seta(e); PD, posterior dorsal plate; PE, pos- terior epimeral plate(s); pgs, perigenital setae. The epimeral plates, legs and their segments are numbered I to IV. The leg segments are trochanter, basifemur, telofe- mur, genu, tibia, and tarsus. The setation formula of the legs presents the number of setae from trochanter to tar- sus. The given number of setae of the tarsi includes the solenidion and parambulacral setae. The position of a se- ta is given in a decimal system with reference to the length of the relevant structure, from its anterior to posterior or proximal to distal end. Voucher specimens of each instar are deposited in the Zoological Museum in Hamburg. 3. DESCRIPTION 3. 1. Larva (Figs 1-8) Idiosoma. Length 207-277 um. Idiosoma pale, almost transparent, with pair of small spots of black eye-pigment. Dorsal plates with delicately reticulate ornamentation, each polygon about 5-7 um in diameter. Anterior margin of AD truncate (Fig. 1). OC rounded, slightly wider than long. PD 1.2 times longer than AD. Idiosoma with four pairs of gland pores, one pair on AD, one on OC, and two pairs on PD. Pore canaliculus immediately posterior to gland pore of OC. Dorsal setae tiny. Pair of ds-1 on AD, ds-2 on OC, ds-3 in striated integument, ds-4 and ds-5 on PD, both anterior to the level of third pair of gland pores, ds-6 in posterior margin of PD. Part of anal sclerites seen in dorsal aspect. AE with pair of epimeral pores (Fig. 2), each slightly ovate and 4-5 um in diameter, and two pairs of long, slender ventral setae. PE short, epimeral plate IV lacking; PE with single pair of slender setae. Anus in pos- terior end of idiosoma. Gnathosoma. Length 117-123 um, 0.44-0.50 of length of idiosoma. Rostrum slender, about 1.5 times longer than gnathosomal base. Gnathosomal base faintly ornamented. Pharyngeal plate large, extending almost to basal margin (Fig. 3). Tectum slightly concave. Basal pair of maxillary setae long, inserted near base of rostrum, apical pair of maxillary setae near tip of rostrum, adjacent to two pairs of rostral setae. Palps four-segmented. P-2 cylindrical, with conspicuously long seta at 0.68 or about 18 um prox- imal to end of segment; apically with a ventromedial lamellar process (Fig. 4). P-3 with dorsomedial lamellar process. P-4 with three slender setae in a basal whorl, a short spine, two stout setae, about 7 um long, and one se- tula. Chelicerae elongate; small apical claw dentate. Legs. With three pairs of five-segmented legs; basi- and telofemora fused. Number of setae per segment: legs I and II, 1, 1+3 (basi- + telofemur), 4, 5, 6; leg III, 1, 1+2, 2, 5, 6. No variation in the 10 larvae studied. Tibiae I to HI 1.2, 1.1 and 1.1 times longer than tarsi. Pair of ventral setae on genu and tibia I pectinate (Fig. 5). Genu II with sin- gle pectinate seta and tibiae II and III each with pair of ventral setae (Figs 6 and 7). All tarsi with pair of small fossa membranes. Tarsus I with dorsolateral papilla prox- imal to dorsal fossary seta. Solenidion on tarsus I and Il in dorsolateral position immediately adjacent to fossa membrane, about 6 tm long. Dorsolateral fossary seta on tarsus I distinctly longer than dorsomedial seta. On tarsi II and HI no such difference in length of fossary setae. On tarsus II interval between basal seta and dorsal fossary seta about 20 um, distance from basal seta to basis of tar- sus somewhat more than height of tarsus; basal seta at 0.27 relative to length tarsus (Fig. 8). All tarsı end with pair of 50 Ilse BARTSCH: The freshwater mite Porolohmannela violacea (Kramer, 1879) 10 «MD Figs 9-16. Porolohmannella violacea, protonymph, 9, idiosoma, dorsal; 10, idiosoma, ventral; 11, leg 1, medial; 12, leg II, me- dial; 13, leg MI, medial; 14, leg IV, medial; 15, tarsus III, lateral (medial parambulacral seta and claw omitted, dorsomedial fossa- ry seta in broken line); 16, palp, lateral. (ep, epimeral pore) Scale = 50 um. parambulacral setae. Tarsı with paired claws. Claw pecten with short tines. Central sclerite minute, on tarsus III with delicate upper tooth. 3. 2. Protonymph (Figs 9-16) Idiosoma. Length 301-347 um. Idiosoma pale, translu- cent; gut contents with red spots. Idiosoma with two lat- eral and one median spot of eye pigment. Dorsal plates delicately reticulate. Anterior margin of AD truncate (Fig. 9) or with minute frontal process. OC longer than wide. PD 1.3 times longer than AD. Pair of ds-3 on OC. AE with pair of small epimeral pores (Fig. 10), 34 um in diame- ter, and three pairs of slender setae. PE including epime- ral plates III and IV; a single ventral (marginal) seta pres- ent. Genital plate and anal plate fused to GA; this plate with one pair of external genital acetabula which are 4 um in diameter. Gnathosoma. Length 145-161 um, 1.e., 0.43-0.49 of length of idiosoma. Seta on P-2 at 0.81 (Fig. 16); distance from seta to end of P-2 15 um. Legs. With four pairs of legs, legs I to III six-segmented (Figs 11-13), leg IV five-segmented (Fig. 14). Leg chaeto- taxy: legs and IL 1,23, 5, 3, 6; leg IM, 1,2,35 23:6; Bonner zoologische Beiträge 55 (2006) Sl Figs 17-24. Porolohmannella violacea, deutonymph, 17, idiosoma, dorsal; 18, idiosoma, ventral; 19, palp, lateral; 20, leg I, me- dial; 21, leg II, medial; 22, leg MI, medial; 23, leg IV, medial; 24, tarsus III, lateral (medial parambulacral seta and claw omitted, dorsomedial fossary seta in broken line). Scale = 50 um. leg IV, 0, 0+2 (basi- + telofemur), 3, 5, 5. Tibiae I to IV 1.5, 1.3, 1.2 and 1.1 times longer than tarsi; telofemora I to III 1.1-1.2 times longer than genua. Genu and tibia I each with pair of bipectinate setae (Fig. 11). Genu II with single pectinate seta. Tibiae II to IV each with pair of ven- tral setae; on tibia III ventromedial seta coarsely pectinate and somewhat shorter than ventrolateral seta; on tibia IV both setae similar in size and not markedly pectinate. Paired fossary setae on tarsus I dissimilar in length, on tar- si II to IV almost equal in length. Basal seta on tarsus II] at 0.38 relative to length of tarsus (Fig. 15), or 1.5 times height of tarsus; interval between basal seta and dorsal fos- sary seta 20 um. 3. 3. Deutonymph (Figs 17-24) Idiosoma. Length 440-487 um. Idiosoma with three spots of eye pigment. Dorsal and ventral plates reticulate. AD with small frontal spine (Fig. 17); AD about as long as wide. OC with cornea, two pairs of minute setae, gland pore and pore canaliculus. Length of PD about 1.4 times that of AD. AE with three pairs of ventral setae; epime- ral pores lacking. PE with short dorsal seta and two long ventral setae. GA with two pairs of setae, situated ante- rior to two pairs of genital acetabula (Fig. 18). Acetabu- la 3-4 um in diameter. Gnathosoma. Length 195-204 um, 1.e., 0.41-0.45 of length of idiosoma. Seta on P-2 at 0.85 (Fig. 19), or 15 um removed from apical end. 59 Ilse BARTSCH: The freshwater mite Porolohmannela violacea (Kramer, 1879) Figs 25-32. Porolohmannella violacea, female, 25, idiosoma, dorsal; 26, idiosoma, ventral; 27, genital sclerites; 28, ovipositor with genital spines; 29, gnathosomal base, dorsal; 30, gnathosoma, ventral; 31, palp, lateral; 32, gnathosoma, lateral. (gae, exter- nal genital acetabula; gai, internal genital acetabulum) Scale = 50 um. Legs. All four pairs of legs six-segmented. Tibiae of legs I to IV 1.7, 1.5, 1.4 and 1.3 times longer than correspon- ding tarsi. Telofemora of all legs about 1.1 times length of genua. Leg chaetotaxy as summarized in Table 1. Gen- ua I and II with two and one pectinate setae, respective- ly. Tibia I in general with four ventral pectinate setae (Fig. 20), rarely with three such setae and one of mid-segmen- tal setae being absent. Tibia II mostly with three, rarely with four ventral setae (Fig. 21). Tibia III with four ven- tral setae, two of them short and pectinate, one seta slen- der and smooth and one rather stout but without distinct pectination (Fig. 22). Tibia IV with single pair of gener- ally long ventral setae (Fig. 23). On tarsi I and II dorso- lateral fossary seta longer than dorsomedial one, on fol- lowing tarsi these fossary setae similar in size. On tarsus III distance between basal seta and dorsal fossary seta 20 um; this basal seta at 0.46 relative to length of tarsus (Fig. 24); distance between basal seta and basal edge of tarsus slightly more than height of tarsus. 3. 4. Female (Figs 25-39) Idiosoma. Length 525-590 um. Integument of idiosoma, gnathosoma and legs pink to violet, gut contents red- brown. With three spots of eye pigment, a pair on OC and a single one on AD. Dorsal and ventral plates distinctly reticulate, mesh size 5-8 um. AD slightly wider than long (Fig. 25). OC longer than wide. Length of PD about 1.7 times that of AD. AE wider than long, with three pairs of ventral setae. PE with short dorsal seta and two long ven- Bonner zoologische Beitráge 55 (2006) 53 Figs 33-39. Porolohmannella violacea, female, 33, leg I, medial; 34, leg II, medial; 35, leg IH, medial; 36, leg IV, medial; 37, tarsus I, lateral (medial parambulacral seta and dorsomedial fossary seta in broken line); 38, tarsus II, lateral (medial parambula- cral seta, claw and dorsomedial fossary seta in broken line); 39, end of tibia and tarsus III, dorsolateral. (p, papilla) Scale = 50 um. tral setae (Figs. 25 and 26). GA large, dominating ventral aspect; plate with three pairs of slender perigenital setae, two pairs anterior to GO, one pair lateral to GO. GO al- most in middle of GA. Genital sclerites with two pairs of external genital acetabula, each acetabulum about 5 um in diameter. A third pair of similar-sized genital acetabu- la in internal position, on inner flank of genital sclerites (Fig. 27). Ovipositor short, with basal pair of bristle-like setae and five pairs of short apical spines (Fig. 28). Basai pair 15 um long and delicately pennate. Apical pairs of genital spines 7-9 um long and tri- or quadrifid; two an- terior pairs somewhat longer (9 um) than three posterior pairs of apical genital spines. Anus in ventral position. Fe- males with zero to three eggs. Gnathosoma. Length 252-275 um, 0.45—0.48 of length of idiosoma. Gnathosomal base coarsely foveate (Fig. 30). Arrangement of maxillary setae as in larva (Fig. 30). Tec- tum concave (Fig. 29). P-2 and P-3 with ventromedial and dorsomedial spiniform process, respectively (Fig. 32). Se- ta on P-2 at 0.86 (Figs 31 and 32), 20 um proximal to end of that segment. Legs. Tibiae of legs I to IV 1.9, 1.6, 1.5 and 1.3 times longer than their tarsi; telofemora 1.1-1.2 times longer than genua. Chaetotaxy as summarized in Table 2. Genu and tibia I with one and two pairs of ventral setae, respec- tively, ventromedial setae distinctly pectinate (Fig. 33), ventrolateral setae weakly pectinate. Genu and tibia II with one and three to four pectinate setae, respectively (Fig. 34). 54 Ilse BARTSCH: The freshwater mite Porolohmannela violacea (Kramer, 1879) Fig. 40. Geographical distribution of Porolohmannella violacea. Table 1. Porolohmannella violacea, deutonymph. Leg segments and their number of setae (number of cases of a given variant in brackets). segment 1 2 3 4 5 6 leg I 1(20) 2(20) 3(2),4(17),501) 6(20) 7(7),8(13) 6(20) leg II 1(20) 2(20) 3(1),4(19) 6(20) 7(18),8(2) 6(20) leg HI 1(20) 2(20) 3(20) 3(20) 7(20) 6(20) leg IV 0(20) 2(20) 3(20) 3(20) 5(20) 5(20) Table 2. Porolohmannella violacea, female. Leg segments and their number of setae (number of cases ofa given variant in brack- ets). segment 1 2 3 4 5 6 leg I 1(20) 2(20) 4(20) 6(20) 7(1),8(19) 6(20) leg II 1(20) 2(20) 4(20) 6(20) 7(3),8(17) 6(20) leg MI 1(20) 1(1),2(17),3(2) 3(20) 3(20) 7(20) 6(20) leg IV 0(19),1(1) 2(20) 2(1),3(19) 3(20) 5(3),6(12),7(5) 5(20) Bonner zoologische Beiträge 55 (2006) 55 Table 3. Porolohmannella violacea. Number of idiosomal setae, epimeral pores, epimeral plates, and genital acetabula (internal and external) in each instar. (—, GP absent in larva. Changes from one instar to the other underlined and in bold). L AE, pairs of setae 2 AE, pairs of epimeral pores | PE including epimeral plates ENI PE, number of setae 1 GP, pairs of setae - GP, pairs of acetabula - Table 4. - PN DN Ad 3 3 3 l 0 0 EMT EV EIII,EIV EII,EIV | 3 3 0 2 3 l 2 3 Porolohmannella violacea. Leg chaetotaxy of larva\protonyph\deutonymph\female. (-, leg IV absent ın larva. Single cases omitted, rare variants in parentheses. Changes from one instar to the other underlined and in bold). leg I L\PN\DN\Ad trochanter I\I\I\ basifemur 1\2\2\2 telofemur 3\3\(3-)4\4 genu 4\5\6\6 tibia 5\5\7-8\8 tarsus 6\6\6\6 Tibia III with four ventral setae, ventromedial setae short and pectinate (Fig. 35). Tibia IV with three dorsal and two to four ventral setae (Fig. 36); pair of apical ventral setae always present; number of mid-segmental setae, one pecti- nate, one slender, variable, often different on right and left leg. Solenidia on tarsi I and II (Figs 37 and 38) in dorso- lateral position, adjacent to fossa membrane, 10 and 9 um in length, respectively. On these tarsi dorsomedial fossary seta distinctly shorter than dorsolateral seta. Tarsus I with papilliform process proximal to dorsal fossary seta (Fig. 37). Basal seta of tarsus III almost in middle of segment, at 0.57 relative to length of tarsus (Fig. 39), interval be- tween basal seta and base of tarsus 1.3 times that of height of tarsus, distance from basal seta to dorsal fossary seta 20-22 um. All tarsi with pair of parambulacral setae and pair of claws. Male unknown. Remarks. In the females, the number and shape of the ventral setae on tibia IV proved to vary from two to four. The apical pair of setae always is present, the two mid- leg II leg III leg IV L\PN\DN\Ad L\PN\DN\Ad L\PN\DN\Ad IMM IMM -\0\0\0 1\2\2\2 1\2\2\2(-3) -\0\2\2 3\3\4\4 2\3\3\3 -\2\3\3 4\5\6\6 2\2\3\3 -\3\3\3 5\5\7(-8)\(7-)8 5\5\7\7 -\S\5\(5-)6(-7) 6\6\6\6 6\6\6\6 -\5\5\5 segmental setae, one bipectinate, one slender, may be lack- ing on either one or both legs. Unilateral differences are very common. In ten females studied the most common combination of setae on tibia IV was, presence of the slen- der ventral mid-segmental seta but absence of the pecti- nate seta (11 legs, 55 %). On five tibiae IV (25 %) both a pectinate and a slender mid-segmental seta were pres- ent, whereas on three tibiae (15 %) both mid-segmental setae were absent. A single tibia IV (5 %) had a pectinate but no slender mid-segmental seta. In material (53 fe- males) from North America (unpublished data), the pres- ence of three ventral setae on tibia IV was the most com- mon state (63 %), without any concentration on the one or other variant in a given locality (population). The four females collected on Hokkaido, Japan, had three ventral setae on tibiae IV, all pectinate (ABE 1990). All instars have a small dorsolateral papilla on tarsus I (cf. Fig. 37), although in dorsomedial aspect the papilla may be obscured. ABÉ (1990) emphasized the similarity with a chemosensory organ. There is a very delicate afferent canal through the integument, so this papilla is not mere- 56 Ilse BARTSCH: The freshwater mite Porolohmannela violacea (Kramer, 1879) ly a part of the reticulum that covers the surface; it may represent a famulus. 3. 5. Development The larva has a short idiosoma with three pairs of five- segmented legs. The dorsal plates AD, OC and PD are present, the PD is short (relative to length of the AD), the ornamentation of the plates is weak. The larva already has the final number of dorsal idiosomatic setae and gland pores. The AE has a pair of epimeral pores and two pairs of setae. The PE is short and bears a single seta. The anal sclerites are partly visible in dorsal aspect. The gnatho- soma is characterized by its slender palps and rostrum. In the protonymphs the epimeral plate IV and leg IV have been added. The OC and PD have increased in size, the OC in the medial and posterior portion, the PD both an- terior to the pair of ds-4 and between the gland pores. The PE that now includes epimeral plate IV is much longer than in the larva but the PE still bears a single seta. On the AE a pair of setae on epimeral plate II has been added. The epimeral pores are still present though less conspic- uous than in the larva. A genital plate, fused with the anal plate, and a pair of genital acetabula have been added. The shape of the gnathosoma resembles that of the larva but the position of the seta on P-2 has changed in this and the following instars. The femora of the three larval pairs of legs are divided, hence the legs are six-segmented, leg IV is five-segmented. The tibiae have increased in length, re- lative to the tarsi. Setae have been added to the basifemo- ra I to III, telofemur HI and genua I and II. The deutonymph differs from the protonymph in that the epimeral pores of the AE are reduced, on the PE a short dorsal and long ventral seta are added and the GA bears two pairs of genital acetabula and two pairs of pgs. Both pairs of setae are anterior to the genital acetabula. All legs are six-segmented. Changes in the number of setae are summarized in Table. 4. Setae have been added on basife- mur IV, telofemora I, II and IV, and genua and tibiae I to IM. In the female both the dorsal and ventral plates are longer and wider and their reticulate ornamentation is much more prominent than in the nymphs, and the areas of striated integument between the plates are narrow. On the large GA a posterior pair of pgs has been added and in the mid- dle of the plate is the large genital opening. Two pairs of genital acetabula have moved onto the genital sclerites, a third pair ıs on the inner flank of the sclerites. The anal sclerites are ın a ventral position. The shape, number of setae and the ratio of gnathosoma to idiosoma is similar to that of the other instars, P-2 has grown in length and the seta, in almost mid-segmental position in the larva, is now in an apical position. All legs, especially their tibi- ae, have increased in length; tibia I is 1.7 times longer than tarsus I (instead of 1.1-1.2 times as in the larva). The tar- si are more slender (length:height ratio) than in the larva, extended in length primarily at their bases, as demonstrat- ed on tarsus Ill. The basal seta of tarsus III is positioned at 0.56 in the female, but at 0.46, 0.38 and 0.27 in the pre- ceding instars, respectively. The setation of the tibiae of- ten differs from that in the deutonymph, that of the other leg segments is the same in both instars. The number of setae on trochanters and tarsi is unchanged throughout all instars. 4. BIOLOGY AND DISTRIBUTION The majority of records of Porolohmannella violacea are from standing surface waters, swamps, ponds and lakes, though the species also has been extracted from the groundwater (GLEDHILL, 1973, 1982; STRAYER, 1988). Porolohmannella violacea inhabits altitudes from the sea level (coastal waters) to mountain lakes in almost 2000 m, e.g., the Lacs de Estibere in the Pyrenees (ANGELIER 1965) and the Statzer See in the Alps (WALTER 1922), and a variety of substrata, sand, flocculent ooze, vascular plants, mosses, algae and also gill chambers of crayfish. Details on the life cycle are not known, other than that fe- males are ovigerous most time of the year, carrying one to six eggs each. The presence of a large number of lar- vae in spring 2006 (beginning of June), as well as deu- tonymphs and females, and the appearance of protonymphs in the sample from July, seems to be an ev- idence for intense reproduction in spring. All present records are from the northern hemisphere (Fig. 40), from Europe, Greenland, North America, and Japan (K. VieTs 1956; K.O. ViETS 1978, 1987; GREEN & MAC- Quitty, 1987; Pex1C 2004; BARTSCH 2006). The data sum- marized in Fig. 40 include unpublished records from Spain, Portugal and Canada (Northwest Territories, British Columbia, Alberta, Manitoba, Ontario, Quebec, Newfoundland). Several of the collecting sites (e.g., in Greenland and Northwest Territories (lake near Tuktuyak- tuk)) are ice-covered for most of the year. In contrast, the small pond near Hamburg, hardly shaded by trees, was ex- posed for several weeks to unusual long-term solar radi- ation, high temperatures and deteriorating oxygen supply. The pond was rapidly evaporating in July. Beneath a cov- er of dried remnants of a former vegetation, in a moist mixture of sediment, debris and mosses, specimens of P. violacea still were alive. At a visit to the pond at the end of August, after repeatedly heavy rainfall and distinctly lowered temperature, the moss was found to be regener- Bonner zoologische Beiträge 55 (2006) 57 ating. In the sample examined Porohalacarus alpinus and Hydrozetes sp. were still present in moderate numbers, adults as well as juveniles. The number of specimens of Porolohmannella violacea had decreased considerably, but still, larvae, nymphs and females were found. 5. DISCUSSION In several descriptions of the freshwater halacarids Lobo- halacarus weberi, Porohalacarus alpinus, Soldanellonyx monardi and S. chappuisi Walter, 1917, the external char- acters of the one or other juvenile have been briefly out- lined (WALTER 1919b; WALTER & BADER 1952; BARTSCH 1973, 1975, 1982); unfortunately some species are de- scribed exclusively on the basis of nymphs (WALTER 1919a; VIETS 1931; JANKOVSKAJA 1967). Now, the devel- opment of external characters from larva to adult of Porolohmannella has been examined. The marine genera best studied in respect to their development are Copido- gnathus, Isobactrus, Rhombognathides Viets, 1927, Metarhombognathus Newell, 1947 and Rhombognathus Trouessart, 1888 (BARTSCH 1998, 2003). In contrast to many other aquatic mite taxa, e.g., Hydrach- nidia and Oribatida, the juvenile instars of halacarıd mites do not differ markedly from adults, though of course they are of smaller size. Halacarids have no apparent change in habitat or life-style. Freshwater halacarids progress through one larval and two nymphal instars until they moult into the adult. A single exception, Astacopsiphagus parasiticus Viets, 1931, a mite found on freshwater crayfish in Australia, has three nymphal instars in its life cycle (VieTs 1931). Marine hal- acarids have one to three nymphal instars. The character development in P. violacea is similar to that known in other halacarid species (cf. BARTSCH 1998, 2003). Minor differences are: (a) In P. violacea epimeral pores are present in the larva but also in the protonymph; they are absent in the deu- tonymph and female. In most marine species in which adults lack epimeral pores, these pores are restricted to the larva. (b) There is a striking change in the length ratio of tibia to tarsus, with a marked growth of the tibiae compared with that of the tarsi. Such a difference in length has not been described before. Adults of P violacea are long- legged, but marine species studied so far are short-legged. Long-legged marine species may show a similar shift in the length relation. (c) From instar to instar, the basal seta on tarsus III and the seta on P-2 are moved to a more apical position. The segments obviously grow in length near their bases. In ma- rine genera, a similar but much less marked growth near the base of the tarsi has been documented. The marine species studied to date are short-legged whereas the nymphs and females of P. violacea have long and slender legs and palps. Karl Viets (1927, 1933) once distinguished between hal- acarid species of marine and freshwater, between Halacari- dae and Porohalacaridae, respectively. That distinction has been abandoned (NEWELL 1947, 1959; BARTSCH 1973, 1989; PETROVA 1974, 1981) because it was not support- ed by synapomorphies. The ontogeny of Porolohmannel- la violacea is similar to the development of characters in marine halacarid species, and hence does not support a tax- onomic separation into marine and freshwater families. According to Fig. 40, P. violacea is wide-spread through- out the northern hemisphere. These data do not imply that P. violacea is restricted to the holarctic. Porolohmannel- la violacea ıs euryvalent and thrives in a variety of habi- tats, in standing as well as in running water, in brackish coastal as well as humic inland waters, 1t can withstand long-term solar radiation with rising water temperature, but also ice-covering for more than six months. One may speculate that high temperature prevents a colonization of the tropics, but the P. violacea population in northern Ger- many proved to withstand the hot summer in 2006. Hence, the temperature in the tropics should not hamper a distri- bution. According to TESCHNER (1963) and SIEMER (1996), in the freshwater halacarıd Lobohalacarus weberi and the marine Metarhombognathus armatus (Lohmann, 1893) a cold season 1s needed in the course of develop- ment. Does the higher and more uniform annual temper- ature cycle in warm-temperate and tropic areas prevent P. violacea from colonization? Presumably not. Lobohal- acarus weberi has been collected in the southern hemi- sphere, in Australia (Queensland, near Brisbane) (unpub- lished), in an area with an annual surface water tempera- ture range between 16 °C and 29 °C (Davie 2004). Sim- ilarly, P. violacea may thrive in the tropics and southern hemisphere. The present geographic distribution, restrict- ed to the holarctic, is merely the result of the extraordi- narily low number of samples taken in the south compared to that in the north. Porolohmannella violacea may be Mesozoic, or even Pre-Mesozoic, in origin (BARTSCH 1996). The species is expected to be found in South Amer- ica, Africa and Australia, either being a remnant of a Pan- gaean fauna or introduced by means of birds, mammals, insects, extreme floods and storms. 58 Ilse BARTSCH: The freshwater mite Porolohmannela violacea (Kramer, 1879) Acknowledgements. Dr. lan M. Smith, Ottawa, collected hal- acarid mites in Canada and the United States over a period of three decades and sent me the mites for study. The data on Porolohmannella violacea from Canada are from these collec- tions, deposited in the Canadian National Collection, Ottawa. Thanks are due to I. M. Smith for forwarding that halacarid ma- terial and two referees for helpful comments on the manuscript. REFERENCES ABÉ, H. 1990. New record of Porolohmannella violacea (KRA- MER, 1879) (Acari, Halacaridae). Acarologia 31: 241-246. ALBERTI, G. 1977. Zur Feinstruktur und Funktion der Genital- napfe von Hydrodroma descipiens (Hydrachnellae, Acari). Zoomorphologie 87: 155-164. ALBERTI G. 1979. Fine structure and probable function of gen- ital papillae and Claparede organs of Actinotrichida. Pp. 501-507 in: RODRIGUEZ, J. (ed.) Recent Advances in Acarol- ogy. II. Academic Press, New York. ANGELIER, E 1965. Les Porohalacaridae de la faune francaise. Annales de Limnologie 1: 213-220. BARTSCH, I. 1973. 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Die Halacaridae der Nordsee. Zeitschrift für wis- senschaftliche Zoologie 130: 83-173. VieTs, K. 1931. Uber eine an Krebskiemen parasitierende Hal- acaride aus Australien. Zoologischer Anzeiger 96: 115-120. VIETS, K. 1933. Vierte Mitteilung über Wassermilben aus unter- irdischen Gewássern (Hydrachnellae et Halacaridae, Acari). Zoologischer Anzeiger 102: 277-288. VIETS, K. 1956. Die Milben des Süßwassers und des Meeres. Ka- talog der Halacaridae. Meeresmilben. II. Abschnitt. Pp 641— | Bonner zoologische Beitráge 55 (2006) 870 in: VIETS, K. (ed.) Die Milben des Süßwassers und des Meeres. Hydrachnellae et Halacaridae, Acari. VEB Fischer Verlag, Jena. VIETS, K. O. 1978. Hydracarina. Pp. 154-181 in: ILLıes, J. (ed.) Limnofauna Europaea. Eine Zusammenstellung aller die eu- ropäischen Binnengewässer bewohnenden mehrzelligen Tier- arten mit Angaben über ihre Verbreitung und Ökologie. 2te Auflage, Gustav Fischer Verlag, Stuttgart. VIETS, K. O. 1987. II. Halacaridae des Süßwasser. Pp. 829-857 in: VIETS, K. O. (ed.) Die Milben des Süßwassers (Hydrach- nellae und Halacaridae [part.], Acari). 2: Katalog. Sonderbán- de des naturwissenschaftlichen Vereins zu Hamburg 8. Paul Parey, Hamburg. WALTER, C. 1919a. Hydracarınen aus den peruanischen Anden und aus Brasilien. Revue Suisse de Zoologie 27: 19-59. WALTER, C. 1919b. Schweizerische Süsswasserformen der Hal- acariden. II. Revue Suisse de Zoologie 27: 236-242. 59 WALTER, C. 1922. Die Hydracarinen der Alpengewásser. Denk- schriften der schweizerischen naturforschenden Gesellschaft 58: 61-252. WALTER, C. & BADER, C. 1952. Mission scientifique de l'Omo — Hydracarina. Mémoires du Muséum nationale histoire na- turelle, Serie A, 4: 87-236. Author’s address: Dr. Ilse BARTSCH, Deutsches Zentrum für marine Biodiversitätsforschung, Forschungsinstitut Senckenberg, Notkestr. 85, 22607 Hamburg, Germany; E- Mail: bartsch@meeresforschung.de. Received: 22.09.2006 Revised: 04.12.2006 Accepted: 12.12.2006 Corresponding editor: B. A. Huber ae Bonner zoologische Beitráge Band 55 (2006) Heft 1 Seiten 61-71 Bonn, Januar 2007 Patterns of Geographic Variation in Body Measures and Plumage Colour of the Brimstone Canary Crithagra sulphurata (Aves: Fringilidae) Sverre KLEMP Hamburg, Germany Abstract. The Brimstone Canary is a geographical highly variable species, distributed throughout southern and eastern Africa. Here I present data on body measures and plumage characteristics of 476 skins from all parts of the species’ ran- ge (Fig. 1). In both univariate and multivariate analyses the wing and beak measures showed the most remarked diffe- rences between populations (Tab. 2, 5). Variation parallels climatic trends with individuals from hot and humid regions have shorter wings and smaller beaks (Tab. 7). In contrast to earlier studies plumage colour should not be used to distin- guish populations, because of high individual variation (Tab. 8). Only skins from South Africa (including Natal and Zululand) could be separated clearly (stronger beaks) from all other populations (shorter wings, less strong beaks). Clinal variation and high individual variation did not allow more diffe- rentiation in the northern part of the range. Therefore, the data are in agreement with the separation of the subspecies sulphurata (in South Africa) and sharpei (all other populations), while wilsoni and all other subspecies are not suppor- ted. Keywords. Multivariate analysis, subspecies, wing dimensions, beak size, individual variation. 1. INTRODUCTION The traditional way to describe spatial variation of species is the recognition of subspecies. Subspecies are more or less clearly separated from other populations of the same species by differences in at least one trait and are distrib- uted in a more or less clearly defined subarea of the species” range. This method gives, however, an incomplete picture of variation, because many traits vary in clines and the spatial characteristics of clines may differ from trait to trait (see GOULD & JOHNSTON 1972 and ZNK & REM- SEN 1986 for reviews). Hence, to distinguish a representative of a subspecies from an individual variant or a local form in a cline profound knowledge of individual and geographical variation within the species is necessary. Such knowledge was not available when most subspecies have been described and is still not available for many species. The Brimstone Canary Crithagra sulphurata ıs such a ge- ographical highly variable species. Within the Canaries five afrotropical species form the monophyletic genus Crithagra as suggested by morphological, ethological and molecular data (van den ELZEN 2000). C. sulphurata in- habits grasslands and savannas with scattered trees throughout southern and eastern Africa (Fig.1). Within this range and especially in its eastern part a high degree of morphological variation has led to the description of sev- en subspecies: — sulphurata (Linnaeus, 1766) (LINNAEUS 1766, Syst. Nat., ed. 12,1: 305), type from Cape of Good Hope, — sharpii (Neumann, 1900) (NEUMANN 1900, J. Ornithol. 48: 287), tt. Marangu, Kilimanjaro, — shelleyi (Neumann, 1903) (NEUMANN 1903, Ornithol. Mber. 11: 184), t.t. Kafuro, NW Tanzania, — frommi (Kothe, 1911) (KoTHE 1911, Ornithol. Mber. 19: 71), t.t. Namanjera, Ufipa, SW Tanzania, — loveridgei (van Someren, 1921) (van SOMEREN 1921, Bull. Br. Ornithol. Club 41: 114), Lumbo, northern Mozambique, — wilsoni Roberts, 1936 (ROBERTS 1936, Ann. Transv. Mus. 18: 216), t.t. Kloof, Natal, and — languens Clancey, 1962 (CLANCEY 1962, Durban Mus. Nov. 6: 193), type from Manhica, Sul do Sava, south- ern Mozambique. Different authors have adopted controversial positions which of these types represent real subspecies, e.g., FRY & KEITH (2004) in their recent review recognized only three of these (sulphurata, sharpei, wilsoni). RAND (1968) and CLANCEY (1972) reviewed available measure- ments and distribution data and concluded, in accordance, that most of the “subspecies” mentioned above are not 62 Sverre KLEMP: Patterns of geographic variation of Brimstone Canary Crithagra sulphurata sharply separated from each other in the traits considered. Due to larger body size and stronger beaks South African nominate sulphurata can be distinguished from all other forms with relative ease. Therefore, RAND accepted only two subspecies: sulphurata and sharpei (all populations except South Africa). In an alternative view he divided sharpei on the basis of subtle differences in plumage colour and size into four subspecies: wilsoni (Natal, south- ern Mocambique), sharpei (Kenya), frommi (Tanzania, Zambia and Angola) and shelleyi (all other populations). In contrast, CLANCEY (1972) separated four subspecies in South Africa alone (sulphurata, wilsoni, languens and shelleyi). Both studies are, however, based on only a few traits and from some regions only a small number of in- dividuals have been included. In this study I present data on a larger variety of meas- ures and plumage characteristics of C. sulphurata skins from all parts of the species’ range. The aim of this study is to describe the extent and pattern of geographic varia- tion in this species. Which of the subspecies described ear- lier represent real taxonomic units and which are interme- diate stages arbitrarily picked out of a continuous cline of variation? In a second step I want to relate morphologi- cal variation to possible causal factors such as ecological conditions (climate, vegetation, topography) and palaeo- graphic events. 2. MATERIAL AND METHODS I measured a total of 485 museum skins of Crithagra sul- phurata, of which all 476 adults (253 males, 171 females and 52 unsexed) were considered in the following analy- ses. Additionally, I studied 24 skins of the sister species (van den ELZEN 2000), Crithagra flaviventris, for compar- ison (15 males, 9 females). The skins are kept by Zoolo- gisches Forschungsmuseum Koenig (ZFMK), Bonn, Zo- ologisches Institut und Museum (ZMH), Hamburg, Zool- ogisk Museum (ZMUC), Copenhagen, Zoologische Staatssammlung (ZSM), Munich, Natural History Muse- um (BMNH), Bird Group, Tring, and Royal Museum of Central Africa (RMCA), Tervuren, respectively. Tarsus and beak measures (length, width and height in both cases) were taken from all specimens. Length of wing, tail and first to ninth primaries as well as Kipp’s dis- tance (wing tip to first secondary) and graduation of tail were measured in all except molting specimens. From the wing measures I calculated Wing Pointedness Index I and II after MLIKOVSKY (1978) and HEDENSTROM (1989), re- spectively. Plumage colours (throat, breast, belly and back) were compared with standardized colour maps (Scandi- navian Colour Institute 2001). For statistical analysis I de- scribed standard colours by four variables: proportion of black, of colour, green (negatively correlated to propor- tion of yellow due to construction of the colour system), and red (C. flaviventris only). Not all traits could be tak- en for all individuals because of skin condition. Therefore, sample sizes differ from analysis to analysis. Skins with geographical origin close to each other have been pooled for analysis, resulting in 15 ‘populations’ (Fig. 1). Individuals which locality information were lacking, unclear or doubtful (e.g. Collection Meinertzhagen; cf. Knox 1993) were included in the analyses, but considered as belonging to none of the populations. To get larger sam- ple sizes some of the populations were pooled to larger “population groups’ (A-G) in some analyses (Fig. 1). F G % 2426 = a 4 x KX x x Fig. 1. Map of collection localities of skins (crosses - single skins; numbers - series with number of collected individuals). Ellipses define “populations”. “Population groups” are marked by letters A to G. The present breeding distribution of C. sulp- hurata (simplified after Fry & KEITH 2004) is shaded. Predominant vegetation structure was revealed for each skin locality using the vegetation map of EC-JRC (2003, map 2) summarizing all categories under ‘dominant tree layer’ as ‘forest’ and all categories under ‘dominant agri- culture’ and ‘dominant shrub or grass layer’ as ‘open habi- tat’. Climatic data of the closest weather station were tak- en from LEBEDEV (1970). Because of the scarcity of weath- er stations, up to 63 skins gathered on one weather sta- tion and, in these cases, I used the mean for each trait. The Bonner zoologische Beiträge 55 (2006) 63 following data were considered: yearly mean of mean, maximum, minimum and range of air temperature per day, yearly sum of precipitation, daily maximum of precipita- tion. Altitudinal effects could not be taken into account because for most skins altıtudinal information were not available with sufficient accuracy. Non-parametric Factor Analysis were performed using the MS Excel Add-In XLSTAT 7.0 (ADDINSOFT 2003). For all other calculations I used the software package SPSS for Windows 11.0 (SPSS 2001). 3. RESULTS 3.1. Data preparation Some variables were transformed to fit a normal distri- bution: beak height (reciprocal square root transforma- tion), first primary length (reciprocal), graduation of tail (natural logarithmic) and the tarsus measures (cube). All other metric variables did not differ significantly from nor- mal (Kolmogorov-Smirnov One-sample-test, p>0.05). The plumage colour variables have been subjected to a non-parametric Factor Analysis based on Spearman's rank correlation. After Varimax rotation the resulting three fac- tors with eigenvalues larger than 1 explain 88 % of the variance (Tab. 1). The first factor (hereafter called ‘front colour’ or *C1”) is highly associated with belly and throat Table 1. Correlations between plumage colour variables and factor scores from a Factor Analysis based on a Spearman's rank correlation matrix. Correlations > 0.6 printed bold. Factor _ a C1 C2 0 belly: % black -0.367 0.226 0.027 belly: % colour 0.751 -0.053 -0.134 belly: % green -0.740 0.045 0.175 breast: % black -0.056 0.248 0.943 breast: % colour 0.336 -0.223 -0.747 breast: % green -0.225 0.010 0.620 back: % black -0.147 0.717 0.332 back: % colour 0.243 -0.951 -0.188 back: % green -0.141 -0.307 0.026 throat: % black -0.567 0.160 0.119 throat: % colour 0.723 0.002 = -0.145 throat: % green -0.776 0.015 0.248 eigenvalue 4.4 1.6 fied % variance explained 387 223 27.8 colour; higher values represent individuals with more yel- lowish and less greenish front plumage. C2 (“back colour”) describes back colour, higher values correspond with more black and less colour components in the back plumage. The colour of the breast is highly correlated with C3 (‘breast colour’). Individuals with high values of C3 have darker and more greenish breasts while low C3 values in- dicate yellowish breasts. For further analysis I used the factor scores of C1-C3 for each skin. 3.2. Morphological differences between C. sulphurata and C. flaviventris At first I looked at differences in morphological charac- teristics between C. sulphurata and its sister species, C. flaviventris. A Discriminant Function Analysis (DFA) re- duces the 21 morphological variables to a single canoni- cal variable with an eigenvalue of 1.13 and a canonical correlation of 73 %. Two beak measures and two wing measures are included in the analysis (standardized canon- ical coefficients in parenthesis): beak length (1.533), beak height (0.696), Ist primary length (0.463) and distance from wing tip to secondaries (0.343). Values for C. fla- viventris are lower, thus, skins of this species are small- er in beak and wing dimensions. 95 % of all skins have been classified correctly by the DFA. An additional trait for differentiation is a red component in the front plumage of C. flaviventris: 13 of 21skins had a slight tinge of red, but none of the C. sulphurata skins. 3.3. Differentiation of C. sulphurata populations Most traits differ significantly between populations (Tab. 2). Wing and beak measures show high levels of differ- entiation (high significance values) in both males and fe- males. For a detailed analysis I tested for each trait if there is a difference between population groups. Again, the most pronounced differences are in the wing and beak meas- ures (Tab. 3). Of 21 population group pairings 16 (males) and 11 (females) differ significantly in at least one wing measure, respectively, and 12 (males) and 6 (females) in at least one beak measure, respectively. The populations of Kenya (group G) and South Africa (A) are separated by differences in most traits from all other population groups, but not from each other. In the central part of the species’ range populations vary hardly. There is a tenden- cy for females to differ in fewer traits than males across the same population group pairings. Many of the studied traits correlate with each other (e.g. wing length and primary length). Intercorrelations may oc- cur between all variables. Therefore, I performed a Prin- cipal Component Analysis (PCA) on all 24 original meas- 64 Sverre KLEMP: Patterns of geographic variation of Brimstone Canary Crithagra sulphurata Table 2. Univariate differences in C. sulphurata traits between populations (one-way ANOVA). *significant differences on the 5%-level (corrected for 24 tests: P<0.05/24=0.0021). males df F wing measures wing length 13/215 19.002 Ist primary length 13/192 8.443 2nd primary length 13/192 7.286 3rd primary length 13/197 7.083 4th primary length 13/189 1139 Sth primary length 13/195 11.383 6th primary length 13/194 15.328 7th primary length 13/195 19.416 8th primary length 13/188 17.909 9th primary length 13/191 12.721 Kıpp 13/213 2.1971 wing index | 9/61 0.534 wing index II 9/61 0.494 tail measures length 13/216 7.475 graduation 13/214 1.230 beak measures length 13/217 9.009 height 12/104 11.610 width 13/218 20.355 tarsus measures length 13/216 4.631 width 13/215 2.576 height 13/215 2.650 Plumage colour front colour 13/218 3.169 back colour 13/218 4.592 breast colour 13/218 4.067 ures (21 morphometric and 3 colour traits). It reduces the data set to five uncorrelated factors that together explain 65 % of the variance (Tab. 4). The correlations between the first factor (PC1) and length of wing, tail and all pri- maries are all relatively large and positive (1st primary ap- pears inverse due to transformation). It can be taken as a multivariate measure of size; individuals with high PC1 values are larger in size. The second factor summarizes the three beak measures: higher PC2 values indicate in- females Sig. df F Sig. 0.000* 13/146 10.548 0.000* 0.000* 11/127 4.040 0.000* 0.000* 11/128 4.660 0.000* 0.000* 11/129 6.708 0.000* 0.000* 11/124 4.818 0.000* 0.000* 11/129 6.202 0.000* 0.000* 11/131 11.721 0.000* 0.000* 11/132 11.504 0.000* 0.000* 11/131 10.368 0.000* 0.000* 11/130 7.282 0.000* 0.001 * 13/142 1.433 0.151 0.844 9/39 1.242 0.299 0.873 9/39 1.082 0.397 0.000* 13/145 5.812 0.000* 0.259 13/143 1.025 0.431 0.000* 13/147 7.826 0.000* 0.000* 13/73 9.792 0.000* 0.000* 12/146 12.610 0.000* 0.000* 13/147 3.196 0.000* 0.002 13/147 2.681 0.002 0.002 13/147 3.632 0.000* 0.000* 13/147 1.487 0.129 0.000* 13/147 2.047 0.021 0.000* 13/147 3.862 dividuals with stronger beaks (beak height transformed as Ist primary). PC3 and PC4 represent wing shape and breast colour, respectively. The fifth factor correlates neg- atively with front plumage and positively with back plumage. Individuals with high PCS scores are, therefore, darker on their back and more greenish on their front, while individuals with low PCS scores are more greenish on their back and more yellowish on their front. Bonner zoologische Beiträge 55 (2006) 65 Table 3. Differentiation of traits (raw measures) between the population groups A to G. Shown are traits with significant diffe- rences on the 5%-level (Mann-Whitney-U-test, significance level corrected for 24 tests as in Tab. 2). The diagonal separates ma- le (right) and female (left) analyses. Abbreviations: | — length, w — width, h — height; 1-9 first to ninth primary length. B, C, G: proportion of black, colour and green of breast, throat, belly and back plumage, respectively. wing: |, Kipp, wing: |, Kipp, 1-9 1-9 tarsus: w, h ; beak: |, w beak: 1, h, w wing: |, Kipp, tail | tail | 1-9 wing: 1,2, 8 wing: |, |, 3, 6 belly: CG breast: BC beak: |, h, w beak: |, w beak: 1, h, w throat: G breast: BC tail | tail | wing: l, 1-9 wing: |, 1-9 tarsus: | tarsus: | beak: l, w tail | tail | throat: G wing: l, 1-9 Schn: L,H, Br wing: 1, 2,7, 8 beak: | wing: |, 2, 5-9 tarsus: | tail | belly: G breast: C wing: 1, 1, 3-9 wing: |, Kipp, beak: 1, h, w 2, 4-9 ing: back: BC beak: |, h, w wing: | tarsus: | beak: 1, h, w tail I wing: l, 1-9 tarsus: l, w breast: BC beak: l, h, w breast: BC tail | back. BC wing: |, Kipp, 2, 7-9 tarsus: ] beak: | wing. |, 2, 3, 6-9 66 Sverre KLEMP: Patterns of geographic variation of Brimstone Canary Crithagra sulphurata Table 4. Correlations between the 24 original measures and principal component scores from a PCA (using Varımax rotation with Kaiser normalization) of the correlation matrix of 476 C. sulphurata skins. Correlations >0.600 are printed bold. Variable PC1 PC2 PC3 PC4 RES wing length 0.864 0.331 0.137 0.105 0.013 Ist primary length -0.726 -0.171 0.199 -0.108 -0.156 2nd primary length 0.780 0.124 -0.274 0.087 0.173 3rd primary length 0.800 0.120 -0.292 0.029 0.120 4th primary length 0.822 0.077 -0.298 -0.013 0.059 Sth primary length 0.858 0.119 -0.236 0.039 0.011 6th primary length 0.891 0.208 -0.012 0.103 0.022 7th primary length 0.919 0.215 0.091 0.100 0.040 8th primary length 0.905 0.224 0.141 0.092 0.050 9th primary length 0.844 0.203 0.137 0.100 0.102 Kipp distance 0.366 0.198 0.234 0.036 -0.208 wing index I -0.111 -0.029 0.946 0.037 0.041 wing index II -0.110 -0.012 0.943 0.038 0.007 tail length 0.636 0.273 0.081 0.125 0.105 tail graduation 0.012 0.182 -0.035 0.461 -0.082 tarsus length 0.286 0.048 0.052 0.625 0.189 tarsus width 0.094 0.369 -0.015 0.106 0.296 tarsus height 0.181 0.294 0.004 0.209 0.319 beak length 0.400 0.697 -0.010 0.148 -0.009 beak height -0.218 -0.823 0.033 -0.020 -0.023 beak width 0.382 0.729 -0.004 0.078 0.096 front colour 0.032 -0.153 0.027 0.133 -0.661 back colour 0.170 -0.055 0.050 0.009 0.683 breast colour 0.036 -0.011 0.071 0.810 -0.081 eigenvalues 8.172 2.463 2.270 1.452 1.296 % variance explained 34.0 10.3 9.5 6.1 5.4 Table 5. Standardized Canonical Discriminant Function Coefficients from a DFA of the five PCA scores of 443 C. sulphurata skins from the population groups A to G. PCA-factor DFI DF2 DF3 DF4 PCI 6.824 -0.633 -0.272 -0.007 PC2 0.816 0.612 -0.32 -0.184 PC4 0.52 -0.026 9.779 -0.455 PCS 0.516 0.178 0.365 0.809 eigenvalues 1.517 0.244 0.136 0.015 % variance explained 793 12.7 7.1 0.8 canonical correlation 0.776 0.443 0.346 0.122 Bonner zoologische Beitráge 55 (2006) 67 Table 6. Proportion (%) of skins classified correctly as belon- ging to population group A-G by the DFA. N - sample size. B € D E Re sG (Keer 20 790 0 0 3 32 14 Bun u97, 0 0 0 0 0 100 0 Gihe 36 0 0 0 0 8 89 3 D 14 | 0 0 0 0 21 79 0 AA E 0 0 0 40 53 4 Ba 23teh 91 0 | 0 3 4 1 GBA NO 0 0 0 18 21 53 3. de 0. x ee 3 | O. 23] : 3 0 3 0 | : ye u Z “3 0 3 Fig. 2. filled symbols - skins from population group A to G, respectively. To determine which of these factors have the highest se- lectivity for the population groups, I performed a stepwise Discriminant Function Analysis (DFA) on the PCA scores (Tab. 5). PC3 (wing shape) does not contribute to differ- entiation between groups and is removed by the analysis. The other four PCA scores are transformed to four DFA factors. 65 % of skins are classified correctly by this analy- sis. There are, however, marked differences between the population groups in the proportion of correct classifica- tion (Tab. 6). Population group A (South Africa) is clear- ly divided off from all other groups. Population group G (Kenya) is separated less, but more than 50 % of skins are classified correctly. Most overlapping of G-skins is with group A. Nearly all other skins from groups B, C, D, E and F are classified as belonging to population group E (Congo, SW Tanzania) or F (Kivu/Uganda/Rwanda). Sep- arating males and females in a single DFA each gives sim- ilar results and is, therefore, not presented here. 3 -3 0 3 = © ® > N © a PC1 (size) Crithagra sulphurata skins in the space defined by principal components | and 2 (see Table 4). Solid symbols - all skins; 68 Sverre KLEMP: Patterns of geographic variation of Brimstone Canary Crithagra sulphurata The groups B to F of eastern Africa which appear homog- enous in the above population group analysis may be dif- ferentiated on a lower level. Therefore, I calculated a DFA at population level on all skins except from population group A and G. Again, most skins are classified as belong- ing to populations of group E or F. 3.4. Pattern of geographic variation The large range of classification accuracy by DFA (Tab. 6) indicates that some population groups are more sepa- rated from their neighbours than others. Scores of all groups overlap to some extend with the characteristics of other populations (Fig. 2). The pattern of variation is much more complex than described by clear-cut population groups. As an example, in Fig. 3 / Fig. 4 wing length and 20°W 30°W 40°W Oo a .. 009 o...00 / oo e / E o... N iR e@ .00 \ e @ @ 000 o o 090. 20°s | sll a \ ® o \ \ | \ ” \ Ó 30°S N O \ / @ 82.0-84.9 mm O 0513 mm @ 750-789 mm ® 73.0-75.9 mm * <72.9 mm Fig. 3. means for 2°30'-fields with at least one studied skin. The solid Variation of wing length of C. sulphurata. Shown are line schematizes the coastline of southern and eastern Africa. beak width are shown against the geographical origin of the skins. The population group with lowest overlap to other groups is South Africa (including Natal; group A). South African C. sulphurata are characterized by strong beaks and large to intermediate body sizes (Fig. 2). Within population group A both wing and beak measures are relatively ho- mogeneous (Fig. 3, Fig. 4). Skins from Natal have a lit- tle bit lower scores in both principal components, thus dis- tinguished by little weaker beaks and little shorter wings/tails. The differences to the next population in the north (group C) are, however, much more remarkable than the differences between the South African and Natal pop- ulation (Figs 2-4). 20°W 30°W 40°W O 0? 10°S O 20°S \ \ ) 30°S ® 8.4-9.0 mm @ 0 @ 7.0-7.6mm O 6.3-6.9 mm * 5.6-6.2 mm Fig. 4. Variation of beak length of C. sulphurata. See Figure 3 for explanation. Bonner zoologische Beiträge 55 (2006) 69 In relation to all other groups Kenyan C. sulphurata (group G) look similar to skins from the other end of the species’ range, group A. Beaks of skins from Kenya are interme- diate between South African birds and all other groups, while body size is larger in Kenya than in all other groups (Fig. 2). Overlap to neighbouring populations is, howev- er, considerable. As shown by Fig. 3 and Fig. 4 the pop- ulation of Kenya is less homogenous in both beak and wing measures. Skins from northern Kenya have as long wings as skins from southern Kenya, but in respect to beak size they resemble birds from the eastern and central part of the species’ range. Outside Kenya and South Africa no remarkable morpho- metric differentiation between population groups exist. There is a tendency, however, for skins from Mocambique (B) and Angola (D) to be smallest in both principal com- ponents (Fig. 2). Wing length increases from the coasts to central Africa and could be described as a quadratic function of geographic longitude (WING=-1.23E- 10*LONG? + 6.87E-05*LONG + 66.25; R2=0.06; ANO- VA df=2/374 F=11.1 P=0.000). Beak width variation is cli- nal, too, and fits a linear function with highest values at the eastern coast (BEAK=-1.51E-06*LONG+6.82; R2=0.03; ANOVA df=1/376 F=11.2 P=0.001). Apart from this clear trend, for both measures areas with low scores alternate with high score areas, however (Fig. 3, Fig. 4). The clinal longitudinal trend is visible in the PC1/PC2 plot as a shift along the axis B (Mozambique) to C (Malawi/Zimbabwe) to E (Congo/Tanzania; Fig. 2). Due to high intra-group variability both population group F (Kivw/Rwanda/Uganda) and D (Angola) overlap with nearly all other central population groups. Skins from these two groups are, therefore, indistinguishable from the other population groups except A and G. Hence, the DFA could not dissolve the population groups B to F (Tab. 6) and classified all skins from the central part of the species” range as belonging to group F. 3.5. Impact of climate and vegetation To reveal associations between variation of morphologi- cal traits of C. sulphurata and climatic conditions I per- formed a PCA with the six climatic variables mentioned above and the five principal components of morphologi- cal measures (cf. Tab. 4). The resulting two factors explain 45 % of total variance (Tab. 7). The first principal com- ponent (cPC1) shows that body size (PC1) decreases along a gradient of increasing temperature. Large C. sulphura- ta can be found under relatively moderate temperature conditions. Beak dimensions decrease as sum of precip- itation increase. Therefore, strong beaks are associated with relatively dry climate. Table 7. Correlations between the 6 climatic variables, the 5 principal components (cf. Table 4; mean for each weather sta- tion) and principal component scores from a PCA (using Vari- max rotation with Kaiser normalization) of the correlation ma- trix of 68 weather stations. Correlations >0.6 are printed bold. variable 6PCl ePG2 mean of temperature 0.898 -0.240 maximum of temperature 0.850 -0.146 minimum of temperature 0.916 -0.089 range of temperature -0.262 -0.057 sum of precipitation -0.041 -0.801 maximum of precipitation -0.468 -0.064 PCI -0.641 0.443 PC2 0.149 0.785 PC3 -0.156 0.016 PC4 -0.089 0,327 PES -0.238 0.329 eigenvalues 3.4 1.3 % variance explained 31.0 139 Table 8. Mean variance of principal components (see Tab. 4) at eight localities (“local”; between 8 and 15 skins, total N=59) and in a random sample of the respective populations (“popu- lation”; same sample sizes). “Significant”: number of the eight local-population pairings with significant differences in varian- ce (Levene’s test, p<0.05). Component Local Population Significant PCI (body size) 0.59 0.93 2 PC2 (beak size) 0.48 0.79 l PC3 (wing shape) 1.03 0.53 | PC4 (breast plumage) 0.77 1.19 | PCS (front/back plumage) 0.80 1.44 l Skins from forested areas have lower scores of the size- related first principal component (thus, are smaller in size; mean -0.12) than skins from open landscape (mean 0.07; t-test t=2.06 d/+416 P=0.040; P all other PC >0.05). This difference is more pronounced in Kenyan skins (popula- tion group G), where it is highly significant (t=3.76 df+ 32 P=0.001). 70 Sverre KLEMP: Patterns of geographic variation of Brimstone Canary Crithagra sulphurata 3.6. Individual variation From seven localities I have series of eight or more skins that had been sampled in the same season (within three month). I compared variability of these samples with vari- ability in a random sample of same sample size taken from the same population. In principle variances of local sam- ples are slightly smaller than population variances (Tab. 8), but this difference 1s significant in only one or two of eight pairings. Mean variances of the eight samples are smaller in principal components 1 and 2 (body and beak size) than in the other variables (plumage colour). This is true for both local and population level samples. 4. DISCUSSION The morphological variability of C. sulphurata is consid- erable. Differences in body dimensions and plumage colour occur not only between populations but also indi- vidually at a single locality (Tab. 8). Therefore, 1f geo- graphical variation is studied attention has to be paid to individual variation, too. In both univariate and multivariate analyses wing and beak measures show the most striking differences between pop- ulations (Tab. 2, 5). Wing and beak measures also exhib- it lowest individual variation at one locality (Tab. 8). Therefore, these traits are suitable to separate populations and subspecies. The revisions of RAND (1968) and CLANCEY (1972) were based on wing and beak measures, too. Additionally, both authors considered slight differ- ences in plumage colouration (a little more greenish/yel- lowish etc.) to recognize subspecies. My study of a larg- er sample of skins shows, however, that plumage colour varies enormously at one site (Tab. 8) and can not be used to distinguish populations. Morphological variation of C. sulphurata parallels climat- ic trends: individuals from hot and humid regions have shorter wings and smaller beaks (Tab. 7). The pattern of variation in wing and beak size (Fig. 3, Fig. 4) could be explained by altitudinal effects to a large extend. Similar climatic conditions may be responsible for similar trait di- mensions of the populations of Kenya and South Africa, resident at the opposite borders of the species’ range. Dif- ferences in body size in relation to predominate vegeta- tion cover are probably a side-effect of climate, too, be- cause also in rain forest areas C. su/phurata settles in the scattered open habitats and not in forests (FRY & KEITH 2004). Especially in Kenya, where C. sulphurata inhab- its a broad range of altitudes, the relation between body size on one hand and climate and vegetation on the oth- er hand is strong. In this population separation of sites close to each other but differing in altitude is obvious. If wing length is taken as a measure of body size, this cli- matic trend parallels Bergmann’s rule, which suggest an increase of body size from cold to warm climates (ZINK & REMSEN 1986). There is little evidence for geographical isolation of pop- ulations due to climatic changes in the past 20.000 years. Palaeovegetation records certify distinct changes in veg- etation cover during this period (ADAMS 2004). Habitats occupied by C. sulphurata today (grasslands, scrub, sa- vannas) have been driven back due to moister conditions during the holocene (about 10.000 years ago), resulting probably in a much smaller range of C. sulphurata at that time. However, open landscapes ranged continuously from southern Africa to Kenya, at the most interrupted by a nar- row forest belt at 20-25" S. Geographic variation reflects adaptation to different en- vironmental conditions within a species’ range. Therefore, it could be seen as a model of evolutionary steps of a sin- gle population over time in a changing environment (GOULD & JOHNSTON 1972). Differences in wing shape and size between populations often parallels migratory behav- iour (LEISLER & WINKLER 1985). FRY & KEITH (2004) de- scribe C. sulphurata as resident throughout its range. Short distance (altitudinal?) movements may occur in some pop- ulations, but at the most occasionally (MACK WORTH-PREAD & GRANT 1963, HARRISON et al. 1997). Therefore, mor- phological differences more probably reflect differences in habitat structure and use since body size and shape af- fect manoeuvrability (ZINK & REMSEN 1986). As LEISLER & WINKLER (1985) have shown in their comparative study, morphological characters are strongly correlated with for- aging techniques. The variability of beak dimensions in this study suggests differences in foraging strategies or food composition of habitats within the range of C. sul- phurata. In all parts of the range C. sulphurata settles in open habitats with scattered trees, food mainly consists of seeds and small fruits (MACKWORTH-PREAD & GRANT 1963, HARRISON et al. 1997, FRY € KEITH 2004). How- ever, detailed information on geographic variation of habi- tat and food preferences is lacking. More field studies of the species are necessary to reveal this relationship. Single traits can be linked due to developmental con- straints. The PCA shows that the different measures of wing and tail size do not differ independently, but repre- sent a single principal component (Tab. 4). This corre- sponds with many other studies, where the first, highly correlated factor could be interpreted as a size factor (GOULD & JOHNSTON 1972). The fact, that beak dimen- sions, wing shape and plumage colour each form separate factors, points to habitat or food related adaptations inde- pendent of size. Bonner zoologische Beiträge 55 (2006) 71 Only the skins from South Africa can be separated clear- ly from all other populations with the data presented here. The clinal nature of variation, the large overlap of trait measures and the large individual variation do not allow the recognition of subspecies in the remaining, northern part of the species” range. C. sulphurata of the South African nominate subspecies sulphurata are characterized mainly by their strong beaks (Fig. 2). The range of this subspecies includes South Africa from the Cape Province to Natal and Zululand. In contrast, RAND (1968) recognized the population of the Cape Province as a separate subspecies, too, but classified the populations of north-eastern South Africa (Natal/Zululand) as belonging to another subspecies which includes all oth- er populations in the north. Already MEEs (1970) point- ed to the fact, that the populations of the Cape Province and Natal are different from each other in plumage colour at the most, but not in other measures (see also RAND’s own measurements). He therefore argued for combining both populations into one subspecies. The statistical analy- sis of skins presented here allows no separation between individuals from the Cape Province and Natal/Zululand, but separates between these and the populations in the north. These are isolated geographically, too, because C. sulphurata ıs lacking along the Limpopo River (HARRI- SON et al. 1997). Individuals distributed in the northern part (sharpei) are — despite marked individual variation — smaller (shorter wings) and have less strong beaks than their South African congeners. Both traits vary outside South Africa in a com- plicated pattern without abrupt changes. Variation in both wing length and beak size have a clinal and an altitudi- nal compound. The separation of subspecies is arbitrari- ly and would not contribute to a better understanding of geographical variation in this case. At best the population of the Kenyan highlands could be recognized by relative- ly strong beaks, but the transition to neighbouring popu- lations is fluid. A possible differentiation in traits not stud- ied here (e.g., genetical or ethological) have to be reserved for future studies. In conclusion, the data presented here are in agreement with the viewpoint of FrY & KEITH (2004) in respect to the subspecies sulphurata and sharpei, while the recog- nition of a subspecies wilsoni is not supported by this da- ta set. A further differentiation of populations in the north- ern part of the range as done by other authors (see above) could not be confirmed in this study. Acknowledgements. I thank Renate van den Elzen for her sup- port of this study. Michelle Louette (RMCA), Renate van den Elzen (ZFMK), Cordula Bracker and Harald Schliemann (ZMH), Mark Adams (BMNH), Jon Fjeldsá (ZMUC) and R. Diesener (ZSM) enabled access to their skin collections. Two referees gave valuable comments on the manuscript. REFERENCES ADAMS, J. 2004. Africa during the last 150,000 years. Website: http://members.cox.net/quaternary/nercAFRICA.html, 21.03.2004. ADDINSOFT 2003. XLSTAT for Windows 7.0. CLANCEY, P. A. 1972. The austral races of Serinus sulphuratus (Linnaeus). Bulletin of the British Ornithologists’ Club: 169-171. EC-JRC 2003, Hrsg. A land cover map of Africa. European Com- mission Joint Research Center, Ispra. ELZEN, R. van den 2000. Systematics and distribution patterns of Afrotropical Canaries (Serinus species group, Aves, Pas- seriformes, Carduelidae). in: RHEINWALD, G. (ed.) Isolated Vertebrate Communities in the Tropics. Proc. 4th Int. Symp., Bonn. Bonner zoologische Monographien 46: 133-143. Fry, C. H. & S. Keir 2004. The Birds of Africa. Vol. VII, Spar- rows to Buntings. Helm, London. GOULD, S. J. & JOHNSTON, R. F. 1972. Geographic variation. An- nual Review of Ecology and Systematics 3: 456-498. Harrison, J. A. et al. 1997. The Atlas of Southern African Birds. Vol. 2. Passerines. BirdLife South Africa. HEDENSTROM, A. 1989. Which wing-index should be used? Ibis 131: 154. Knox, A. G. 1993. Richard Meinertzhagen — a case of fraud ex- amined. Ibis 135: 320-325. LEBEDEV, A. N. 1970, Ed..The climate of Africa. Part 1. Air tem- perature, precipitation. Israel Progr. Sci. Transl., Jerusalem. LEISLER, B. & H. WINKLER 1985. Ecomorphology. Current Or- nithology 6: 143-174. MACKWORTH-PREAD, C. W. & GRANT, C. H. B. 1963. Birds of the Southern third of Africa. Longmans, London. MLIKOVSKY, J. 1978. Die Flügelformel der Vögel und ihre Aus- wertung. Vogelwarte 29: 268-272. RAND, A. L. 1968. Geographical Variation in the Canary Seri- nus sulphuratus. Fieldiana, Zoology 51: 119-124. Scandinavian Colour Institute 2001. Natural Colour System. In- dex edition 2. Stockholm. SPSS Inc. 2001. SPSS for Windows. Standard Version. Release 11.0.0. ZINK, R. & REMSEN, J. V. jr. 1986 Evolutionary processes and patterns of geographic variation in birds. Current Ornithol- ogy 4: 1-69. Author’s address: Sverre KLEMP, Zoologisches Institut und Museum, Universität Hamburg, Abteilung Ornitholo- gie. Present address: Rahlaukamp 4, 22045 Hamburg; E- Mail: klemp@gmx.de. Received: 15.06.2004 Revised: 09.11.2004 Accepted: 12.05.2005 Corresponding editor: Dr. Renate van den Elzen ae ne . e i ; a YN heno ny y e an Bonner zoologische Beiträge | Band 55 (2006) Heft 1 | Seiten 73-77 Bonn, Januar 2007 A New Species of Pseudachorutella Stach, 1949 (Collembola: Neanuridae) from Poland ADRIAN SMOLIS & DARIUSZ SKARZYNSKI Wroclaw, Poland Abstract. A new species of the genus Pseudachorutella Stach, 1949, P. bescidica is described from the Polish Carpathi- ans. It is related to P assigillata (Börner, 1901) and P balcanica Cassagnau et Peja, 1978 described from Germany and Albania respectively. The new species is characterized by moderate plurichaetosis on dorsal side of the body, elongated labium, the presence of the male ventral organ and four teeth on claws. Keywords. Entomology, taxonomy, Collembola, Neanuridae, Pseudachorutella, new species, Poland. 1. INTRODUCTION The genus Pseudachorutella was established by STACH (1949) with Pseudachorutes assigillatus Bórner, 1901 as the type species. Till now ten species have been described as Pseudachorutella, including four from Europe (MARI Mutt & BELLINGER 1990; MAssouD 1967; NAJT & WEIN- ER 1997; NAJT et al. 1990; WEINER & NAJT 1991). Mor- phologically, Pseudachorutella refers to Arlesia Handshin, 1942 (North and South America) and Cephalachorutes Bedos & Deharveng, 1991 (Asia, Africa). It differs from them by the set of subtle features in chaetotaxy of anten- nal segments III and IV (BEDOS & DEHARVENG 1991). Ac- cording to Deharveng (pers. comm.) it seems most prob- able that Pseudachorutella species occurring outside Eu- rope belong to two mentioned genera or a new undescribed genus. During investigations in the Polish Carpathians we have found specimens of the genus Pseudachorutella very sim- ilar to two European species, P assigillata (Börner, 1901) and P. balcanica Cassagnau et Peja, 1978 described from Germany and Albania, respectively. Examination of the material of the mentioned taxa from Börner’s and Cassag- nau’s collections has enabled us to ascertain that the Pol- ish specimens represent a new species. Its description is presented below. 2. MATERIALS AND METHODS Litter and rotting wood samples collected in the Polish Carpathians were extracted using a Tullgren apparatus. Obtained specimens of the new species were cleared in potassium hydroxide and chloral phenol and finally moun- ted on slides in Swan’s medium (distilled water, chloral hydrate, glacial acetic acid, glucose, Arabic gum) and ob- served using a phase contrast microscope Nikon Eclipse E 600. All drawings were prepared using a camera lucida. In order to compare morphology of related species the fol- lowing material was examined: P. assigillata: 4 adult ma- les, 2 adult females, Rotenburg, Germany, 27.03.1927, det. Börner (Börner’s collection, housed in Staatliches Mu- seum für Naturkunde Görlitz); P. balcanica: subadult male, Greece, det. Cassagnau (Cassagnau’s collection, housed in Museum National d Histoire Naturelle, Paris). 3. TAXONOMY Pseudachorutella bescidica sp. n. (Figs 1-10, 16-21) Types. Holotype: adult female on slide, litter and fine de- bris, deciduous forest, ca. 400 m a.s.l., N slope of the sum- mit Ostra in the “Przelom Jasıolki” reserve near Tylawa village (Beskid Niski Mountains, Carpathians, SE Poland), 16.06.2001, leg. A. Smolis. Paratypes. Adult male, subadult male, 4 juveniles on slides, same data as holotype. Other material: adult female, in piece of rotting wood, deciduous forest (Tilio-Carpine- tum), ca. 500 m a.s.l., “Obrozyska” reserve near Muszy- na, (Beskid Sadecki Mountains, Carpathians, S Poland), 03.05.2004, leg. A. Smolis, D. Skarzynski. The material is deposited in the collection of the Department of Bio- diversity and Evolutionary Taxonomy, Wroclaw Univer- sity, Poland. Description. Habitus as in Fig. 2. Body length (without antennae) 1.02-2.15 mm (holotype: 1.5 mm). Colour of 74 ADRIAN SMOLIS & DARIUSZ SKARZYNSKI: New Species of Pseudachorutella o) W ras OQ 2 a EI O Ú Nu 5 ) J El 5 UNA, Fig. 14. Pseudachorutella bescidica sp. n. 1. dorsal chaetotaxy of central part of head. 2. habitus and dorsal chaeto- taxy. 3. chaetotaxy of lateral part of tergum III. 4. chaetotaxy of abdominal sterna II-VI. 75 Bonner zoologische Beitráge 55 (2006) 0.1 mm a nm YAA MAA J Y au UL UL EU J NR Y = E S O f=) I-IV of right antenna, dorsal view. 6. antennal se , ventral view. 7. furca, genital plate and male ventral organ. 8. furca, dorsal view. 9. furca, ventral r H ai Pseudachorutella bescidica sp. n. 5. antennal segments Fig. 5-10. ments II-IV of right antenna view. 10. furca, lateral view. ADRIAN SMOLIS & DARIUSZ SKARZYNSKI: New Species of Pseudachorutella 76 > HN Perec Sag Fast e tó ( bee | 1 | labium, 12. labrum, 17. labrum, 18. claw, ıtella bescidica sp. n. (16-21). 11. igillata (11-15) and Pseudachort Pseudachorutella assi dorsolateral view; 14. tibiotars lateral view: 19. tibiotarsus III, ventral view; 20. Fig. 11-21. lateral view. 16. labium, claw, dorsolateral view; 21. tibiotarsus III, dorsal view. us III, ventrolateral view; 15. claw, 13. tibiotarsus III, Bonner zoologische Beitráge 55 (2006) 77 the body grayish-blue, eyes dark. Granulation coarse, rather homogenous (Fig. 1). Antennae shorter than head (Fig. 2). Antennal segment I with 7 chaetae, antennal seg- ment II with 13 chaetae (Figs 5, 6). Antennal segments III and IV fused dorsally. Antennal Ill-organ with two small internal curved sensilla, one ventral microsensillum and two cylindrical guard sensilla, ventral one slightly longer than dorsal (Figs 5, 6). Chaetotaxy of antennal seg- ments II-IV as in Figs 5, 6. Antennal segment IV with trilobed apical vesicle, subapical organite, microsensillum, chaeta i and 8 relatively thick, cylindrical sensilla (Fig. 5). Antennal segment IV ventrally with numerous modi- fied small sensilla (Fig. 6). Postantennal organ absent. Area ocularıs with 8 + 8 large, pigmented eyes (Figs 1, 2). Buccal cone long. Mandible delicate with 3-4 teeth, maxilla styliform. Labium with 12 + 12 chaetae, 2 + 2 small papillae x and I + | subapi- cal denticles (Fig. 16). Labium distally distinctly elongat- ed, distance between chaetae A and B greater than length of chaeta B (Fig. 16). Labrum elongated, chaetotaxy: 4-5/2, 3, 2, 2, 2. Labral edge non ogival (Fig. 17). Dorsal chaetotaxy as in Figs 1, 2. Chaeta d, on the head present or absent, unpaired chaeta ag absent. Chaetae a, on thoracic tergum II and m, on abdominal tergum IV present. Dorsal chaetotaxy moderately plurichaetotic, vari- able. 4-5 ordinary chaetae on lateral parts of terga II-III (Fig. 3). Sensorial formula of the body 022/11111. Sen- silla twice as long as ordinary chaetae. Microsensillum on thoracic tergum II present. Thoracic sterna without chaetae, ventral tube with 3-5 + 3-5 chaetae. Ventral chaetotaxy of abdominal sterna I-VI as in Figs 4, 7. Fur- ca relatively long. Dens with 6 chaetae (Figs 7, 8, 10). Mu- cro 2—2.5 times shorter than dens, triangular in shape (Figs 8, 10). Dens with ventro-apical hyaline area (Figs 9, 10). Retinaculum with 3 + 3 teeth. Male ventral organ (visi- ble only in adult male with ductus ejaculatorius) built of thickened and slightly forked chaetae on abdominal ster- na III-V (Fig. 7). Tibiotarsi I, I, IM with 19, 19, 18 chaetae respectively (Figs 19, 21). Femora I, II, III with 13, 12, 11 chaetae res- pectively. Trochanters with 6 chaetae each. Coxae I, II, III with 3, 7-8, 7-8 chaetae respectively. Subcoxae 2 I, I, III with 1, 2-4, 2-4 chaetae respectively. Subcoxae | I, II, III with 1, 2-3, 2-3 chaetae respectively. Claws with one tooth on inner edge, one tooth on outer edge and two lateral teeth (Figs 18-21). Empodial appendage absent. Derivatio nominis. Named after its terra typica, the Beskidy Mountains in Poland. Discussion. Among known members of the genus the new species is most similar to P. assigillata (Börner, 1901) and P. balcanica Cassagnau et Peja, 1978, but differs from them in the following characters: 4—5 ordinary chaetae on lateral parts of terga II-III (moderately plurichaetotic form) (P. assigillata and P. balcanica: 2 chaetae), distal- ly elongated labium (P. assigillata and P. balcanica: labi- um of normal size, see Figs 11, 12), non-ogival edge of labrum (P. balcanica also non-ogival, but in P. assigilla- ta distinctly ogival, see Figs 11, 12), labral chaetotaxy: 4-5/2, 3, 2, 2, 2 (P assigillata and P. balcanica: 4/2, 3, 5, 2, see Fig. 12), male organ present (in P. assigillata ab- sent, in P balcanica unknown), four teeth on claws (in P balcanica three teeth, outer tooth absent; in P. assigilla- ta only inner tooth present, see Figs 13-15). Acknowledgements. We wish to express our sincere thanks to Dr. H.-J. Schulz for supplying Borner’s mate- rial of P. assigillata and to Dr. L. Deharveng for loaning the specimen of P. balcanica from the collection of Prof. P. Cassagnau. We also thank to L. Deharveng and Ulrich Burkhardt for insightful comments on the manuscript. The study was sponsored by the University of Wroclaw (grant 2020/W/1Z/2004). REFERENCES BEDOS, A. & DEHARVENG, L. 1991. Cephalachorutes gen.n., a new genus of tropical Neanuridae (Collembola). Tijdschrift voor Entomologie 134: 145-153. Mari Mutt, J. A., & BELLINGER, P. 1990. A catalog of the Neotropical Collembola, including Nearctic areas of Mexico. Gainesville (Sandhill Crane Press): 237 pp. Massoup, Z. 1967. Monographie des Neanuridae, Collemboles Poduromorphes a pieces buccales modifiées. In: DELAMARE DEBOUTTEVILLE, C. & RAPOPORT, E. (eds.) Biologie de 1? Amé- rique Australe, Paris (Centre National de la Recherche Scien- tifique). NAIT, J. & WEINER, W. M. 1997. Collembola Poduromorpha de Nouvelle-Calédonie. Mémoires du Muséum National d’ His- toire Naturelle 171: 9-44. NAIT, J., THIBAUD, J.-M., WEINER, W. M. 1990. Collemboles (In- secta) Poduromorphes de Guyane francaise. Bulletin du Museum National d’ Histoire Naturelle, Paris , 4, 12A: 95-121. STACH, J. 1949. The Apterygotan fauna of Poland in rein rela- tion to the world fauna of this group of insects. Families: Anuridae and Pseudachorutidae. Acta monographica Musel Historiae Naturalis, Krakow: 122 pp. WEINER, W. M. & Nat, J. 1991. Collembola Poduromorpha of South Africa. Bonner zoologische Beitráge 42 (3-4): 369-387. Authors’ address: Adrian SmOLIs; E-Mail: adek(abiol.uni.wroc.pl; Dariusz SKARZYNSKI; E-MAIL: hy- pogast@biol.uni.wroc.pl; Zoological Institute, Wroclaw University, Przybyszewskiego 63/77, 51-148 Wroclaw, Poland. Received: 10.06.2005 Revised and accepted: 26.08.2005 Corresponding editor: B. A. 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N 088 394 Inhalt 1 3 a PFEIFFER, Martin; SCHULTZ, Roland; RADCHENKO, Alexander; YAMANE, Seiki; 1 \VOYCIECHOWSKI, Michal; ULYKPAN, Aibek & SEIFERT, Bernhard: A Critical Checklist of the Ants of Mongolia (Hymenoptera: Formicidae) VOIGLÄNDER, Karin: 9 The Life Cycle of Lithobins mutabilis L. Koch, 1862 (Myriapoda: Chilopoda) HERRMANN, Hans-Werner; SCHMITZ, Andreas; HERRMANN, Patricia A. & BÖHME, Wolfgang: Dili Amphibians and Reptiles of the Tchabal Mbabo Mtns, Adamaoua Plateau, Cameroon RHEINWALD, Gotz: 37 The Position of Trochiliphagus Carriker within the Ricinidae (Insecta: Phthiraptera) BARTSCH, Ilse: 47 The Freshwater Mite Porolohbmannella violacea (ISramer, 1879) (Acari: Halacaridae), Description of Juveniles and Females and Notes on Development and Distribution KLEMP, Sverre: 61 Patterns of Geographic Variation in Body Measures and Plumage Colour of the Brimstone Canary Crithagra sulphurata SMOLIS, Adrian & SKARZYNSKI, Dariusz: 7) A New species of Pserdachorutella Stach, 1949 (Collembola: Neanuridae) from Poland Titelbild/Cover illustration: Istylosternus sp. from Mayo Selbe (ca. 800 m elevation) (see contribution of Hans-Werner HERRMANN et al., pp. 27-35) FORSCHUNGS x useu : KOENIG Bonner zoologische Beiträge Zoologischen Forschungsmuseum Alexander Koenig, Bonn Band I) Heft 2 2007 (2006) ra! 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Organise manuscripts for Scientific Notes as follows: Title, Author Name(s), Text, Acknowledgements, References, Ad- dress(es), Tables, Figure Captions, Figures. Electronic submission. Submission of manuscripts via e-mail or on diskette is encouraged. Please, contact the editor-in-chief in advance. Reprints. 50 reprints or a PDF-file of the article are supplied free of charge. Additional reprints may be purchased by ordering when page proofs are returned. Questions we ask our reviewers Authors preparing manuscripts for Bonner zoologische Beitrage should keep in mind the following questions which referees for the journal are asked: — Is the paper of interest to the readers of Bonner zoologische Beiträge? Otherwise, suggest another journal. — Is it substantial and novel enough to justify publication? — Would the reader easily grasp the significance of the work from the introduction and discussion? — Do the conclusions follow from the information presented? — Can the paper be made clearer and more concise? — Could the title be shortened or made more informative? — Does the abstract indicate the methods used and the important re- sults? — Are alternative interpretations sufficiently considered? — Are contents and length of the paper well balanced? — Are there any important references to the subject that have not been considered? — Are all tables and illustrations necessary? — Are the tables and figures sufficient? Bonner zoologische Beiträge Band 55 (2006) Heft 2 Seiten 81-87 Bonn, Juli 2007 A new species of the genus Trapelus Cuvier, 1816 (Squamata: Agamidae) from arid central Africa LSO Philipp WAGNER & Wolfgang BOHME JAN 2 2 7008 ) Bonn, Germany / A oA LIRR Di Abstract. The Saharo-Sindian genus Trapelus contains 14 species, four of which occur in northern Africa. One of these taxa, Trapelus mutabilis, has a very widespread distribution from West to East Africa along the northern Saharan border. It has been identified as a species complex that includes several cryptic taxa. Together with a key of the so far described African species of the genus, the description of the first of these cryptic taxa is presented here. Keywords. Squamata, Agamidae, Trapelus sp. n., Africa, Chad, Ennedi mountains. 1. INTRODUCTION In his guiding work Moopy (1980) revised the family Agamidae and resurrected the genus Trapelus Cuvier, 1816. The taxa of the genus are characterized by short and thick heads and a small, deeply sunk tympanum with some spiny scales above the ear-opening. About 14 species of the genus are recognized and are distributed from north- western Africa, along the Saharan border, through the Near East to southwest and central Asia. Four of them occur in Africa [Trapelus mutabilis (Merrem, 1820): northern Africa; Trapelus pallidus (Reuss, 1833): Egypt; Trapelus savignii (Duméril & Bibron, 1837): Egypt; Trapelus tournevillei (Lataste, 1880): Algeria, Tunisia.] with a dis- tribution centre in north-eastern Africa. Most of the Egypt- ian taxa occur eastwards to the Near East and Egypt is the westernmost border of their distribution. The synonymi- sation of T. flavimaculatus with T. savignii by SALEH (1997) is not followed by us in this article, because SALEH (1997) failed to give reasons for this important taxonom- ic step. The two taxa are clearly distinct both in morpho- metrics and colouration (BAHA EL Din 2006), but the for- mer taxon is restricted to Arabia and does not extend in- to northeastern Africa. Trapelus mutabilis ıs perhaps the most complicated and widespread species within the African group. However, it is evident that this taxon represents a species complex including several cryptic taxa. The variability of 7. muta- bilis is already indicated by the high number of synonyms and different descriptions of the species in the relevant lit- erature (WERMUTH 1967). For instance, SCHLEICH et al. (1996) referred to blue throated specimens from Cyrenaica (Libya) and mentioned that the taxonomic status of these specimens remained unclear. Further work on a revision of this complex will be done by the authors in the near future. Trapelus pallidus was also for a time (e. g. WERMUTH 1967) considered a junior synonym of 7. mutabilis but is now again regarded as a valid species (e.g. PASTEUR & Bons 1960, MARX 1968, SALEH 1997). BAHA EL DIN (2006) also discussed this topic and drew attention to dif- ferences in morphology, colouration, behaviour and habi- tat. He also mentioned that 7. pallidus occurs exclusive- ly east of the Nile, whereas 7. mutabilis occurs west of the Nile. The new species described herein is closely related to 7. mutabilis but differs significantly in morphology, body proportions and colour pattern from all other known species of the genus. The 7. mutabilis complex is distrib- uted in northern Africa [Western Sahara (GENIEZ et al. 2004), Mauritania (PADIAL 2006), Morocco (PASTEUR & Bons 1960), Algeria (DOUMERGUE 1901), Tunisia (JOGER 2003), Libya (SCHLEICH et al. 1996), Egypt (BAHA EL DIN 2006), Mali (JOGER & LAMBERT 1996), Sudan (GENIEZ et al. 2004)]. Some previously described taxa are currently regarded as synonyms [7rapelus aegyptius Cuvier, 1829; Agama inermis Reuss, 1833; Agama gularis Reuss, 1833; Agama aspera Werner, 1893; Agama latastii Boulenger, 1885] of this widespread species. Records from the Near East are now known to belong to Trapelus pallidus [e. g. Israel, Jordan and Iraq fide KHALAF (1959)]. In addition to the relevant literature we used the materi- al housed in ZFMK (see Appendix) to compare the new species with voucher specimens of the relevant taxa. The synonymy follows WERMUTH (1967). Measurements and scale counts were done according to GRANDISON (1968) and Moopy & BOHME (1984). Measurements were taken with a dial calliper to the nearest 0.1 mm. 82 Philipp WAGNER & Wolfgang BÖHME: A new species of Trapelus from arid central Africa 2. RESULTS € DISCUSSION Trapelus schmitzi sp. n. Holotype. ZFMK 2590; Guelta Archei, Ennedi Moun- tains, Chad; leg. G. Niethammer, April 1954. Diagnosis. A small species of Trapelus, with a short and thick head, a dark throat and a homogenous, smooth to feebly keeled dorsal scalation intermixed with a few larg- er keeled scales of the same shape. Scalation of hindlimb homogenous. The new species differs from all described African taxa of the genus by its body proportions, a short tail, its dorsal scalation and the uniformly dark-coloured throat. Trapelus schmitzi sp. n. differs: — from 7. savignii (type locality: Egypt) and 7. flavima- culatus (type locality: Djetta, Arabien) in having smooth ventral scales and in having a very small gular pouch instead a large one. — from T. mutabilis (type locality: Egypt) in having no keeled, enlarged dorsal scales, in a higher number of preanal pores (8 to 12 in 7. mutabilis [SCHLEICH at al. 1996] instead of two rows of 18 [10+8] in 7! schmitzi sp. n.), a shorter tail (average of 102.6 mm in 7. muta- bilis and 82.65 mm in 7. schmitzi sp. n.) and a more or less homogenous dorsal scalation. — from 7: pallidus (type locality: ‘Oberägypten’= sou- thern Egypt) in having a homogenous hindlimb scala- tion. Fig. 1. Holotype of Trapelus schmitzi sp. n. Bonner zoologische Beiträge 55 (2006) 83 — from 7. tournevillei (type locality: Ouargla, Algeria) in having shorter body and head proportions, a smaller gu- lar pouch, and smooth to feebly keeled dorsal scales in- stead of keeled to strongly keeled ventral scales in 7. tournevillei. From the available and potentially valid synonyms of T. mutabilis (synonymy after WERMUTH 1967) the new species differs as follows: — from Agama inermis Reuss, 1833 (type locality: *Ober- ägypten’= southern Egypt) in having smooth to feebly keeled dorsal scales and a more or less homogenous sca- lation. i — from Agama gularis Reuss, 1833 (type locality: *Ober- ägypten’= southern Egypt) in having no strongly en- larged dorsal scales and smooth to feebly keeled dor- sal scales. — from Agama latastii Boulenger, 1885 (type locality: Egypt) in not having four pairs of quadrangular dark spots, body not depressed and in having a homogenous dorsal scalation intermixed with larger scales. — from Agama leucostigma Reuss, 1833 (type locality: “Oberágypten'= southern Egypt) in shorter head propor- tions. The latter has a long head with the broadest point before the ear hole and additionally only two mucronate scales on the superiorly margin of the ear hole. — from Agama aspera Werner, 1893 (type locality: Alger- ian Sahara between Kef-el-Dhor and Chegga; Biskra- Bordj-Saada; Zab-el-Zig south of El Meranyer) in ha- ving smooth or feebly keeled dorsal scales and in ha- ving two rows of preanal pores. Description of the holotype. Habitus stout, tail moder- ately longer than the body, limbs long. Measurements. Snout-vent length 69.1 mm; tail length 82.65 mm; head length 20.89 mm; head height 11.7 mm; head width 18.91 mm; length of forelimb 36.84 mm; length of hindlimb 51.5 mm. Scalation. Nostril on canthus rostralis, pierced in the pos- terior part of a large, flat nasal scale, directed obliquely upwards. Irregularly arranged smooth scales between nos- trils; interorbital region a median row of three more-or- less longitudinal scales separating the sideward originat- ing scales. Supraoculars smooth. Parietal scale more or less round, not enlarged; pineal organ visible, pierced in the middle; parietal scale surrounded by seven slightly en- larged scales. Scales originating from both sides of the parietal midline have their imbrications anteriorly direct- ed, free anterior margins of the scales rarely with senso- ry pits. Eyelids with a series of mucronate scales form- ing a ring. Ear-opening small, tympanum sunk, not visi- ble, about one third of the size of eye, its superior mar- gin with four spiny, mucronate scales. Rudimentary nuchal crest of only one spiny, mucronate scale. Gular scales flat, smooth, slightly imbricate at their posterior margins, be- coming somewhat smaller towards the gular fold. Gular pouch small. Dorsal scales homogenous, smooth to fee- bly keeled, partly mucronate, intermixed with few larger and feebly keeled, mucronate scales. Scales on tail smooth, becoming keeled and mucronate posteriorly, not arranged in whorls. Tail cylindrical, 20 % longer than the snout-vent Fig. 2. Holotype of Trapelus schmitzi sp. n. in comparison with other African taxa of the genus. From left to right: 7! schmitzi sp. n., holotype; 7. mutabilis, Egypt, ZFMK 2520; 7: aff. mutabilis, Sudan, ZFMK 2530; T. aff. mutabilis, Algeria, ZFMK 49664; 7. sp. n., Morocco, ZFMK 49751; 7. sp. n., Morocco, ZFMK 49741; T pallidus, Egypt, ZFMK 2537; T. tournevillei, Tunisia, ZFMK 17986. 84 Philipp WAGNER & Wolfgang BÖHME: A new species of Trapelus from arid central Africa Fig. 3. Dorsal scalation of A: 7! schmitzi, holotype; B: 7. mu- tabilis, Egypt, ZFMK 2520; C: T. aff. mutabilis, Sudan, ZFMK 2530; D: T. aff. mutabilis, Algeria, ZFMK 49664; E: T. sp. n., Morocco, ZFMK 49751; F: 7. spec. nov., Morocco, ZFMK 49741; G: 7. pallidus, Egypt, ZFMK 2537; H: 7. tournevillei, Tunisia, ZFMK 17986. length. Ventral scales smooth and slightly imbricate. Two rows of 18 (10 anterior, 8 posterior) preanal pores. Up- per forelimbs with strongly keeled scales becoming fee- bly keeled beneath, homogenous in size. 41 finger longest, digital length decreasing 3-2-5-1, plantar scales and sub- digital lamellae strongly keeled. Scales on hindlimbs ho- mogenous, smooth to feebly keeled and mucronate, be- coming more strongly keeled beneath, on the femora as large as the dorsals, becoming slightly larger towards the tibiae and feet. 4 toe longest, digital length decreasing 3-5-2-1, hindlimb long, reaching the eye with the tip of the longest digit. Colouration in alcohol. Dark grey above without any dar- ker markings, pale vertebral stripe visible, tail annulated dark grey and white, belly and under parts of the tail whitish-grey, gular region uniform dark grey to black, laterally with pale reticulations. Habitat. If the voucher was collected at Guelta Archei (see below) the habitat is thornbush savannah with sandy soils (see figs. 4 and 5). Etymology. We dedicate this new species to our good friend and colleague Dr. Andreas Schmitz, Research Of- ficer of Herpetology at the Muséum d Histoire Naturelle. Geneve, Switzerland, in recognition of his valuable con- tributions to African herpetology. Distribution and habitat. So far, the new species is on- ly known from the holotype. It was collected by the ‘Koll- mannsberger International Sahara Expedition’ in the Enne- di Mountains in Chad. The specimens collected during this expedition were divided up between several institutions. The specimens collected by Gúnther Niethammer, a well known German ornithologist, Professor at the University of Bonn and Curator of Ornithology at the Zoologisches Forschungsmuseum A. Koenig, are deposited in Bonn, while specimens collected by Franz Kollmannsberger are deposited in the collection of the University of Saarbrück- en. Because the holotype of 7. schmitzi sp. n. was cata- logued shortly after Niethammer’s return to Bonn, it be- longed to his portion of the material and was thus collect- ed by him. In the Ennedi Mountains, the expedition took two different routes (KOLLMANNSBERGER 1957). Niethammer remained at Guelta Archei to collect birds, while Kollmannsberger crossed the mountains to the northern parts of the Ennedi. Taxonomic relationships & biogeography. As men- tioned in the introduction, 7. mutabilis is a species com- plex with several cryptic taxa. 7. schmitzi sp. n. differs strongly from the typical 7. mutabilis from Egypt but re- sembles in some parts of its morphology a population of T. aff. mutabilis from the Sudan. This hitherto undescribed form (WAGNER unpubl. data) is probably the immediate sister species of 7. schmitzi sp. n. This correlates with a biogeographic pattern of the sub- Saharan savanna belt. BÖHME (1985) and Moopy & BÖHME (1984) recognized a distribution gap of typical rep- tile species of the sub-Saharan belt roughly between Ndele (Central African Republic) and El Fasher (Sudan). The Uromastyx acanthinura-geyri-dispar complex shows nearly the same distribution pattern as the Trapelus mu- tabilis complex. WILMS & BOHME (2000) differentiated three morphologically distinct clades of which the east- ern one, from Tibesti and Ennedi Mountains to Sudan, be- longs to U. dispar dispar, whereas the western populations belong to U. dispar maliensis and U. dispar flavifascia- ta. This distribution pattern suggests that Trapelus schmitzi sp. n. might also occur in the Sudan. Bonner zoologische Beiträge 55 (2006) 85 Key to the African species of Trapelus: if = Ventrals keeled; nuptial colouration of males blue, spotted white, tail reddish; females grey with dark crossbars. T. savignii — Ventrals smooth. 2 2 —Hindleg scalation heterogeneous; flat nasal scale, nos- tril on canthus rostralis, occiput with few spines, no gular pouch, dorsal scales smooth or indistinctly keeled, hindleg scalation heterogeneous, 3rd finger shorter than 4th, T. pallidus — Hindleg scalation homogenous. 3 3 -— Dorsal scalation heterogeneous; 3 finger longer Fig. 4. than 4!h. Swollen nasal scale, nostril dorsal posi- tioned, occiput without spines, ventrals smooth or feebly keeled, dorsal scales heterogeneous inter- mixed with larger scales, hindleg scalation homog- enous, small gular pouch in males. T. mutabilis complex — Dorsal scalation sub-homogenous; 3'4 finger short- er than 4th, 4 — Dorsal head scales smooth, four spinose scales on the upper border of the ear-opening dorsal scales equal, smooth to feebly keeled and mucronate, with some intermixed enlarged scales, small gular pouch, dark coloured throat, belly whitish, tail short, little longer as snout-vent-length, two rows of preanal scales in males. T. schmitzi sp. n. — Upper head scales smooth, not enlarged on occiput, no spinose scales on head, very slight fringe of point- ed scales on the upper border of the ear-opening, dorsal scales equal, granular, strongly keeled and not mucronate, one row of preanal scales in males, large gular pouch, belly whitish with dark longitudinal stripes, tail long, two and a half as long as the dis- tance from gular fold to vent, covered with equal keeled scales. T. tournevillei Habitat of T. schmitzi sp. n. around Guelta Archei, Ennedi Mts., at the time of the Kollmannsberger Expedition. 86 Philipp WAGNER & Wolfgang BÖHME: A new species of Trapelus from arid central Africa Fig. 5. Habitat of 7 schmitzi sp. n. at Guelta Archei, Ennedi Mts., at the time of the Kollmannsberger Expedition. Acknowledgements. We are grateful to Dr. Rainer Hutterer (ZFMK, Bonn) for his supply with the relevant literature and im- ages of the Kollmannsberger expedition and to Dr. Andreas Schmitz (MHNG, Geneve) for the review of an earlier draft of the manuscript. REFERENCES BAHA EL Din, S. 2006. A Guide to the Reptiles and Amphibians of Egypt. The American University ın Cairo Press, 359 pp. Boume, W. 1985. Zoogeographical patterns of the lizard fauna of the African subsaharan savanna belt, with preliminary de- scription of anew chameleon. Proceedings of the Internation- al Symposium on African Vertebrates, Bonn: 471-478. DOUMERGUE, F. 1901. Essai sur la Faune Erpétologique de l'Ora- nie. Bulletin de la Société de Geographie et d'Archéologie d’Oran 19-21: 1-404. GENIEZ, P., MATEO, J. A., GENIEZ, M. & PETHER, J. 2004. The Amphibians and Reptiles of the Western Sahara. Frankfurt Contributions to Natural History, Edition Chimaira, Volume 19, 229 pp. GRANDISON, A. G. C. 1968. Nigerian lizards of the genus Aga- ma (Sauria: Agamidae). Bulletin of the British Museum of nat- ural history, Zoology 17: 67-90. JOGER, U. 2003. Reptiles and amphibians of southern Tunisia. Kaupia 12: 71-88. JOGER, U. & LAMBERT, M. R. K. 1996. Analysis of the herpeto- fauna of the Republic of Mali, I. Annotated inventory, with description of a new Uromastyx (Sauria: Agamidae). Journal of African Zoology 110: 21-51. KHALAF, K.T. 1959. Reptiles of Iraq with notes on the amphi- bians. Bagdad (Ar-Rabitta), 96 pp. KOLLMANNSBERGER, F. 1957. Drohende Wüste. Erlebnisse und Ergebnisse der Internationalen Sahara-Expedition 1953/54. F. A. Brockhaus, Wiesbaden, 240 pp. Marx, H. 1968. Checklist of the reptiles and amphibians of Egypt. Special Publication, United States Naval Medical Re- search Unit 3, 91 pp. MERREM, B. 1820. Versuch eines Systems der Amphibien I (Ten- tamen Systematis Amphibiorum). J. C. Krieger, Marburg, 191 PP- Moopy, S. M. 1980. Phylogenetic and Historical Relationships of the Genera in the Family Agamidae (Reptilia: Lacetrilia). Unpublished Ph.D. Thesis, Univ. of Michigan. MoobY, S. M. & BÖHME, W. 1984. Merkmalsvariation und taxo- nomische Stellung von Agama doriae BOULENGER, 1885 und Agama benueensis MONARD, 1951 (Reptilia: Agamidae) aus dem Sudangürtel Afrikas. Bonner zoologische Beiträge 35: 107-128. Bonner zoologische Beiträge 55 (2006) 87 PADIAL, J. M. 2006. Commented distributional list of the repti- les of Mauritania (West Africa). Graellsia 62 (2): 159-178. PASTEUR, G. & Bons, J. 1960. Catalogue des reptiles actuels du Maroc. Travaux de l'Institut scientifique cherifien, Série Zoo- logie 21: 1-132. Reuss, A. 1933 (1934). Zoologische Miscellen, Reptilien. Ab- handlungen aus dem Gebiete der beschreibenden Naturge- schichte. Museum Senckenbergianum 1 (6): 27-62. SALEH, M. A. 1997. Amphibians and reptiles of Egypt. Publica- tion ofthe National Biodiversity Unit, Egyptian Environmen- tal Affairs Agency, no. 6, pp. 283. SCHLEICH, H. H., KÄSTLE, W. & KABISCH, K. 1996. Amphibians and reptiles of North Africa. Koeltz Scientific Books, Koenigstein, 627 pp. WILMS, T. & BÖHME, W. 2000. Revision der Uromastyx acant- hinura-Artengruppe, mit Beschreibung einer neuen Art aus der Zentralsahara (Reptilia: Sauria: Agamidae). Zoologische Ab- handlungen des Museum für Tierkunde Dresden 51: 73-104. WERMUTH, H. 1967. Liste der rezenten Amphibien und Repti- lien: Agamidae. Das Tierreich 86: 1-127. Authors’ address: Philipp WAGNER (corresponding aut- hor), Wolfgang BÖHME, Zoologisches Forschungsmuseum A. Koenig, Adenauerallee 160, D-53113 Bonn, Germany. philipp.wagner.zfmk@uni-bonn.de; w.boehme.zfmk@uni-bonn.de Received: 09.03.2007 returned for revision: 09.03.2007 accepted: 17.04.2007 Corresponding Editor: R. van den Elzen APPENDIX Material examined Trapelus mutabilis & Trapelus aff. mutabilis: ALGERIA: Bechar (ZFMK 49664); Biscra (ZFMK 2487-2490); Bou Sada (ZFMK 49828); Dra el Kastir (ZFMK 2491-2492); El Beioth (ZFMK 2497); Ghardaia (ZFMK 49653-657); Hoggar: Arak (ZFMK 20079), In Eker (ZFMK 20080), Oued Dahim (ZFMK 2498- 2499), Thar-emert-Ak (ZFMK 2501-2502), Amguid (ZFMK 2500), Hoggar (ZFMK 2503); Laghout (ZFMK 19416-418); Ou- argla (ZFMK 2494-2496); Ounif (ZFMK 7431); Tadmeit (ZFMK 2493); Touggourt: Djamda (ZFMK 2452-2486, 17988). EGYPT: Assuan (ZFMK 2517-2518); Kairo (ZFMK 2514-2516, 2519-2526, 2527-2528, 64395-396); Sinai: Abu Muilah (ZFMK 2573), Ain Mouta (ZFMK 2569), Nachla (ZFMK 2565-2568), Sinai (2532-2551), Sudar (ZFMK 64400), Wadi Chbed (ZFMK 2562-2564), Wadı el Arısch (ZFMK 2552-2561), Wadı Feran (ZFMK 77473), Wadi Ramleh (ZFMK 2571-2572), Wadi Schech (ZFMK 2570); Ras Matarma (ZFMK 65474). LiBya: Tripolis (ZFMK 20848). Morocco: Akka (ZFMK 49751-754); Erfoud (ZFMK 49741); Ksar EsSouk (ZFMK 7432); Tafilalt: Erfoud (ZFMK 26183). SUDAN: Bajuda desert: Chor Abu Harraq (ZFMK 2529-2531); Dafur: Rahib Wells (ZFMK 32471-476). TUNISIA: Sousse: Sebkha (ZFMK 2504-2505); Tozeur: Oasis Stil (ZFMK 17987), El Hamma du Djerid (ZFMK 29048-049); Tu- nisian Sahara (ZFMK 2506-2513). Trapelus pallidus: EGYPT: (ZFMK 2537); JORDANIA: Azrag (ZFMK 44317); Shawbak (ZFMK 44320); Wadi Khanzira (ZFMK 44321). Trapelus sa- vignyi: EGYPT: Sinai: Ghaza (ZFMK 2574-2580). Trapelus tour- nevillei: ALGERIA: Ain Taiba (ZFMK 2600); Bir Laif: El Alia (ZFMK 2594); El Beioth (ZFMK 2599); El Muilah (ZFMK 2591-2593); El Oued (ZFMK 17984-985); Ghardaia (ZFMK 19415); Ouargla (ZFMK 2597-2598); Touggourt (ZFMK 2595- 2596). Tunisia: Hazoua (ZFMK 49571); Oasis Nefta (ZFMK 17986); Tunisian Sahara (ZFMK 2601-2604). 88 Buchbesprechung JUNKER, Thomas (2004): Die zweite Darwinsche Revolution — Geschichte des Synthetischen Darwinismus in Deutschland 1924 bis 1950. Basilisken-Presse, Marburg. 635 S., 25 Abbildungen, mit CD-ROM. ISBN 3-925347-67-4. Preis 124,00 €. Wo und wann immer die Geschichte der evolutionáren Synthe- se beschrieben wird, geraten nahezu ausschließlich angelsách- sische Autoren ins Blickfeld, allenfalls der Russe Sergej S. Chet- verikov und — als einziger Deutscher — Bernhard Rensch wer- den noch als Begründer oder „Architekten“ erwähnt (z.B. MAYR 1984, S. 456, HULL 1988, S. 57ff. oder SMocovitis 1996, S. 21). Dass im deutschen Sprachraum sowohl wesentliche Beiträge zur Modernen Synthese entwickelt worden waren, als auch nahezu parallel eine entsprechende umfassende Sicht auf die Evolution der Organısmen entstanden war, ıst wenig bekannt. Ein erster umfangreicherer Anlauf zur wissenschaftshistorischen Aufarbei- tung dieses Abschnitts der Biologiegeschichte war ein Workshop zum Thema „Gab es eine Moderne Synthese in der deutschen Evolutionsbiologie?“ am 6. und 7. Dezember 1996 in Tübingen auf Einladung des Lehrstuhls für Ethik in den Biowissenschaf- ten unter Leitung von Eve-Marie Engels. Die Vorträge dieses Workshops wurden 1999 ım Verlag für Wissenschaft und Bil- dung publiziert. (JUNKER & ENGELS 1999). Das vorliegende umfangreiche Werk von Thomas Junker ist ein ungemein detailreiches und gleichzeitig übersichtliches Kompen- dium aus inhaltlichen Kompaktfassungen wissenschaftlicher Ent- wicklungen, Biographien beteiligter Wissenschaftler und histo- rischen Analysen von Prozessen und Konsequenzen. Im Ab- schnitt 1 („Namen — Interpretationen — Rekonstruktionen“) wer- den Begriffe geklärt und werden vor allem die Abläufe in den USA und England denen in Deutschland gegenübergestellt. Der Abschnitt 2 („Die Darwinisten“) enthält 29 teils ausführliche Biographien deutschsprachiger Persönlichkeiten aus Genetik, Verhaltensforschung, Morphologie und Systematik, Anthropo- logie und Wissenschaftstheorie. Unter diesen sind neben den üb- lichen Prominenten (Konrad Lorenz, Erwin Stresemann, Bern- hard Rensch, Gerhard Heberer, Adolf Remane etc.) auch weni- ger bekannte wie Gertraud Haase-Bessell und Ludwig Rüger. Der Abschnitt 3 behandelt „Die Evolutionsfaktoren” und venti- liert besonders die Formulierung und Gewichtung verschiede- ner Faktoren in Deutschland. Der vierte Abschnitt (,,Evolutions- theorie“) enthält Würdigungen der deutschsprachigen Beiträge zur Evolutionstheorie allgemein, zur Debatte um Mikro- und Ma- kroevolution, zu Diskursen in der deutschsprachigen Palaonto- logie und der Beziehungen zwischen der Evolutionstheorie und der Eugenik. Im fünften und letzten Abschnitt („Internationaler Darwinismus“) wird — etwas entgegen dem Wortlaut der Über- schrift — hauptsächlich die Formierung einer Synthetischen Theo- rie der Evolution in Deutschland durch die „Architekten“ Erwin Baur, Nikolai W. Timofceff-Ressovsky, Walter Zimmermann und Bernhard Rensch rekonstruiert, und erst auf den letzten zehn Sei- ten geht es explizit um die internationale Rezeption der deutschsprachigen Synthese. Abgerundet wird die riesige Daten- und Ideenfülle durch ein Li- teraturverzeichnis von 83 Seiten — das entspricht knapp 1000 auf- geführten Quellen. Das Buch enthält keinen Index, aber eine CD ROM mit dem kompletten Text des Buches als PDF-Datei. Na- túrlich erschließt sich der ungeheure Vorteil dieser Kombinati- on nur, wenn man gerade Zugang zu einem Computer mit CD ROM-Laufwerk hat. Das dürfte meistens der Fall sein, denn man wird das vorliegende Buch mit seinen Außenmaßen von 24,8 x 17,5 x 4,5 cm und seinem Gewicht von 1,4 kg wohl eher am Schreibtisch als in der Straßenbahn lesen. Für alle, die bisher innere Widerstände gegen das Medium „Digitales Buch“ hegen, sei die gewählte Verbindung von (hervorragend) auf Papier ge- drucktem Buch und beigefügter digitaler Version gepriesen. Um wie viel leichter und umfänglicher die Möglichkeiten zur Re- cherche auf diese Weise als auf herkömmliche sind, kann schwer- lich unterschätzt werden. Der Text liest sich bei aller Komplexität angenehm. Gut gewähl- te Abbildungen und Tabellen lockern nicht nur das Erscheinungs- bild auf, sondern erleichtern es mitunter erheblich, den Überblick über Zusammenhänge zu erwerben oder zu behalten. Die 357 Anmerkungen können zum Glück bei der Lektüre getrost igno- riert werden, sie enthalten in der Tat nur Zusatz-Information. Wä- re dem nicht so, hätte es wegen des dauernden Zwangs zum Um- blättern von bis zu 500 Seiten eine bedauerliche mechanische Belastung des wunderschön produzierten Buches zur Folge. Autor und Verleger sind aus Sicht des Rezensenten keine we- sentlichen Fehler oder Versäumnisse vorzuhalten. Dass trotz der offenkundig immens ausführlichen Literatur-Recherche und der gründlichen Darstellung des Lebens und des Werks von Wilhelm Ludwig die ungewöhnlich detaillierte und einflussreiche Arbeit von Janis Antonovics nicht zitiert ist, fällt bei der Fülle sonsti- ger Quellen kaum ins Gewicht. Es kann kaum unbemerkt blei- ben, dass ım Vergleich zu Genetik und Systematik die Paläon- tologie deutlich weniger genau beleuchtet wird, und dass zwar „Phylogenie“ als Überschrift auftaucht, „„Phylogenetik“ als Zweig der Evolutionsforschung aber gar nicht behandelt wird. Dies jedoch passt zu einer unverkennbaren Neigung des Autors, Ernst Mayr’s Sicht auf die Geschichte der Biologie zu akzep- tieren, z.B. in der Kennzeichnung der von Wolf Herre, Adolf Re- mane und Kurt Kosswig begründeten Phylogenetischen Sympo- sien als „antidarwinistisch“. Letztlich liegt es allein in der Kom- petenz des Autors, derlei Wertungen vorzunehmen. Hervorzuheben ist, dass die persönlichen und institutionellen Verstrickungen von Biologie bzw. Biologen und nationalsozia- listischer Ideologie und Praxis durchgängig offengelegt und nüchtern dargestellt werden. Dabei ist höchst anerkennenswert, dass der Autor bei aller Offenheit stets jede — möglicherweise ungerechte — moralische Verurteilung der erwähnten Personen vermeidet. Der doch erhebliche Kaufpreis wird die Verbreitung dieses Wer- kes bedauerlicherweise deutlich beschränken. Nichtsdestoweni- ger ist es zur eklektischen Lektüre ebenso uneingeschränkt zu empfehlen wie zum intensiven Studium. ANTONOVICS, J. 1990. Wilhelm Ludwig and his contributions to population genetics. Trends in Ecology and Evolution 5: 87-90. Hutt, D.L. 1988. Science as a Process — An Evolutionary Ac- count of the Social and Conceptual Development of Sci- ence. Chicago University Press, Chicago - London. JUNKER, T. & ENGELs, E.-M. (eds.) 1999. Die Entstehung der Synthetischen Theorie — Beiträge zur Geschichte der Evolutionsbiologie in Deutschland 1930-1950 (Verhand- lungen zur Geschichte und Theorie der Biologie 2). Ver- lag für Wissenschaft und Bildung, Berlin. MAYR, E. 1984. Die Entwicklung der biologischen Gedanken- welt — Vielfalt, Evolution und Vererbung. Springer, Ber- lin etc. Smocovrris, V.B. 1996. Unifying Biology — The Evolutionary Synthesis and Evolutionary Biology. Princeton Univer- sity Press, Princeton NJ. M. SCHMITT, Zoologisches Forschungsmuseum Alexander Koe- nig, Bonn (m.schmitt@uni-bonn.de) Bonner zoologische Beltráge Band 55 (2006) Heft 2 Seiten 89-94 Bonn, Juli 2007 Populationsgenetische Untersuchungen zur Differenzierung des schleswig-holsteinischen Rehwildes (Capreolus capreolus Linnaeus, 1758; Artiodactyla: Cervidae) Frank E. ZACHOS, Marthe OTTO, San San HMWE & Günther B. HARTL Kiel, Germany Abstraet. Fifty-nine roe deer specimens from four populations in Schleswig-Holstein, northern Germany, were analy- sed with regard to variability at eight polymorphic microsatellite loci. Two populations were island populations with known reintroduction and translocation histories and two were located on the mainland. The island populations exhibited lower variability as measured by heterozygosity, allelic diversity and allelic richness. Their demographic history is mirrored by the genetic data. Population differentiation was relatively high in terms of occurrence of private alleles (alleles exclusi- ve to only one population) but less so in terms of the overall fixation index, which was only about 6 %, and pairwise ge- netic distances. Overall deviations from HARDY-WEINBERG expectations were found in all populations studied. Possible explanations include population subdivision (WAHLUND effect) and inbreeding due to small population sizes. The relati- onship between population viability as seen from the hunters” vs. the geneticists’ viewpoint is briefly discussed in the light of the present study. Keywords. Capreolus capreolus, Schleswig-Holstein, island populations, microsatellites. 1. EINLEITUNG Das Reh (Capreolus capreolus Linnaeus, 1758) ist mit e1- nem geschätzten Bestand von allein in Mitteleuropa etwa 15 Millionen Tieren (MITCHELL-JONES et al. 1999) die häu- figste Huftierart des Kontinents. Nach dem beinahe völ- ligen Zusammenbruch des schleswig-holsteinischen Be- standes in der Mitte des 19 Jh. hat das Rehwild mittler- weile ein Rekordhoch erreicht (JESSEN 1988). Entspre- chend hoch ist seine Bedeutung auch für die Jagd (Strek- ke in Schleswig-Holstein im Jagdjahr 2002/2003 über 50.000 Tiere, Jagd und Artenschutz Jahresbericht 2003). Die vorliegende Arbeit soll einen Eindruck der genetischen Variabilität und Differenzierung, gemessen an acht poly- morphen Mikrosatellitenloci, verschiedener Rehwildpo- pulationen Schleswig-Holsteins vermitteln und überdies die beiden interessanten Inselpopulationen von Föhr und Fehmarn, die sich trotz Isolation und Gründereffekt (s. u.) durch hohe jagdliche Qualität auszeichnen, genetisch cha- rakterisieren. Der gegenwärtige Rehbestand auf Fehmarn geht auf acht 1935 eingeführte Gründertiere, drei Böcke und fünf Rik- ken, zurück, nachdem die bereits vor dem Ersten Weltkrieg dort ausgesetzten Rehe wieder ausgestorben waren (NIET- HAMMER 1963). Die acht Tiere stammen aus einem Revier von der dänischen Insel Seeland, das sich durch beson- ders vitales Rehwild (gemessen an Gewicht und Geweih) auszeichnet (ibid.), was die entsprechende Qualität des Fehmaraner Bestandes erklärt, wenn auch bemerkenswert ist, dass sich die Qualität trotz der geringen Größe der Gründerpopulatıon gehalten hat. Nach Auskunft der orts- ansässigen Jäger gelangen keine Rehe vom Festland auf die Insel. Die Analyse dieser Population dient einerseits dazu, die genetischen Folgen der beinahe 70-jahrigen Iso- lation zu charakterisieren, und andererseits kann sie An- haltspunkte dafür liefern, ob die jagdliche Beurteilung ei- nes Bestandes auf der Basıs von Wildgewicht und Trophä- enqualität Schlußfolgerungen auf die genetische Variabi- lität erlaubt oder nicht. Die Föhrer Population geht auf ein wissenschaftliches Ex- periment zurück (NIETHAMMER 1963, RIECK 1956): Zur Abschätzung des relativen Einflusses von genetischer Ver- anlagung und Umweltbedingungen sollte ein neuer Be- stand aus besonders starkem Rehwild in einem ungünsti- gen Habitat und einer aus besonders schwachem Rehwild in einem günstigen Habitat gegründet werden. Derjenige aus starkem Rehwild, für den die Insel Norderney ausge- wählt wurde, ging bald zugrunde, so dass der Föhrer Ver- such mit schwachem Rehwild blieb. Im Jahr 1939 wur- 90 Frank E. Zachos et al.: Populationsgenetische Untersuchungen zur Differenzierung des schleswig-holstein. Rehwildes den zwei Böcke und zwei Ricken aus einem Gebiet nord- westlich von Köln sowie ein Bock aus einem Revier west- lich von Düren auf der Insel ausgesetzt, jedoch verende- ten ein Bock sowie eine Ricke noch in demselben Jahr, so dass die Gründerpopulation aus nur zwei Männchen und einem Weibchen bestand (RIECK 1956). In den Jah- ren 1958/59 erfolgte eine „Blutauffrischung‘ mit zehn dá- nischen Tieren (jeweils fünf Männchen und Weibchen), weil Degenerationserscheinungen vorgelegen hätten (An- onym 1990), die von RIECK (1956) allerdings nicht er- wähnt werden. Gleichzeitig bemühte man sich um einen Abschuss des ursprünglichen Bestandes, was aber nicht gelang (ibid.). Zwanzig Jahre später, 1979, wurden aber- mals fünf dänische Rehe, zwei Bócke und drei Ricken von der Insel Fünen, auf Föhr ausgesetzt, von denen zwei je- doch bald verendeten. Interessant ist, dass die (jagdliche) Qualität der Föhrer Rehe trotz Isolation und Inzucht über dem Landesdurchschnitt liegt und dass bereits in den Fünf- ziger Jahren, also bevor weiteres Rehwild auf die Insel verbracht wurde, eine deutliche Steigerung von Geweih- größe und Körpergewicht gegenüber den Ursprungsgebie- ten eingetreten ist (Anonym 1990, RIECK 1956). Zum Vergleich mit diesen beiden Inselpopulationen wur- den zwei durch den Nord-Ostsee-Kanal, der trotz gelegent- licher Überwindung sicherlich ein Migrationshindernis darstellt, getrennte Populationen des schleswig-holsteini- schen Festlands herangezogen: eine aus Nordfriesland so- wie eine aus dem Kreis Pinneberg (fortan als „‚Rantzau“ bezeichnet). fo) LA AULA Abb. 1. Geographische Lage der Probengebiete. 1: Föhr, 2: Nordfriesland, 3: Rantzau, 4: Fehmarn. HH = Hamburg, KI = Kiel. Bonner zoologische Beiträge 55 (2006) 9] 2. MATERIAL UND METHODEN Es wurden insgesamt 59 Rehe aus vier Populationen un- tersucht (Abb. 1): Föhr (n = 7), Nordfriesland (n = 10), Rantzau (n = 23) und Fehmarn (n = 19). Die DNA-Isola- tion erfolgte aus Lebergewebe (entnommen 2003/2004) mit dem SuperQuikGene Kit sowie mit dem DNeasy Tis- sue Kit. PCR-Amplifikation und -Überprüfung sowie Län- genanalyse der Mikrosatelliten erfolgten wie in ZACHOS et al. (2003) beschrieben. Die Annealing-Temperaturen entsprachen den in der Literatur für den jeweiligen Lo- cus angegebenen Werten, wurden bei Problemfällen aber gesenkt. Folgende Mikrosatellitenloci wurden analysiert: OarFCB304 (BUCHANAN & CRAWFORD 1993), RT1, RT7 (WILSON et al. 1997), ILSTSO08, ILSTSO58 (Kemp et al. 1995), NVHRT16, NVHRT21 und NVHRT24 (ROED & MIDTHJELL 1998). Mit dem Programm Genepop (RAYMOND & ROUSSET 1995) wurde ein Test auf Kopplungsungleichgewicht (,,lin- kage disequilibrium“) durchgeführt, um auszuschließen, dass gekoppelte Loci als scheinbar unabhängige Marker in die Analyse eingingen. Die Alleldiversitäten der Popu- lationen wurden berechnet als die durchschnittliche An- zahl unterschiedlicher Allele pro Mikrosatellitenlocus. Da dieser Index von der Stichprobengröße abhängt, wurde au- Berdem die sogenannte „allelic richness“ mit der Software Fstat (GOUDET 1995) berechnet. Dieser Wert ist ein Maß fúr die Allelvielfalt, der um Unterschiede in der Stichpro- bengröße bereinigt ist, so dass die Werte direkt vergleich- bar sind. Beobachtete (Ho) und erwartete (Hp) Heterozy- gotiewerte sowie ein Test auf signifikante Unterschiede zwischen ihnen (und damit vom HARDY-WEINBERG- Gleichgewicht) wurden mit Arlequin (SCHNEIDER et al. 2000) durchgeführt. Mit Fstat wurde außerdem ein Abwei- chen vom HARDY-WEINBERG-Gleichgewicht über alle Lo- cı getestet. Als ein Indikator für die Isolation der Popula- tionen voneinander wurden die exklusiven Allele (,,priva- te alleles“ in der englischen Literatur, SLATKIN 1995), al- so diejenigen, die in nur einer der untersuchten Popula- tionen auftraten, gezählt. Der Gesamtfixationsindex For, der denjenigen Anteil der Gesamtvariation angibt, der auf Unterschiede zwischen den Populationen entfällt (im Ge- gensatz zu Variation innerhalb der einzelnen Populatio- nen), wurde mit Arlequin berechnet. Um ein weiteres Maß für die Differenzierung der Populationen zu erhalten, wur- den mit der Genetix-Software (BELKHIR 2000) paarweise genetische Distanzen berechnet: die Chorddistanz nach CAVALLI-SFORZA und EDWARDS (1967) sowie die für klei- ne Stichprobengrößen korrigierte Nei-Distanz (NEI 1978). 3. ERGEBNISSE Alle Loci waren polymorph und wiesen 8 bis 17 Allele auf. Es trat kein Kopplungsungleichgewicht auf (weder mit noch ohne Bonferroni-Korrektur für multiple Tests), so dass alle acht Mikrosatelliten in die Multi-Locus-Ana- lyse einbezogen werden konnten. In den vier untersuch- ten Populationen fanden sich insgesamt 39 (Föhr), 60 (Nordfriesland und Fehmarn) bzw. 72 (Rantzau) Allele. Die Werte für beobachtete und erwartete Heterozygotie, Alleldiversität, Allelic Richness sowie exklusive Allele sind in Tabelle 1 zusammengefasst. Die Festlandpopula- tionen weisen — vor allem in bezug auf die Allelic Rich- ness — eine höhere Variabilität auf als die beiden Inselpo- pulationen, die ein ähnliches Variabilitátsniveau zeigen. In allen vier Populationen traten für einzelne Loci wegen Heterozygotendefizits auch nach Bonferroni-Korrektur si- gnifikante Abweichungen vom HARDY-WEINBERG-Gleich- gewicht auf. Auch über alle acht Loci gemeinsam zeig- ten alle Populationen nach Bonferroni-Korrektur signifi- kante Abweichungen. Der Gesamt-Fgrp-Wert war signifikant von Null verschie- den und betrug 0,059. Das bedeutet, dass ca. 6 % der Ge- samtvariation auf Unterschiede zwischen den vier Popu- lationen entfallen und ca. 94 % in der Variabilität der In- dividuen innerhalb der Populationen begründet liegen. Ta- belle 2 gibt eine Übersicht über die paarweise errechne- Tabelle 1. Beobachtete (Ho) und erwartete (H¡.) Heterozygotie, Alleldiversitát (AD), Allelic Richness (AR), Anzahl exklusiver Allele (“private alleles”, P) sowie relativer Anteil der exklusiven Allele für die untersuchten Populationen (FOE = Föhr, NF = Nord- friesland, RAN = Rantzau, FM = Fehmarn). Ho, Hp, AD und AR sind über alle 8 Loci gemittelt. Pop. Ho Hp AD FOE 0.58 0.80 4.9 NF 0.64 0.86 15 RAN 0.62 0.82 9.0 FM 0.55 0.78 TES AR P %P 4.5 4 10.3 mye) 10 16.7 52 18 25.0 4.6 14 23.3 92 Frank E. Zachos et al.: Populationsgenetische Untersuchungen zur Differenzierung des schleswig-holstein. Rehwildes Tabelle 2. CAvALLI-SFORZA und EDWARDS” Chorddistanzen (oberhalb der Diagonalen) sowie Distanzen nach NE! für klei- ne Stichprobengrößen (unterhalb der Diagonalen) auf der Basis der Allelfrequenzen von acht polymorphen Mikrosatellitenloci. FOE NF RAN FM FOE --- 0.419 0.386 0.382 NF 0.119 --- 0.295 0.156 RAN 0.108 0.098 --- 0.254 FM 0.117 0.082 0.081 --- ten genetischen Distanzen zwischen den Populationen. Die Stabilitat dieser Werte wird von dem Programm Genetix mit einem Permutationstest geprtift. Dieser ergab sowohl für die Nei- als auch für die Chorddistanz, dass mit Aus- nahme der Distanz zwischen Nordfriesland und Fehmarn alle Werte signifikant waren (p < 0.05). Beide Distanzma- Be zeigen das gleiche Bild: eine geringere Differenzierung zwischen Nordfriesland, Rantzau und Fehmarn als zwi- schen diesen drei Populationen und Fohr, wobei Nordfries- land jeweils die hóchste Distanz zu Fóhr aufweist. 4. DISKUSSION Da die meisten der bisher publizierten populationsgene- tischen Arbeiten zum Reh auf Alloenzymen oder Sequenz- daten basieren (z. B. HARTL et al. 1991b, 1993, HEWISON 1995, VERNESI et al. 2002), sind direkte Vergleiche mit der vorliegenden Arbeit nur eingeschränkt möglich sind. Beim Vergleich mit den Ergebnissen von LORENZINI et al. (2002) sowie WANG & SCHREIBER (2001), die Mikrosatellitenana- lysen an Rehen aus Italien bzw. Mitteleuropa durchgeführt haben, fällt auf, dass die erwartete Heterozygotie und die Alleldiversitát der Rehe aus Schleswig-Holstein höher aus- fielen — sowohl für die Festland- als auch für die Insel- populationen (die Stichprobenunterschiede sınd zu gering, als dass sie verantwortlich zu machen wären). Die beiden schleswig-holsteinischen Populationen, die WANG und SCHREIBER 1m Rahmen ihrer Studie untersucht haben, wie- sen erwartete Heterozygotiewerte von 0,50 und 0,54 so- wie Alleldiversitáten von jeweils 4,0 auf. Erstaunlich ist, dass für den Locus ILSTSOS8, den sowohl WANG und SCHREIBER als auch wir analysiert haben, in beiden Stu- dien 16 Allele gefunden wurden, und das obwohl in der vorliegenden Arbeit sowohl der geographische Rahmen als auch die Gesamtzahl untersuchter Tiere (59 gegenüber 492) wesentlich geringer waren. Offensichtlich sind die schleswig-holsteinischen Rehe an diesem Locus besonders variabel. Die auf ein Heterozygotendefizit zurückzuführenden si- gnifikanten Abweichungen vom HARDY-WEINBERG- Gleichgewicht können mehrere Gründe haben. Neben der Möglichkeit des Auftretens von Nullallelen spielen bei den Inselpopulationen natürlich die geringe Populationsgrö- ße, der Gründereffekt und Inzucht eine Rolle. Schließlich kommt möglicherweise noch die Biologie des Rehs zum Tragen: Die ausgeprägte Territorialität kann dazu führen, dass sich innerhalb eines als panmiktische Population be- trachteten Bestandes Untergruppen bilden, was zu einem relativen Anstieg der Inzucht und folglich zu einem He- terozygotendefizit führt (WAHLUND-Effekt). Die Differenzierung zwischen den Populationen ergibt für beide berechneten genetischen Distanzen ähnliche Ergeb- nisse. Für die korrigierte NEI-Distanz liegen Vergleichs- werte aus der Untersuchung von WANG & SCHREIBER (2001) vor: Die Werte der vorliegenden Studie liegen zwar im Bereich der Werte aus der zitierten Arbeit, sind insge- samt aber — vermutlich wegen des kleineren geographi- schen Rahmens — durchschnittlich deutlich niedriger. Die Tatsache, dass Föhr die größten Distanzen gegenüber den anderen Populationen aufweist, liegt eventuell darin be- gründet, dass die Föhrer Population eine Hybridpopula- tion aus westdeutschen und dänischen Tieren ist. Der Fgr- Wert von nur 6 % spricht, im Gegensatz zu der hohen An- zahl an exklusiven Allelen, für eine geringe Differenzie- rung zwischen den Populationen: LORENZINI et al. (2002, 2003) fanden für italienische bzw. spanische Rehe Werte zwischen 14 und 15%. Ein Test auf Korrelation zwischen genetischen und geographischen Distanzen (,,isolation by distance“) wurde wegen des Inselstatus sowie des alloch- thonen Charakters der Föhrer und Fehmaraner Rehe nicht durchgeführt. Unsere Ergebnisse zeigen, dass die Inselpopulationen Föhr und Fehmarn verglichen mit den beiden Festlandpopula- tionen niedrigere Variabilitátswerte haben. Insbesondere die Fehmaraner Population aber, die auf nur acht Grün- dertiere zurückgeht, weist eine recht hohe Variabilität im Verhältnis zum erwarteten Grad an Inzucht und Driftef- fekten auf. Angesichts dessen könnte man vermuten, dass doch das eine oder andere Reh den Fehmarn-Sund über- quert hat oder auf der Insel ausgesetzt wurde, ohne dass schriftliche Vermerke existieren, doch bleibt dies bloße Spekulation. Andere Inselpopulationen zeigen eine deut- lich geringere Mikrosatellitenvariabilitát, so z. B. die sar- dischen Rothirsche (ZACHOS et al. 2003 sowie unveröf- fentlichte Daten) oder, um ein Extrembeispiel zu nennen, die Graufüchse der San-Nicolas-Insel vor Kalifornien, die komplett monomorph sind (cf. AGUILAR et al. 2004)! Die typischen Verinselungseffekte Inzucht und Verlust gene- tischer Variabilität durch Drift sind auf Föhr sicherlich durch das wiederholte Aussetzen dänischer Rehe aufge- fangen worden, doch ist auch hier in der Zukunft mit ver- Bonner zoologische Beiträge 55 (2006) 93 mehrter Inzucht zu rechnen. Bemerkenswert ist die Zu- nahme der (jagdlichen) Qualität des Föhrer Rehwildes be- reits relativ kurze Zeit nach der Aussetzung (s. 0.), wor- an zu sehen ist, dass die Realisierung des genetischen Po- tentials in bezug auf Merkmale wie Körpergewicht und Geweih sensibel auf äußere Einflüsse reagiert. Jagdliche Qualität und genetische Variabilität müssen also nicht zwingend miteinander korreliert sein. Das darf indes nicht darüber hinwegtäuschen, dass Inzucht und genetische Ver- armung in kleinen, isolierten Beständen unweigerlich zu- nehmen. Dazu kommt, dass an Geweihmerkmalen ausge- richtete selektive Bejagung mit diesen Merkmalen gekop- pelte Loci ebenfalls einem gerichteten Selektionsdruck aussetzt (cf. HARTL et al. 199la, 1995a, b sowie SCRIB- NER & SMITH 1990), der nicht unbedingt den Selektions- bedingungen im Naturzustand entspricht. Eine gute jagd- liche Qualität des Wildes sollte auf keinen Fall mit feh- lender Inzuchtgefährdung gleichgesetzt werden, deren Fol- gen oft erst relativ spät im Zusammenspiel mit genetischer Verarmung phänotypisch sichtbar werden, und auch dann nicht notwendigerweise an den Merkmalen, die der jagdlichen Beurteilung zugrunde liegen. Danksagung. Die Autoren danken folgenden Personen für die Unterstützung bei der Probenbeschaffung sowie für Informatio- nen über den Rehwildbestand in Schleswig-Holstein: Herrn Ewaldsen (Nordfriesland), Herrn Gröning (Landesjagdver- band), Herrn Hewicker (Forstamt Rantzau), Herrn Pedersen (Föhr), den Herren Kühl, Reese, Serck und Wilder (Fehmarn) sowie den vielen kooperierenden Jägern. Einer der Autoren (F. E. Z.) dankt der Studienstiftung des deutschen Volkes für die fi- nanzielle Unterstützung während der Durchführung dieses Pro- jektes. LITERATUR AGUILAR, A., ROEMER, G., DEBENHAM, S., BINNS, M., GARCELON & WAYNE, R. K. 2004. 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The impact of habitat frag- mentation and social structure on the population genetics of roe deer (Capreolus capreolus L.) in Central Europe. Hered- ity 86: 703-715. WILSON, G. A., STROBECK, C., Wu, L. & CorFIN, J. W. 1997. Characterization of microsatellite loci in carıbou Rangifer tarandus, and their use in other artiodactyls. Molecular Ecol- ogy 6: 697-699. ZACHOS, F., HARTL, G. B., APOLLONIO, M. & REUTERSHAN, T. 2003. On the phylogeographic origin of the Corsican red deer (Cervus elaphus corsicanus): evidence from microsatellites and mitochondrial DNA. Mammalian Biology 68: 284-298. Anschrift des Autors: Dr. Frank E. ZACHos, Christian- Albrechts-Universität Kiel, Zoologisches Institut, Olshau- senstraße 40, D-24118 Kiel. Tel.: +49/(0)431-8804529, Fax: +49/(0)431-8801389, E-Mail fzachos@zoologie.uni- kiel.de Received: 05.07.2004 Revised: 23.11.2004 Accepted: 24.11.2004 Corresponding editor: G. Peters Bonner zoologische Beiträge Band 55 (2006) Heft 2 Seiten 95-100 Bonn, Juli 2007 Bioacoustic Characteristics and Population Numbers of Endemic Cinclodes oustaleti baeckstroemii (Aves: Furnariidae) Lönnberg, 1921 of Alejandro Selkirk Island, Chile Ingo Hann !) 2) 1) Pontifical Catholic University of Chile, Santiago, Chile 2) Present address: Institute of Landscape Ecology, University of Münster, Münster, Germany Abstract. Two types of vocalisation of the endemic Masafuera Grey-flanked Cinclodes Cinclodes oustaleti baeckstroe- mii Lönnberg, 1921 are distinguished: contact call and song. The call is a whistle-like “chic” similar to that of the main- land taxon C. o. oustaleti. The song is described for the first time and illustrated by sonagrams. It is a low, melodic se- ries of pearling notes uttered early in the morning. Song sequences vary in duration, being about two to five seconds long. The frequency amplitude of elements ranges from 5.3 to 7.7 kHz, showing core frequencies from 5.8 to 6.8 kHz. Elements are usually separated by pauses of about 0.03 seconds. At the beginning and end of a sequence the song pau- ses are extended up to 0.15 seconds. The song is low and only heard for a short distance, maybe as it is addressed to the partner at one specific site. The first census counts of this endemic species were carried out using the line transect me- thod. Both estimates of its population size (in the years 1994/95 and 2002) resulted in a total number of approximately 1500 individuals. Thus no negative population trend was detected; however, this small population confirms the “vulne- rable” conservation status of the Masafuera Cinclodes. Conservation management should include further population mo- nitoring and eradication of introduced mammals. Keywords. Vocalisations, song, call, Masafuera Grey-flanked Cinclodes, Juan Fernandez, Cinclodes oustaleti baeck- stroemii, transect method, island census, conservation basis. Resumen. Dos tipos de vocalización del endémico Churrete chico de Másafuera Cinclodes oustaleti baeckstroemii Lónn- berg, 1921 fueron distinguido, grito de contacto y canto. El grito es un gorjeo (“chic”) similar a eso del C. o. oustaleti en tierra firme. El canto ha sido descrito para primera vez y esta ilustrado por sonagrames. El canto es una serie de to- nos bajos y melódicos cantado temprano en la mañana. Secuencias de canto pueden variar en su duración, por lo menos de dos a cinco segundos. La frecuencia de los elementos tiene un rango de 5,3 a 7,7 kHz con una frecuencia principal de 5,8 a 6,8 kHz. Elementos usualmente están separados por pausas de próximamente 0,03 segundos. En el inicio y fin del canto pausas están alongadas hasta 0,15 segundos. La rara y baja presentación del canto da indicación para la ausencia de una adaptación vocal compensativa a las condiciones isleñas ventosas. Usando el método de transectos lineares, dos censos de la población han sido realizados (en los años 1994/95 y 2002), que han sido resultados en un numero total de próximamente 1500 individuos en Isla Alejandro Selkirk. Por eso no tendencia critica de la población ha sido detectada, pero el Churrete chico de Másafuera es clasificado como vulnerable. Manejo de conservación debe incluir monitorio de su población e erradicación de mamíferos introducidos. Zusammenfassung. Zwei Typen der Lautäußerung des endemischen Masafuera-Grauflanken-Uferwippers (Cinclodes oustaleti baeckstroemii Lönnberg, 1921) werden unterschieden: Kontaktruf und Gesang: Der Ruf ist ein pfiffartiges „tschick“ ähnlich dem von C. o. oustaleti auf dem Festland. Der Gesang wird hier erstmals beschrieben und durch Sonagramme illustriert. Er ist eine leise und melodisch vorgetragene Serie perlender Töne, die am frühen Morgen geäußert wird. Ge- sangssequenzen können in der Länge von etwa zwei bis fünf Sekunden variieren. Die Frequenzamplitude von Elemen- ten reicht von 5,3 bis 7,7 kHz bei einem Kernfrequenzbereich von 5,8 bis 6,8 kHz. Elemente werden durch kurze Ge- sangspausen von ca. 0,03 Sekunden voneinander getrennt. Am Anfang und Ende der Gesangsstrophe sind die Pausen bis zu 0,15 Sekunden verlängert. Der leise und nur in geringer Distanz vernehmbare Gesang ist möglicherweise an den Part- ner an einem bestimmten Ort adressiert. Mittels der Linien-Transekt-Methode wurden die ersten Populationserfassungen dieses Endemiten durchgeführt. Die darauf basierenden Gesamtbestandszahlen führten in den Jahren 1994/95 sowie 2002 jeweils zu einer Anzahl von etwa 1500 Individuen auf Alejandro Selkirk. Demnach wurde zwar kein negativer Popula- tionstrend festgestellt, aufgrund der kleinen Populationsgröße ist jedoch der Gefährdungsstatus („vulnerable“) für den Maäsafuera-Uferwipper zutreffend. Schutzmaßnahmen sollten ein weiteres Bestands-Monitoring und die Bekämpfung ein- geschleppter Säugetiere beinhalten. 96 Ingo HAHN: Bioacoustics of Cinclodes oustaleti baeckstroemii of Alejandro Selkirk Island 1. INTRODUCTION The Juan Fernandez Islands are the most important endem- ic bird area of Chile and host more endemic bird taxa than any other region of this country (cf. WEGE & LonG 1995). However, they face severe conservation problems as for example seen by the low population size of another en- demic landbird species, the Masafuera Rayadito Aphras- tura masafuerae (Philippi & Landbeck, 1866) (PHILIPPI $e LANDBECK 1866; BROOKE 1987; HAHN 1998). For the en- demic Grey-flanked Cinclodes Cinclodes oustaleti baeck- stroemii Lónnberg, 1921, a population census had never been carried out. Together with Aphrastura masafuerae it represents a biogeographical outpost of the Neotropi- cal ovenbird family (Aves: Furnartidae), being limited to only Juan Fernandez in the South-east Pacific (cf. SCHLATTER 1987). The collector Bäckström (LÖNNBERG 1921) and later ob- servers (cf. JOHNSON 1967) reported Cinclodes oustaleti baeckstroemii from Alejandro Selkirk as well as from Robinson Crusoe islands. Interpreting these sources, the population on Robinson Crusoe appears to have been smaller and more oscillating than that of Alejandro Selkirk. However, probably due to lack of adequate data, recent reviews do not differ between islands, and simply state the entire Juan Fernandez Archipelago as the distri- bution area, therefore including also Santa Clara (JARAMIL- LO 2003; REMSEN 2003). Due to its insular distribution area and the general conservation problems of the archipela- go, this endemic cinclodes was already classified as “vul- nerable” in the red list of terrestrial vertebrates of Chile by GLADE (1993). Little is known about its population and biology in gen- eral, partly in relation to the remoteness of the islands and the badly accessible terrain. An additional difficulty ham- pering any population census was the fact that its vocal- isations had not been described. However, reliable cen- sus data are fundamental for any conservation manage- ment. Presenting the first description of vocalisations and the first population data, I aim to provide basic informa- tion for the endemic Masafuera Cinclodes. It may help re- searchers and managers to acoustically identify individ- uals in the field, and to monitor the population in the fu- ture. 2. METHODS The study area is Isla Alejandro Selkirk (formerly Masa- fuera), the westernmost island of the Juan Fernandez Arch- ipelago (33°45’S and 80°45’W). Alejandro Selkirk is in the South-east Pacific 167 km west of the other major is- land Robinson Crusoe, and 769 km off the coast of Chile. The archipelago is biogeographically isolated by lacking any neighbouring islands within a 500 km-zone, and the cool Humboldt Current separating it from the South Amer- ican continent. Alejandro Selkirk (44.64 km?) is of young volcanic origin, shaped like a big rock dome and is one of the steepest islands worldwide. It is entirely part of the Juan Fernandez national park founded in 1935, and UN- ESCO Biosphere Reserve since 1977. More detailed ge- ographical descriptions may be taken from CASTILLA (1987) and SKOTTSBERG (1920-1956). Field work was carried out on the island during the aus- tral summers from 25 November 1992 to 1 February 1993, 15 December 1994 to 9 February 1995, and 23 January to 8 February 2002. Visual field identification of birds was straight foreword, basing on ARAYA et al. (1992; cf. also JARAMILLO 2003) and the original description (LÓNNBERG 1921). Acoustic identification was possible after learning the bird vocalisations while they were under visual obser- vation and taping with a DAT-Recorder (Sony, Type HD-S100). An ordinary external microphone with addi- tional wind protection was placed in suitable field posi- tion. Evaluation of the tape recordings was done with the program AVISOFT-SONAGRAPH PRO. For the estimation of bird populations a number of meth- ods have been developed during the past decades (overviews in RALPH & SCOTT 1981; BiBBY et al. 2000). The abundance data should reach a high level of compa- rability and cover a maximum area. Therefore the line tran- sect method was chosen (description by EMLEN 1971, 1977). Although a quantitative method is determined to be efficient if two-thirds of the present birds are record- ed (EBERHARDT 1978), I aimed to record a significantly higher portion. Thus, I chose a small transect width, and tried to record birds by visual as well as vocal contact. On the Juan Fernandez Islands a total of 111 line transect counts were carried out from 1992 to 2002. Different tran- sect areas and habitat types were covered (for details see HAHN 1998). 3. RESULTS Cinclodes oustaleti baeckstroemii is a small brownish-grey bird of generally inconspicuous appearance, showing long white-framed tail feathers, a whitish superciliar stripe, and a small body size (cf. also FIELDSA & KRABBE 1990; RIDGELY & TUDOR 1994; REMSEN 2003). It flicks its tail up and down like a wagtail, often in the vicinity of water in a brook. It searches rocks, boulders, stone walls, moss- es, and lichens for arthropod prey, flying mostly for short duration and distance only. Even in flight the bird reminds to wagtails (Motacilla), for example when dynamically manoeuvring with its long tail to follow a flying insect. Bonner zoologische Beiträge 55 (2006) It mostly lands on top of stones, rocks or other open places. Only rarely it may land on ferns or other tall plants. No individuals were observed inside the vegetation. Description of vocalisation types Two principal types of vocalisation were identified: call and song. The call is a monotonous “chic” or may also be titled a whistle-like call. It ıs uttered very similar throughout the whole genus, either single or in a loose se- ries of calls. The song of this endemic taxon was record- ed for the first time and is described here through son- agrams (Figs. 1 & 2). It is uttered in the early morning by only one individual of the pair (probably the male). The song is a melodic series of pearling notes. 97 Song elements are limited to frequencies between 5.3 and 7.7 kHz (Fig. la). Within this amplitude the extension of elements varies and a wave-like structure is recognised. In Figure Ib it is visible that the frequency of song ele- ments is slightly different. Single elements are mostly open below or sometimes have the structure of an “S” laid on its side (e.g., last element of Fig. 1b). The short sequence (Fig. 2) illustrates the most covered frequency level of 5.8 to 6.8 kHz. This corresponds to frequency amplitudes of single elements covering about one kilo hertz (kHz). At the beginning and end of the sequence single elements are arranged looser/slower (0.1-0.15 sec.), whereas in the cen- tral song elements are closer to each other, and thus fol- lowing faster with pauses of approximately 0.03 seconds. aa, aT ee có" ton eb Setyrta f A gfltal l (41 i a arar LEE 0,1 0,2 0,3 0,4 0,5 Fig. 1 (a & b). 0,6 0,7 0,8 09 4 sec. Song of the Mäsafuera Grey-flanked Cinclodes Cinclodes oustaleti baeckstroemii Lönnberg, 1921 on Alejandro Selkirk Island. A) Long sequence of 5.5 seconds (N=256, F=100, O=0, kHz =22); B) Section of A of I seconds duration (from 4.75 to 5.75 sec. on the time scale) (N=512, F=50, O=93.5, kHz =22 PRO are given in parentheses. .05); Evaluation levels of the program AVISOFT-SONAGRAPH 98 Ingo Hann: Bioacoustics of Cinclodes oustaleti baeckstroemii of Alejandro Selkirk Island = NY wo 2 0 O ı oo © 0,25 0,5 0,75 Fig. 2. Song of the Másafuera Grey-flanked Cinclodes Cinclodes oustaleti baeckstroemii Lönnberg, 1921 on Alejandro Selkirk ETE “N 1,25 1,5 1,75 2 Island. Short sequence of 1.8 seconds duration (N=512, F=50, O=88, kHz =22). Cinclodes oustaleti baeckstroemii 1600 1400 1200 1000 specimen numbers Census 1992-95 Fig. 3. Census 2001/02 Total population numbers of the Másafuera Grey-flanked Cinclodes Cinclodes oustaleti baeckstroemii Lönnberg, 1921 on Alejandro Selkirk Island in comparison of two census periods. Censuses were carried out using a modified version of the line trans- ect method (cf. HAHN 1998). Population numbers During census counts and additional observations from 1992 to 2002 no Masafuera Cinclodes specimen was ob- served on the islands Robinson Crusoe and Santa Clara. This is an important result, as C. o. baeckstroemii was re- peatedly observed on Robinson Crusoe during the last cen- tury (LÖNNBERG 1921; JOHNSON 1967), and even recent reviews state the whole Juan Fernandez Archipelago as the distribution area (JARAMILLO 2003; REMSEN 2003). On Alejandro Selkirk they were recorded during eight tran- sect counts with 49 specimens, and mapped during numer- ous non-transect excursions. Thus the Masafuera Cin- clodes is presently interpreted to be endemic only to Ale- jandro Selkirk, in spite of the fact that single individuals also have been seen on Robinson Crusoe in the past (LONNBERG 1921). They inhabit all elevation levels up to the top of the summit of Los Inocentes (1320 m) and may be found in all habitat types. Preferred landscape struc- tures are the washes and V-shaped Quebradas, which al- so carry water in the summer; for example, the Quebradas Tongo, Varadero, Inocentes, Vacas, Casas, Sanchez, and Guaton. The rocky beaches are only occupied where fresh- Bonner zoologische Beiträge 55 (2006) 99 water is nearby, for example at the end of Quebrada In- ocentes. However, Másafuera Cinclodes may be abundant in some upland regions too, where no permanent waters exist, namely in moist areas often covered with clouds. Few specimens were seen resting on the top of tall plants or visiting dry grasslands where no washes are nearby. Only a very few indicators exist to reconstruct the popu- lation development of Cinclodes oustaleti baeckstroemii. In the year 1917, Backstrom (LONNBERG 1921) called them “common” in certain regions of Alejandro Selkirk. In the year 1986, BROOKE (1987) made a similar general state- ment. In 1994/95 I carried out the first census of this cin- clodes, which resulted in an estimate of about 1500 indi- viduals. On this set of data no population trend or any type of fluctuation can be reconstructed for historical times. It seems that this endemic taxon was widely distributed and quite common on Alejandro Selkirk during the last cen- tury. The 2002 census also led to a size of approximate- ly 1500 specimens. Thus no changes in the population size have taken place within this seven year period. Also no significant differences were found by comparing exactly the same three transect routes on Alejandro Selkirk in 1994/95 (20 individuals) and in 2002 (19 individuals). 4. DISCUSSION The common call of C. o. baeckstroemii sounds very sim- ilar to that of C. o. oustaleti, and even to some other species of Cinclodes on the mainland. Its primer function seems to keep vocal contact between two partners or mem- bers of a family party. The song of C. o. baeckstroemii is a fine and rather low vocalisation type that 1s not heard beyond a distance of more than 100 m. Thus, no obvious audible adaptation to the harsh and windy conditions on this oceanic island can be identified. In contrast, two oth- er endemic landbirds (Aphrastura masafuerae and Sephanoides fernandensis) have developed significantly louder and/or more intense vocalisations on the Juan Fer- nandez Islands (e.g., HAHN & MATTES 2000). Another remarkable fact is that only a very few singing C. o. baeckstroemii were observed, and few songs were heard throughout the entire study time. This is remarkable as cinclodes were not rare on the island, present in many habitat types, and holding territories in most regions. Maybe singing is more frequent in early spring and large- ly decreased when males are bound in pairs. This repre- sents a difference to the island’s other endemic furnariid (Aphrastura masafuerae), which continues to sing every morning even during breeding and nestling time. Son- agram comparisons with mainland C. o. oustaleti are not yet possible, as tape-recordings of this form are lacking. The total population size of about 1500 corresponds to an average density of 3.4 individuals per 10 hectares. This seems to represent a realistic value, taking into account that the well-watered canyon bottoms are the preferred habitat, and that pastures or other dry areas remain unin- habited. Through the impact of fires and introduced goats the open habitats increased on the island since discovery in 1574. Through such destruction of vegetation, several of the canyons in the North of the island carry no water during summer, but probably have done so when the veg- etation was intact. Thus, many northern canyons present- ly are not inhabited by C. o. baeckstroemii due to lack of freshwater. Another threat comes from introduced predators. Cats Fe- lis catus f. catus (Linnaeus, 1758), Norway rats Rattus norvegicus (Berkenhout, 1769), Ship rats R. rattus (Lin- naeus, 1758), and House mice Mus musculus (Linnaeus, 1758) have landed on Alejandro Selkirk, and have grown wild (HAHN & ROMER 2002). At least cats and mature rats are able to prey on adult passerine birds (ROMER 1995), but all four are known to prey on bird broods. The impacts of these pest species on the population of the Masafuera Cinclodes have not yet been investigated. Taking into ac- count its small distribution range, especially the absence from Robinson Crusoe, this endemic species should be up- graded in the Chilean Red List from “vulnerable” (GLADE 1993) to “endangered”. The population size shows no crit- ical trend, but should be monitored for a five-year inter- val. The principal conservation goal must be the eradica- tion of all introduced mammals, but for efficiency reasons campaigns should start with goats. Acknowledgements. I am grateful to the CONAF for permis- sions (V. Region: 03/94 & 011/01), and to the Rojas and Lopez families on Juan Fernandez for hospitality, and to M. Fernan- dez in Santiago (UC) for discussion. H. Mattes (Múnster), K. Conrads, W. Beisenherz, and U. Rómer (all Bielefeld) comment- ed on terminology and literature. Thanks to DAAD (1994) and AvH (2001-03) for funding. REFERENCES ARAYA B., MILLIE, G. & BERNAL M. 1992. Guia de campo de las aves de Chile. Editorial Universitaria, Santiago de Chile. Bipsy, C. J., BURGESS, N. D., HILL, D. A. & MUSTOE, S. H. 2000. Bird census techniques. Academic Press, London & San Diego. Brooke, M. de L. 1987. The Birds of the Juan Fernandez Is- lands, Chile. ICBP Study Report No. 16: 1-50. CASTILLA, J. C. 1987. Islas oceánicas Chilenas: conocimiento científico y necesidades de investigaciones. Ediciones Univ. Católica de Chile, Santiago de Chile. EBERHARDT, L. L. 1978. Transect methods for population stud- ies. Journal of Wildlife Management 42: 1-31. EMLEN, J. T. 1971. Population densities of birds derived from transect counts. The Auk 88: 323-342. 100 Ingo Hann: Bioacoustics of Cinclodes oustaleti baeckstroemii of Alejandro Selkirk Island EMLEN, J. T. 1977. Estimating breeding season bird densities from transect counts. The Auk 94: 455-468. GLADE, A. A. 1993. Libro rojo de los vertebrados terrestres de Chile. Impresora Creces Ltda., Santiago Chile. Hann, 1. 1998. Untersuchungen zur Ökologie und zum Lebens- raum der Landvogelgemeinschaften des Juan Fernández-Ar- chipels (Chile). WWU-Munster, Münster. Hann, I. & MATTEs, H. 2000. Vocalisations of the Masafuera Rayadito Aphrastura masafuerae on Isla Alejandro Selkirk, Chile. Bioacoustics 11: 149-158. Hann, I. & RÖMER, U. 2002. Threatened avifauna of the Juan Fernändez Archipelago, Chile: the impact of introduced mam- mals and conservation priorities. Cotinga 17: 56-62. JARAMILLO, A. 2003. Birds of Chile. Princeton Univ. Press, Princeton. JOHNSON, A. W. 1967. The birds of Chile and adjacent regions of Argentina, Bolivia and Peru. Volume 2. Platt Establecimien- tos Gräficos S. A., Buenos Aires. LÖNNBERG, E. 1921. The birds of Juan Fernandez Islands. Pp. 1-17 in: SKOTTSBERG, €. (ed.). The natural history of Juan Fer- nandez and Easter islands: Zoology, Vol. 3. Almquist & Wik- sells Boktryckeri, Uppsala. PhiLippr, R. A. & LANDBECK, L. 1866. Beiträge zur Fauna Chiles. Archiv fúr Naturgeschichte 32: 121-132. RALPH, C. J. & SCOTT, J. M. 1981. Estimating the number of ter- restrial birds. Studies in Avian Biology 6: 1-630. Remsen, J. V. 2003. Family Furnariidae (ovenbirds). Pp. 162-357 in: DEL Hoyo, J., ELLIOTT, A. & CHRISTIE, D. (eds.) Handbook of the birds of the world: Broadbills to tapaculos, Vol. 8. Lynx Editions, Barcelona. RIDGELY, R. S. & TUDOR, G. 1994. The Birds of South Ameri- ca. Univ. Texas Press, Austin. RÖMER, U. 1995. Wanderratten (Rattus norvegicus) erbeuten Haussperlinge (Passer domesticus). Charadrius 31: 175-180. SCHLATTER, R. P. 1987. Conocimiento y situación de la ornito- fauna en las islas oceánicas Chilenas. Pp. 271-285 in: CASTIL- LA, J. C. (ed.) Islas oceánicas Chilenas: conocimiento cienti- fico y necesidades de investigaciones. Ediciones Univ. Católica de Chile, Santiago de Chile. SKOTTSBERG, C. 1920-1956. The natural history of Juan Fernan- dez and Easter islands, 3 Vols. Almquist & Wiksells Boktryck- eri, Uppsala. WEGE, D. C., & LONG, A. J. 1995. Key Areas for Threatened Birds in the Neotropics. Burlington Press, Cambridge. Author’s address: Ingo HAHN, Department of Ecology, Pontifical Catholic University of Chile, P.O. Box 114—D, Santiago, Chile. Present address: Institute of Landscape Ecology, University of Münster, Robert-Koch-Str. 26, D- 48149 Minster, Germany; E-mail: geoeco.ingo@web.de. Received: 20.12.2004 Revised: 20.06.2006 Accepted: 20.09.2006 Corresponding editor: R. van den Elzen Bonner zoologische Beitráge Band 55 (2006) Heft 2 Seiten 101-103 Bonn, Juli 2007 Biogeographical Isolation and Bioacoustics: the Juan Fernandez Firecrown, Sephanoides fernandensis (Aves: Trochilidae) (King, 1831), of Robinson Crusoe Island, Chile Ingo Hann !) 2) 1) Pontifical Catholic University of Chile, Santiago, Chile 2) Present address: Institute of Landscape Ecology, University of Múnster, Múnster, Germany Abstract. Three types of vocalisation of the endemic Juan Fernandez Firecrown, Sephanoides fernandensis (King, 1831), are described: courtship song, warning whistle, and territorial call. The first sonagrams of this hummingbird species are presented. Territorial vocalisations are most commonly uttered and are important to localise individuals on Robinson Cru- soe Island, Chile. Although in the bioacoustic analysis significant differences between the three vocalisations are identi- fied, they have one general character in common: audibility over a relative long distance. In this point these vocalisati- ons also differ from those of its closest continental ally, the Green-backed Firecrown 5. sephaniodes, and may represent an adaptation to the harsh island conditions and oceanic winds. Keywords. Vocalisation, song, call, Sephanoides fernandensis, island adaptation. 1. INTRODUCTION The endemic Juan Fernández Firecrown, Sephanoides fer- nandensis (King, 1831), is biogeographically the most iso- lated species of all hummingbirds (Aves: Trochilidae), be- ing limited to only one small island in the South-east Pa- cific Ocean (Isla Robinson Crusoe: 47.11 km2, 915 m high, 567 km distant to the mainland at 33°30°S & 78°50’ W). The species has been separated from its main- land ancestors for some 600,000 years (Roy et al. 1998). Males and females show the most significant sexual di- morphism and dichromatism of all investigated Trochili- dae (COLWELL 1989), and also seem to differ from each other in various ecological aspects. However, little is known about the ecology of this island endemic in detail, except that it is in threat of extinction and therefore clas- sified as “critically endangered” (Roy et al. 1999; BIRDLIFE INTERNATIONAL 2004). This partly may be due to anthropogenic habitat destruction, competition, and pre- dation by introduced mammals (BUSSE 1970; HAHN & ROMER 2002). Its vocalisations are still undescribed (cf. JARAMILLO 2003). However, a successful conservation management requires basic information on aut-ecology. Attempts to reliably estimate the population size (BROOKE 1987; MEZA 1989; STILES 1987; STONE et al. 1989; HAHN 1998) require the knowledge of its vocalisations. This is especially important as its closest mainland relative, the Green-backed Firecrown Sephanoides sephaniodes (Les- son & Garnot, 1827), reached Robinson Crusoe between 1574 and 1830 (cf. AUDOUIN 1830) and now competes with it for food sources. Presenting the first data of the Juan Fernández Firecrown’s bioacoustics, I aim to add more scientific information of this endemic and to pro- vide basic material that may help future researchers and managers to acoustically identify individuals in the field. On the basis of four field campaigns from 1992 to 2002, three main types of vocalisation have been identified: courtship song, warning whistle, and territorial song/call. The courtship song is a purring series of syllables, uttered without pause at comparatively low-pitched frequency lev- els (around 4-5 kHz) and high temporal rates, slightly falling and rising in pitch. It was recognised from the male, which was slowly flying behind the female and follow- ing her from a distance of about 50 cm. The warning call is a short and very high-pitched (8 kHz and more) whis- tle, which is uttered by both sexes. It may be heard dur- ing intra- and inter-specific conflicts with other (hum- ming)birds or in case of the presence of potential preda- tors (e.g., cats, rats, humans). This whistle is difficult to distinguish from Green-backed Firecrown vocalisations in the first instance. Most characteristic and most common- ly uttered are territorial song and calls. Male Juan Fernan- dez Firecrowns are much more abundant than females on the island and, regarding to their larger size and physical dominance, are much more territorial. Thus, most such songs / calls which a listener may hear in the field come from males (Fig. 1), but females utter them too. 102 Ingo Hann: Bioacoustics and Biogeography of Sephanoides fernandensis of Robinson Crusoe Island oO FI wad — Mo pen = 0.25 05 025 10 1.25 ah & hen nn um co 0.25 OS 06 1.0 Fig. 1. o CERA EAN e nen >? - Met, oe N 5 115 2.0 2.25 25 2.15 125 15 1.15 S Territorial vocalisations of the Juan Fernandez Firecrown Sephanoides fernandensis (KING, 1831) on Robinson Crusoe Island. Above: Sequence of the song uttered by a male in its territory. Below: call uttered by a male in its territory. The call may be uttered solely or loosely attached initially/finally to the song. Tape recordings were carried out with a DAT-Recorder (Sony; Ty- pe HD-S100) with external microphone and wind shield. Illustration bases on the program AVISOFT-SONAGRAPH PRO; pro- gram levels are: N=256, F=100, O=75, KHZ=16. The sequence (Fig. |, above) is composed of about 50 syl- lables. Every syllable is built up by one higher and one lower element. The temporal rate of syllables stays very stable throughout the whole sequence (1 syllable per 0.05 sec.). However, frequency range and amplitude of sylla- bles vary during the song, clearly falling towards the end of the sequence. The song generally 1s very variable: 1t may continue for longer time durations than shown in Fig- ure 1, repeatedly showing a wave-like structure of slight- ly falling and rising in pitch. The song reaches extreme- ly high frequency levels. Therefore this firecrown belongs to the few bird species that have voices surpassing the mark of 8 kHz. BERGMANN & HELB (1982) state that this is normally the case with very small birds of low weight. This corresponds to the weight of Sephanoides fernanden- sis (9 = 10.9 g; 9 = 7.4 g according to MEZA 1989). The call (Fig. 1, below) 1s illustrated separately from the song in this case. Although it may be attached to the latter in some cases, it represents an independent feature. It is characterised by two horizontal elements of two frequen- cy ranges (lower element on 5-5.5 and higher on 6.8-7.3 kHz). These elements function as connective structures throughout most of the call, although they end before the last vertical element. The vertical elements cover wide fre- quency ranges from at least 3.5 up to 8 kHz, without be- ing split up in a higher and a lower part. At the end of the call (1.1 to 1.5 on the time scale) four elements form a unit. The temporal rating of the vertical elements in the call is slower than that of the song, about half of the speed with one element being uttered per 0.1 seconds. The low- er temporal rate leads to a more defined structure of sin- gle elements and the overall unit than in the song. The illustrated vocalisations probably have the function of marking the territory and warning competitors not to enter the area of flower agglomerations, as concluded from observations at flowering Dendroseris and Eucalyptus plants (pres. obs. in 1994 & 2001). Probably the described vocalisations can also have the function of signalising presence to potential partners. The structure of elements, and the pauses between them, turned out to be different in song and call. This is an indication that their functions may differ, even if sometimes uttered one after another. However, yet such functional differences have not been Bonner zoologische Beiträge 55 (2006) 103 cleared up. Both territorial vocalisation types of 5. fernan- densis are similar in being piercing and loud. They should be identifiable on basis of the presented sonagrams, even by persons not familiar with this species. The field obser- vations showed song/call audibility over a relatively long distance. This may be an evolutionary adaptation of the Juan Fernandez Firecrown to the harsh island conditions and counteract the generally reduced audibility through strong oceanic winds (cf. HAHN & Mattes 2000). Mechanisms behind this may be the wide frequency range of both song and call. The quick repetition of well defined elements additionally contributes to an intensive sound- ing, heard even over long distances. Field comparisons of audibility to its nearest relative, the Green-backed Fire- crown (COLWELL 1989; Roy et al. 1998), showed larger distances for endemic Juan Fernandez Firecrowns. Green- backed generally seem to use higher frequencies and tem- poral ratings (cf. EGLI 2002) but a less extended frequen- cy range, and therefore are not heard as far as Juan Fer- nandez Firecrowns. In turn, the invasive Green-backed fly more and longer distances than Juan Fernandez Fire- crowns, and thus compensate this vocal disadvantage through their higher mobility. Acknowledgements. I am grateful to the CONAF for permis- sions (V. Region: 03/94 & 011/01), and the Rojas and Lopez fam- ilies on Juan Fernandez. K. Conrads, W. Beisenherz, and U. Römer discussed previous drafts and terminology. Grants: DAAD (1994) and Humboldt Foundation (2001-3). REFERENCES AUDOUIN, M. M. 1830. Notice sur I Histoire naturelle de l‘ile Juan Fernandez. Annales des Siences Naturelles, 1830: 21. BERGMANN, H.-H. & HELB, H.-W. 1982. Stimmen der Vögel Eu- ropas. BLV, Miinchen. BIRDLIFE INTERNATIONAL 2004. Threatened Birds of the World 2004 — CD ROM. BirdLife International, Cambridge. Brooke, M. de L. 1987. The Birds of the Juan Fernandez Is- lands, Chile. ICBP Study Report No. 16: 1-50. Busse, K. 1970. Nota preliminar sobre las poblaciones de col- ibries de las Islas Juan Fernández. Boletín Ornitológico Chileno 2: 2-3. COLWELL, R. K. 1989. Hummingbirds of the Juan Fernandez Is- lands: natural history, evolution and population status. Ibis 131: 548-566. EGLt, G. 2002. Voces de Aves Chilenas. UNORCH Publ., San- tiago. HAHN, I. & Marres, H. 2000. Vocalisations of the Masafuera Rayadito Aphrastura masafuerae on Isla Alejandro Selkirk, Chile. Bioacoustics 11: 149-158. Hann, I. & RÖMER, U. 2002. Threatened avifauna of the Juan Fernández Archipelago, Chile: the impact of introduced mam- mals and conservation priorities. Cotinga 17: 56-62. Hann, I. 1998. Untersuchungen zur Ökologie und zum Lebens- raum der Landvogelgemeinschaften des Juan Fernández-Ar- chipels (Chile). WWU-Münster, Münster. JARAMILLO, A. 2003. Birds of Chile. Princeton Univ. Press, Princeton. KING, P. P. 1831. Notes on birds collected by Capt. King in Chile. Proceedings of the Committee of Science and Correspondence of the Zoological Society of London, 1831: 29-30. Meza, J. 1989. Informe anual del proyecto “Conservación del Picaflor de Juan Fernández Sephanoides fernandensis”: Invier- no 1988-Otono 1989. CONAF, Vina del Mar. Roy, M. S., TORRES-MURA, J. C. & HERTEL, F. 1998. Evolution and history of hummingbirds (Aves: Trochilidae) from the Juan Fernandez Islands, Chile. Ibis 140: 265-273. Roy, M. S., TORRES-MURA, J. C., HERTEL, F., Lemus, M. & SPON- ER, R. 1999. Conservation of the Juan Fernandez firecrown and its island habitat. Oryx 33(3): 223-232. STILES, F. G. 1987. Observaciones sobre la situación actual del Picaflor rojo de Juan Fernández ($. fernandensis), con re- comendaciones para un estudio integral de su ecología y bi- ología poblacional. CONAF 8 FAO, Santiago de Chile. STONE, T., ROBERTS, P., GUNSTONE, K., CHISHOLM, A. & WOOLFE, M. 1989. Sin-K-Tam “88” final report. Oxford University, Ox- ford. Author’s address: Ingo HAHN, Department of Ecology, Pontifical Catholic University of Chile, P.O. Box 114-D, Santiago, Chile. Present address: Institute of Landscape Ecology, University of Münster, Robert-Koch-Str. 26, D- 48149 Münster, Germany; E-mail: geoeco.ingo(@web.de Received: 20.12.2004 Revised: 20.06.2006 Accepted: 20.09.2006 Corresponding editor: R. van den Elzen 104 Buchbesprechnung Merz, B. (ed.) 2006. Phylogeny, Taxonomy, and Biology of Tephritoid flies (Diptera, Tephritoidea). Proceedings of the 3rd Tephritoid Taxonomist’s Meeting, Geneva, 19-24. July 2004. In- strumenta Biodiversitatis 7, 274 pp. ISBN 2-88139-012-9. Price CHF 100.00. This interesting soft-cover book contains the Proceedings of the 3rd Tephritoid Taxonomist’s (sic) Meeting and is the second pub- lished volume from this series of conferences. The results of the first and second meetings held in 1998 and 2000, respectively, have also been published (ALUJA & NORRBOM 1999; FREIDBERG 2006). For the third conference the theme was expanded to in- clude researchers working with other families of the superfam- ily Tephritoidea. In the present proceedings there are 11 articles by 18 authors from eight different countries. Four families, as well as the tephritid subfamily Tachiniscinae, are examined, con- tributing to the taxonomy and phylogeny (8 articles), biology (1 article), faunistics (1 article) and morphology (1 article) of the Tephritoidea. As stated in its introduction, this volume reflects the high productivity of tephritid workers, the international co- operation among many different laboratories and covers taxa from all biogeographic regions of the world. This well bound book is published by the Muséum d’histoire naturelle, Geneve, on glossy paper with excellent black and white line drawings, including many superb habitus illustrations and photographs. The first contribution revises the eastern Asian and Papuan species of Herina of the Ulidiidae (E.P. Kameneva), with the de- scription of 10 new species and the first species-level key from this region. The second paper proposes a classification of the bizarre head structures (appendages, tubercles, stalks, antlers, etc.) that occur in the acalyptrate Diptera (A.E. Whittington), with special emphasis on the extreme head morphology of the Plastotephritinae (Platystomatidae). In the subsequent contribu- tion, the European fossil specimens from the families Pallopteri- dae, Ulidiidae and Tephritidae are reviewed (G. Gentilini, V.A. Korneyev & E.P. Kameneva), with the description of a new genus and two new species. The next paper revises the genera and species of the Tachiniscinae, treated here as a subfamily of Tephritidae (V.A. Korneyev & A.L. Norrbom). Two new gen- era and two new species are described; a key to genera and a cladistic analysis are included in this thorough study. The remain- ing contributions deal exclusively with the more typical tephri- tids. The fifth article analyses the phylogeny of the subtribe Pel- matopina, an odd group of stalk-eyed tephritids (X.-I. Chen & X.-J. Wang). In a contribution toward resolving the classifica- tion of the Dacini, 10 subgenera and 67 species groups of Da- cus are proposed (D.L. Hancock & R.A.I. Drew), based on an analysis of 32 morphological characters and includes 14 new generic synonymies and a key to subgenera. The following ar- ticle describes the 344 instar larva and biology of one of the most enigmatic Palacarctic tephritids, Malica caraganae (S.V. & V.A. Korneyev). The eighth contribution describes three new species of Gymnocarena (A.L. Norrbom), extending the distribution of this New World genus to Costa Rica. The next article describes a new Afrotropical genus and three new species of the Schistopterini (I. Zonstein & A. Freidberg). The book finishes with two articles on the ısland faunas of Tephritidae. In the first article, a checklist of 16 species from the Madeiran archipela- go ıs presented (J.T. Smit), including host plant records, ecolo- gy and island records. The second article analyses the phyloge- ny of the Hawailan tephritids based on mtDNA (J.M. Brown, M. Todd-Thompson, A. McCord, A. O’Brien & B. O’Fallon) and postulates that the 25 endemic species are the result of a single colonization event. In summary, this book ıs a stimulating collection of articles cov- ering a large aspect of the Tephritoidea. It has been very care- fully edited and the illustrations are all of high quality. This book is not only essential for all tephritoid taxonomists”, but will ap- peal to workers in all aspects of fruit fly research and in the fields of morphology, sexual selection, evolution, biogeography and ecology. The next tephritid taxonomist's meeting is planned for 2008 and I look forward to the subsequent proceedings volume and the excellent reputation that has been developed from the previous two books. B. J. SINCLAIR, Entomology Laboratory - CFIA, Ottawa REFERENCES ALUJA, M. & NORRBOM, A. L. (ed.) 1999. Fruit flies (Tephriti- dae): Phylogeny and evolution of behavior. CRC Press, Boca Raton, xvi + 944 pp. FREIDBERG, A. (ed.) 2006. Biotaxonomy of Tephritoidea. Pro- ceedings of the First Tephritidologist Meeting Israel, 28 May-2 June 2000. Israel Journal of Entomology 35- 36 (2005/6): 1-599. Band 55 (2006) Bonner zoologische Beitráge Seiten 105-112 Bonn, Juli 2007 A New Species of Living Peccary (Mammalia: Tayassuidae) from the Brazilian Amazon Marc G. M. VAN ROOSMALEN!) Lothar FRENZ2), Pim van HoorT?), Hans H. DE IoNGH4) & Herwig Leirs>) 1)Amazon Association for the Preservation of High-Biodivers Areas (AAPA), Manaus-AM, Brazil 2)Hamburg, Germany 3)Wageningen NEY and Research Centre, Wageningen, The Netherlands - Leiden University, Leiden, The Netherlands ‘University of Antwerp, Antwerpen, Belgium Abstract. Here we report on the existence of a new species of even-toed ungulate in the Brazilian Amazon, which we name Pecari maximus, the giant peccary. It represents the largest of living peccary species. One complete mitochondri- al D-loop and two nuclear SINE PRE-1 DNA sequences of giant peccary compared with that of the sympatric and mor- phologically most similar collared peccary (Pecari tajacu) support full species status. The divergence time is estimated at 1.0-1.2 million years before present. In contrast to other peccaries, which are gregarious and range semi-nomadical- ly in large home ranges, giant peccaries appear to.live in family groups containing only a pair of adults, with or without 1-2 offspring. In view of recent developments in the interfluves where it lives and due to its limited distribution, we con- sider the giant peccary endangered. Keywords. New species, Artiodactyla, Tayassuidae, Pecari maximus, giant peccary, Brazilian Amazon. 1. INTRODUCTION Until recently only three extant species in the New World Artiodactyla family Tayassuidae (peccaries) were known to science, belonging to three genera (GRUBB 2005): the collared peccary Pecari tajacu (Linnaeus 1758), the white- lipped peccary Tayassu pecari (Link 1795), and the Chacoan peccary Catagonus wagneri (Rusconi 1930). A living population of the latter which was thought to have gone extinct was discovered in 1974 in the Chaco region on the borders of Bolivia, Paraguay and Argentina (WETZEL et al. 1975). During transect surveys of megafauna and fruits conducted in January 2000 in the Rio Aripuana basin, the first author saw a group of three peccaries, not belonging to any of the known peccary species. Since then, the first author has had several encounters with such peccaries and collected some basic ecological and behavioural data. In March 2003, the first two authors succeeded in filming a group of four such peccaries and collecting zoological material from game brought into the village of Arauazinho. 2. DESCRIPTION OF SPECIES Class Mammalia Order Artiodactyla Family Tayassuidae Palmer, 1897 Genus Pecari Reichenbach, 1835 Pecari maximus sp. nov. (Figs. 1-3) Material. Five skins in the possession of hunters along the lower Rio Aripuana and a complete skull without ex- act locality data obtained from them (MR316); an incom- plete skull with skin (MR315) from an adult specimen killed in December 2003 along the Rio Arauazinho; a com- plete skull (INPA4272) from an adult male killed by lo- cals on March 12, 2003, along the left bank of the Rio Aripuana near the settlement of Arauazinho. Several skulls and mandible parts of all three peccary species occurring in the area were examined as found in the kitchen mid- dens of some local communities. Diagnosis and discussion. A species of Pecari differing from the only other known species in this genus, Pecari tajacu, in being much larger but less robust, with much longer legs and a proportionally small head only slightly 106 Marc G. M. VAN ROOSMALEN et al.: A new species of Peccary from the Brazilian Amazon Fig. 1. Wild giant peccary Pecari maximus sp. nov. visiting a pond in the middle of continuous high rain forest; a. and b. show two different adult individuals belonging to the same family of four giant peccaries (Copyright NDR Naturfilm/Roland Gockel). Fig. 2. bigger than that of Pecari tajacu. Most of the body thin- ly bristle-haired, overall colour brown mixed with dirty white, a black mid-dorsal mane running from between the ears as far as the rudimentary tail. Ears small and whitish at distal surface. Nasal disc pinkish, relatively small and soft. Collar running over the shoulders very faint, dirty white, or absent (Figs. 1 and 2). Whitish, thinly haired cir- cum-ocular rings (Fig. 1). Distinguished from all other peccary species by its larger size; thin fur; proportional- ly longer legs giving it a more gracious general appear- ance; proportionally smaller head - skull length one fifth of total body length in Pecari maximus, one quarter in Pecari tajacu and Tayassu pecari (WOODBURNE 1968), and a. (left) Hunters with freshly killed adult male specimen of Pecari maximus. The skull of this individual was saved and is here designated as the holotype (INPA4272). (Copyright by Frieder Salm); b. (right) Skin of Pecari maximus paratype speci- men MR315, lacking even a faint collar. nearly one third in Catagonus wagneri (WETZEL 1977; WRIGHT 1989); less developed nasal disc; smaller ears. The new species is assigned to the genus Pecari, because it possesses a number of traits — in particular cranial fea- tures (Fig. 3 and Table 1) — in common with sympatric Pecari tajacu, from which it differs by its much larger size and weight (40-50 versus 22 kg), less stocky and longer- legged general appearance, thin, grizzled brown and white fur (instead of thick, dense, strongly speckled dark black- ish-grey fur), blacker on the limbs and along the dorsal crest, and a very poorly expressed, sometimes absent, dirty white instead of strikingly contrasting bright white collar passing across the chest from shoulder to shoulder (BOD- Bonner zoologische Beitráge 55 (2006) 107 Fig. 3. CLEAN a. Lateral view of the complete skulls of an adult male Pecari tajacu from Rio Demeni (INPA283) (left) and an adult male Pecari maximus from Rio Aripuana (holotype INPA4272) (right). b. Dorsal view of the same skulls, Pecari tajacu (top), Pe- cari maximus (bottom). e. Ventral view of the crania of the same skulls (in same arrangement). d. Dorsal view of mandibles of the same skulls (in same arrangement). MER & SOWLS 1993; GRUBB & GROVES 1993). Distin- guished from Tayassu pecari, with which it is also sym- patric, by its larger body size and weight (40-50 vs. 28 kg), and grizzled-brown thin fur instead of thick, long, evenly coloured blackish brown fur becoming grizzled or light-coloured only in the pectoral and inguinal regions. The forelimbs and legs are only distally black, while they are grizzled black and tan on the lateral and hind surfaces of the forelimbs in Tayassu pecari (GRUBB & GROVES 1993; MARCH 1993). In contrast to the general blackish brown body colour, the chin, cheeks and sides of the muz- zle are white or yellowish-white in white-lipped pecca- ries. Distinguished from Catagonus wagneri, with which it is allopatric, by its larger body size and weight (40-50 vs. 29-38 kg), grizzled-brown thin fur with or without a faint dirty white collar instead of brownish-grey thick fur with a distinct bright white collar (WETZEL 1977; WRIGHT 1989; GRUBB & GROVES 1993). Overall, the cranium of Pecari maximus seems more re- lated to that of Pecari tajacu than that of Tayassu pecari or Catagonus wagner, but it differs in the following char- acters: it is clearly longer, wider and more robust; the frontal bone between the postorbital processes 1s wider; the rostrum behind the canines is wider, the dorsal sur- face is broader and flatter; the canine buttresses are larg- er; and the canines are more pointed with sharper edges. Comparing the cranial morphometrics of Pecari maximus with Pecari tajacu from Brazil (i.e., INPA283, MR317, MR318), where the species appears to grow bigger, es- pecially where it is sympatric with Pecari maximus, it may be noted that breadth measurements in general differ more significantly between the species than length measure- ments. For example, length of cranium, mandible, and mandibular diastema do not show much difference, where- as breadth across zygomatic arches and between postor- bital processes of frontals are much greater in Pecari maxi- 108 Bonner zoologische Beitráge 55 (2006) Table 1. Cranial and dental measurements [mm] of Pecari maximus sp. nov., Pecari tajacu and Tayassu pecari, following Woop- BURNE (1968). Pecari maximus is represented by the holotype INPA4272 and the paratypes MR316 and MR315 (incomplete skull); Pecari tajacu is represented by INPA283, an adult male from Rio Demeni, MR317, an adult male (head-body length: 1100 mm) from Rio Arauazinho, and MR318, an adult female (head-body length: 1080 mm) from Rio Arauazinho. The second column of measurements of Pecari tajacu is of specimens collected outside Brazil in the northern part of the species” range where individuals are on average smaller than those from central Amazonia; these and the measurements for Tayassu pecari were taken from Woop- BURNE (1968) Cranium P. maximus sp. nov. P. tajacu P. tajacu T. pecari (N=3) (N=3) mean mean (N=71) (N=41) Length of cranium (= length anterior tip 262;260;--- 230;255;258 202 235 of I! to rear of condyles) Length of diastema from C to P? 31;30;--- 21:23:26 18 26 (distance from rear edge of alveolus of C) Width between alveoli of P? 18;20;--- 14:17:16 18 21 Width between alveoli of M3 20;19;19 18;18;18 18 20 Least width of rostrum behind canines 39,5;40;--- 39335535 31 53 Height from condyles to nuchal crest 85:85:85 73:80:80 81 99 Breadth across zygomatic arches 119;120;119 104;110;95 101 118 Breadth between postorbital 88;88;89 78,78;71 73 92 processes of frontals Mandible Length of mandible 185;180;182 165;170;173 162 198 (= length from I! to rear of condyles) Length of diastema from C to P> 40:35:36 31537537 25 34 Depth from tip of coronoid process 88;85;84 73;82;77 75 93 to ventral angle Depth below P> 36:39:40 32:40:37 32 36 Depth below rear of M; 45:45:45 32533535 34 41 Width between alveoli of P> 20;25;24 20;22;21 22 26 Width between alveoli of My 25325324 26;27;26 27 31 Width between condyles 49°:55;--- 52:47:48 49 58 Length from P> to Mz 65:68:08 63:62:64 69 85 mus. The width between alveoli of M3 is slightly greater in Pecari maximus implying a differently shaped palate. Whereas the depth below P, is the same in the two species, the depth below rear of My is much greater in Pecari max- imus, so that the lower jaw deepens posteriorly (for a more detailed comparison, see Table 1). GRUBB (2005) lists the following 19 synonyms for Pecari tajacu: angulatus, bangsi, caitetu, crassus, crusnigrum, humeralis, macrocephalus, minor, modestus, nanus, nel- soni, niger, nigrescens, patira, sonoriensis, tajassu, torquatus, torvus, yucatanensis. Three subspecies groups have been differentiated in the past — the grey forms (‘an- gulatus” group) from Central America (Texas, Mexico and Honduras), the blackish forms with poorly expressed col- lar and dorsal stripe (‘patira’ group) from the Guianas, Colombia, Ecuador and Panama, and the buff forms with clearly marked pale collar and black dorsal stripe from South America (the nominate *tajacu” group). The syn- onyms of the South American forms apply to specimens Bonner zoologische Beiträge 55 (2006) 109 Pecari tajacu' (Arizona) Pecari tajacu? (Origin unknown Pecari tajacu” (Colombia) Pecari maximus Tayassu pecari* (Colombia) Catagonus wagneri’ (Paraguay) Sus scrofa® Phacochoerus aethiopicus’ Fig. 4. Neighbour-joining tree of combined nuclear PRE-1 P27 and P642 sequences. The tree was constructed with MEGA3 (Kumar et al. 2004), assuming the TN93 Gamma model (sha- pe parameter alpha = 0.6). Bootstrap values (10,000 replicates) are indicated on branches. Geographic origins are mentioned bet- ween brackets. The superscripts refer to Genbank accession num- bers. 1: AY546529/AY726778, 2: AB000378/379, 3: AY546532/AY726779, 4: AY546528/31, 5: AY546527/30, 6: AB000365/370, 7: AB000377/678. collected in Colombia, Ecuador and the Guianas, areas north of the Amazon River, being geographically too dis- tant from the range of the new taxon, Pecari maximus, for these synonyms to be taken into account. Linnaeus’ name Sus tajacu 1s based on the tajacu of Marcgraf (GRUBB 2005), from the State of Pernambuco, Brazil, a region that is not considered part of Amazonia. Measurements. Five skins from hunters along the lower Rio Arıpuanä were measured, total length 120, 127, 133, 135 and 137 cm. Collar when present 35 cm. Nasal disc 5.3 x 4.0 cm. Mane bristles 10.5-12.0 cm, proximally with 2-3 white bands at the lower half and 3 brown bands, dis- tally with 3 white and 4 brown bands, the white bands on- ly on lower half of bristles. Bristles to the side of the mane 6-7 cm, the dorsal ones with 3 whitish bands, the more ventral ones 4.7 cm, with only 2 whitish bands. Collar bris- tles 3.6-4.3 cm, with or without 1-2 white bands on dis- tal part. The upper legs have black, fine, non-annulated, 4.0 cm long hairs. Belly almost naked, the hairs vaguely banded or only with one whitish band at the base. Bris- tles on nape 8.5 cm, with one whitish band only. Proxi- mal bristles 8.0 cm, with one whitish narrow band only. Based on film and photo material of wild animals we es- timated the following average measurements for adults: total body length 127 cm; ear length 13 cm; shoulder height 85 cm. Body weight was not taken but according to local hunters ranges from 40-50 kg. Mean skull length 261 mm, mean mandible length 182 mm. For additional skull measurements of Pecari maximus n.sp., P. tajacu and Tayassu pecari and definitions of measurements see Table 1. Type Material. Holotype: INPA4272 (Mammal Collec- tion of the National Institute for Amazon Research, Man- aus, Amazonas, Brazil): complete cranium and mandible of an adult male (Fig. 3), killed for food by a local hunter, Luís Corréa Bastos, on March 12, 2003, along the left bank of the Rio Aripuana near the settlement of Arauazinho. Figure 2a shows the freshly hunted animal from which on- ly the skull could be saved that is here designed as the holotype. Paratypes: MR315, an incomplete skull with skin (Fig. 2b), obtained from hunters who had killed the animal in December 2003 along the Rio Arauazinho; MR316, a complete cranium and mandible collected from hunters living along the lower Rio Aripuanä, but lacking exact lo- cality data (both paratype specimens kept by the first au- thor). Type locality. Left bank of the Rio Aripuana, close to the settlement of Arauazinho, situated at the mouth of the Rio Arauazınho, a left bank tributary of the lower Rio Aripuana, State of Amazonas, Brazil (06°16'94’'S, 60220"87""W). Etymology. The name is Latin meaning “the largest’, re- ferring to the size of the new species that is the largest among living peccaries. The gender is masculine. Vernacular name. Pecari maximus 1s locally known as ‘caitetu-munde’. Locals claim that in the Tupi-Indian tongue (lingua geral) it means “the collared peccary that is bigger and goes in pairs”. This allows them to distin- guish it from Pecari tajacu commonly known as “caitetú- de-bando’ (“the collared peccary that goes in herds”). 3. PHYLOGENY Mitochondrial cytochrome b (THEIMER & KEIM 1998), control region and 12S r RNA sequences, and nuclear SINE PRE-1 sequences (GONGORA & MORAN 2005) in- dicate that Zavassu pecari and Catagonus wagneri are more closely related to each other than to Pecari tajacu. A complete mitochondrial D-loop sequence (1383 bp) and two nuclear SINE PRE-I sequences (P27, 322 bp and P642, 386 bp) (homologous sequences described in Su- LANDARI et al. [1997] and GONGORA & MORAN [2005]) from our skin sample (MR315) support the status of the new peccary species as belonging to the genus Pecari, closely related to Pecari tajacu. Sequences are deposited in GenBank under the accession numbers: DQ009006, DQ016371 and DQ016372. Net maximum likelihood (TN93) sequence divergence with Pecari tajacu is 2.0 % (Colombia, GenBank accession numbers: AY546522, AF276938) and 1.1 % (GenBank accession numbers: AY 26778-79, AY 546529/32, AB000378-79), respective- ly. For the D-loop it is of the same order as observed be- tween the European and Asian pig (1.7 %) (KIM et al. 2002), for which the divergence time has been estimated 110 Marc G. M. VAN ROOSMALEN et al.: A new species of Peccary from the Brazilian Amazon AS os % WZ os > a $ p 10 O ANO a om? 5 \ C; > S Ot 20 e o) x N) G 2 Va; a or ( y A or Fig. 5. e Pecari maximus en x A Rio Aripuana sightings Rio Xapuri o 75150 300 450 600 = a Kilometers N Le A Assumed distribution (shaded) of Pecari maximus sp. nov. in the Brazilian Amazon, based on sightings in the wild and interviews with locals. A question-mark indicates the Rio Xapuri where an American rubber cutter working in the area from 1906— 1916 described game that is similar to the giant peccary both in general appearance and behaviour (YUNGJOHANN 2003). . Borba 1. Arauazinho, type and paratype locality 2. Ilha das Pombas 3. Boca do Rio Juma 4. Prainha and Lago Cipotuba 5. Nova Olinda „ Novo Aripuanä Rio Anpuanä 0 40Km ee] Fig. 6. Collecting and sighting localities for Pecari maximus sp. nov. along the Rio Aripuana. at 860,000 years before present on account of near-com- plete mtDNA genome sequences (KIJAS & ANDERSSON 2001). This indicates a divergence time between Pecari tajacu and Pecari maximus of 1.0 million years before present. The combined SINE PRE-1 sequences indicate a similar divergence time of 1.2 million years before pres- ent assuming a mutation rate of 4.6x10-? per year per site (SULANDARI et al. 1997). Figure 4 shows a neighbour-join- ing tree including all known Tayassuidae species. 4. DISTRIBUTION, ECOLOGY AND CONSERVA- TION STATUS Distribution The larger geographical distribution of the giant peccary is thought to be the interfluve delineated by the Rio Madeira in the west, the Rio Tapajós-Juruena in the east, the Rio Amazonas in the north and the Rio Guaporé in the south (Fig. 5). Since Pecari maximus appears to be con- fined to terra firme rainforest habitat we assume that its real distribution is much smaller and does not extend in- to the northern part of the Rios Madeira/Tapajös interfluve, Bonner zoologische Beitráge 55 (2006) 111 where there are many open savannahs and extensive flood- plains. We have observed the species in the wild only along both banks of the Rio Arıpuanä (Fig. 6); 1t swims well (pers. obs.). The species might also occur in the up- per Rio Madeira, State of Acre (Fig. 5). This assumption is based on the story of an American, John C. Yungjohann, who worked as a rubber cutter in the Rio Xapurí area from 1906-1919, written down in the book “White Gold” (YUNGJOHANN 2003). He describes three types of “bush pigs”. The description of one of these closely resembles Pecari maximus: “there is a great, big one porcäo, they travel in pairs, and are very lively. They will attack on sight — either you have to be swift and a sure shot or climb a tree”. This behaviour — going in pairs and being really ag- gressive when attacked — is also reported by the hunters along the Rio Aripuana. Ecology Both Pecari tajacu and Tayassu pecari roam semi-nomad- ically in a highly variable landscape in noisy, widely spread herds of up to 30, respectively 200 individuals, which stay in permanent contact by loud tooth-clicking. In strong contrast, Pecari maximus seems to walk silent- ly through its preferred habitat — dense terra firme climax forest — in small family groups that contain only an adult pair with or without 1-2 offspring. Pecari maximus ap- pears to perform little or no uprooting, whereas the gre- garious peccary species largely forage for subsoil seed shadows, seedlings, roots and tubers. Instead, it has been seen feeding predominantly on freshly fallen fruits and seeds exposed on the forest floor. If this is its predomi- nant foraging technique, its different dietary specialisation might well explain the fact that among the several skull and mandible parts of the three local species combined, the molars of Pecari maximus invariably showed a less dramatic wearing pattern. This could reflect its much smaller intake of sand and dirt during feeding. It appears that the larger Pecari maximus has abandoned social groupings, group defence and territorial scent marking. Its scent gland is thought to be rudimentary. None of the skins examined emitted the typical peccary scent, suggesting that if any secretory liquid is produced it is scentless, at least for the human nose. Conservation status All three peccary species occurring sympatrically in the Rio Aripuana region are the favourite game of the locals, but only Pecari maximus is hunted with dogs since it does not go in herds like the other peccaries which are known to defend themselves fiercely. Although human occupa- tion in this part of the Amazon is presently very low, this situation might soon change. In the Rio Aripuana region unprecedented illegal extraction of timber and gravel is taking place. Recent road building through the area is in- tended to connect the town of Manicoré on the right bank of the Rio Madeira with the boomtown of Apui at the bor- der of the Tenharim Savannah and the State of Mato Grosso, areas of large-scale soybean agriculture. In view of these recent developments, we fear that commercial hunters using trained dogs will focus first on Pecari max- imus to feed hungry settlers. Taking increasing hunting pressure and the species’ limited distribution into account, we consider Pecari maximus endangered. We recommend inclusion of this new species in the IUCN Global Red List, based on criterion D (very small or restricted population). Besides the giant peccary, the Rio Aripuana region is thought to harbour a number of floral and faunal elements new to science. The first author has identified so far a new species of dwarf porcupine, Coendu roosmalenorum Voss & da Silva, 2001 (Voss & DA SILVA 2001), and seven new primate species, four of which are already described (VAN ROOSMALEN et al. 1998; VAN ROOSMALEN et al. 2000; VAN ROOSMALEN et al. 2002; VAN ROOSMALEN & VAN ROOS- MALEN 2003). Among these primates, the dwarf marmoset Callibella humilis, represents a new genus never collect- ed before. Most surprisingly, not a single area protected by Brazilian environmental law exists in the region. Giv- en the uniqueness of the region in terms of biodiversity and its current status of biological terra incognita, we here encourage UNESCO to urge the Brazilian government to declare the entire region a World Heritage Site. Acknowledgements. We thank Jórn Róver and Tom Syn- natzschke from NDR (Norddeutscher Rundfunk) Natur- film, Hamburg, Germany, who made two expeditions to the Aripuana region possible in order to document new plant and animal species, and Roland Gockel and Frieder Salm, who accompanied the first two authors in the field for a total of three months and filmed and photographed giant peccaries in the wild for NDR. REFERENCES BODMER, R. E. & Sow ts, L. K. 1993. The collared peccary (Tayassu tajacu): status and action plan summary. Pp. 7-13 in: OLIVER, W. L. R. (ed.) Pigs, Peccaries and Hippos. Status Survey and Conservation Action Plan. IUCN, Gland. GONGORA, J. & MORAN, C. 2005. Nuclear and mitochondrial evolutionary analyses of Collared, White-lipped, and Chacoan peccaries (Tayassuidae). Molecular Phylogenetics and Evolution 34: 181-189. GRUBB, P. 2005. Family Tayassuidae. Pp. 643-644 in: WILSON, D. E. & REEDER, D. M. (eds.) Mammal Species of the World. John Hopkins University Press, Baltimore. 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The white-lipped peccary (Tayassu pecari): status and action plan summary. Pp. 13-22 in: OLIVER, W. L. R. (ed.) Pigs, Peccaries and Hippos. Status Survey and Conservation Action Plan. IUCN, Gland. SULANDARI, S., HARUMI, T., WADA, Y. & YASUE, H., MULADNO & YANAI, S. 1997. Localization of swine PRE-1 homologues in 13 loci of Phacochoerus aethiopicus and Tayassu tajacu genomes, and their sequence divergence. Animal Genetics 28: 210-215. THEIMER, T. C. & KEim, P. 1998. Phylogenetic relationships of peccaries based on mitochondrial cytochrome b DNA sequen- ces. Journal of Mammalogy 79: 566-572. VAN ROOSMALEN, M. G. M., VAN ROOSMALEN, T., MITTERMEIER, R. A. & DE FONSECA, G. A. B. 1998. A new and distinctive species of marmoset (Callitrichidae, Primates) from the lower Rio Aripuana, State of Amazonas, central Brazilian Amazo- nia. Goeldiana Zoologia 22: 1-27. VAN ROOSMALEN, M. G. M., VAN ROOSMALEN, T., MITTERMEIER, R.A & RYLANDS, A. B. 2000. 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The Chacoan peccary, Catagonus wagne- ri (Rusconi). Bulletin of Carnegie Museum of Natural Histo- ry 3: 1-36. WETZEL, R. M., DuBos, R. E., MARTIN, R. L. & Myers, P. 1975. Catagonus, an ‘extinct’ peccary, alive in Paraguay. Science 189: 379-381. WOODBURNE, M. O. 1968. The cranial myology and osteology of Dicotyles tajacu, the collared peccary, and its bearing on classification. Memoirs of the Southern California Academy of Sciences 7: 1-48. WRIGHT, B. 1989. Phylogenetic relationships of Catagonus wag- neri: sister taxa from the Tertiary D of North America. Pp. 281-308 in: ReDFORD, K. H. & EISENBERG, J. F. (eds.) Ad- vances in Neotropical Mammalogy. Sandhill Crane Press, Gainesville. YUNGJOHANN, J. C. 2003. White Gold: The Diary of a Rubber Cutter in the Amazon 1906-1916. Synergetic Press, Santa Fe. Authors’ addresses. Dr. M. G. M. VAN ROOSMALEN, Amazon Association for the Preservation of High- Biodivers Areas (AAPA), Alameda B, QD G, c.09, Conjunto Álvaro Neves, Dom Pedro, CEP 69.042-000 Manaus-AM, Brazil: E-mail: garciaroosmalen@- hotmail.com or RoosmalenGarcia(anetscape.net; L. FRENZ, Heussweg 72, 20255 Hamburg, Germany; E-mail: Lofrenz@aol.com; Dr. P. van Hoort, Wageningen University and Research Centre, Resource Ecology Group, Bornsesteeg 69, 6708PD Wageningen, The Netherlands; E-mail: Pim.vanHooft@wur.nl; Dr. H. DE IONGH, Leiden University, Institute of Environmental Sciences, PO Box 9518, 2300RA Leiden, -The Netherlands; E-mail: Delongh@cs.com; Prof. Dr. H. LEIRS, University of Antwerp, Evolutionary Biology Group, Groenenborgerlaan 171, B-2020 Antwerpen, Belgium; E-mail: herwig.leirs@ua.ac.be Submitted: 27.03.2006 Revised: 15.09.2006 Accepted: 16.10.2006 Corresponding editor: G. Peters Bonner zoologische Beiträge Band 55 (2006) Heft 2 Seiten 113-121 Bonn, Juli 2007 A new species of the Creatonotos transiens-group (Lepidoptera: Arctiidae) from Sulawesi, Indonesia Vladimir V. DUBATOLOV* & Jeremy D. HOLLOWAY** * Siberian Zoological Museum of the Institute of Animal Systematics and Ecology, Novosibirsk, Russia ** Department of Entomology, The Natural History Museum, London, U.K. Abstract. A new species from Sulawesi (Celebes), Creatonotos kishidai Dubatolov & Holloway is described. It is cha- racterized by the presence of a sclerotized spine-bearing band which envelops the aedeagus apex, presence of spining on the juxta, and small peniculi (finger-like processes) at the bases of the transtilla, which are longer than in the widespre- ad Oriental C. transiens (Walker, 1855) and shorter than in C. wilemani Rothschild, 1933 from the Philippines. The ho- lotype of the new species is deposited in the Siberian Zoological Museum of the Institute of Animal Systematics and Ecology, Novosibirsk, Russia. Study of a topotype of C. transiens vacillans (Walker, 1855) showed that it is a senior syn- onym of C. t. orientalis Nakamura, 1976. C. ananthakrishanani Kirti et Kaleka, 1999 is synonymized with the nomino- typical subspecies of C. transiens (Walker, 1855). The lectotype of C. transiens (Walker, 1855) is designated in the BMNH collection from “N. India, Kmorah” (misspelling of “Almorah”). Keywords. Tiger-moth, Arctiidae, Arctiinae, Creatonotos, Creatonotos transiens-group, new species, Sulawesi, Indo- nesia. 1. INTRODUCTION The genus Creatonotos Hübner, [1819] 1816 consists of a set of unrevised A frotropical species (GOODGER & WAT- SON 1995) and seven species from South Asia and neigh- bouring territories. Two species are distributed through- out the whole Oriental region: C. transiens (Walker, 1855) and C. gangis (Linnaeus, 1763), the latter reaches its lim- its in Northern Australia, South Iran and North-Eastern Arabia (1 9, 1 9, United Arab Emirates, | km W Marbad, 25°20’N 56°07’E, 28.111 — 5.IV 2006, leg. M. Fibiger & C. Gielis, ex Zoological Institute (St.-Petersburg, Russia) collection). Two species are much more localized within the Oriental Region: C. fasciatus (Candeze, 1927) was de- scribed from Cambodia, C. wilemani (Rothschild, 1933) inhabits the Philippines. To these must now be added €. kishidai sp. n. from Sulawesi. Two species are known from Western Arabia: C. albidior Wiltshire, 1986 and C. leu- canioides Holland, 1893. C. arabicum (Hampson, 1896) from South-Western Asia has been assigned to a new genus Creataloum Dubatolov, 2004. The Creatonotos species from Asia fall into two main groups defined by the forewing pattern and male genitalia structure; these could be treated as subgenera. Species of the nominotyp- ical subgenus have light wings with a dark streak along the posterior vein of the cell, and have long needle-like cornuti on the vesica: C. gangis, C. albidior, C. leucan- ioides and, taking into account the forewing pattern on- ly, C. fasciatus. The C. transiens group might be referred to subgenus Phissama Moore, 1860, the species having forewings without a streak along the posterior vein of the cell, and having short, drawing-pin-like cornuti on the vesica that are sometimes fused into bands. 2. TAXONOMY NAKAMURA (1976) reviewed the latter group and recog- nised two species. On the basis of male genital structure, he divided C. transiens (Walker, 1855) into four sub- species, and described a sibling species, C. philippinense Nakamura, 1976, from the Philippines. HOLLOwAY (1988) and INOUE (1988) independently found an older name for the latter, C. wilemani Rothschild, 1933, which was orig- inally described as a distinct species. HOLLOWAY (1988) also found that there was a further taxon of this group on the Indonesian island of Sulawesi, but did not describe it at the time. After a careful study of Creatonotos specimens kindly sent by Mr Y. Kishida to the collection of Siber- ian Zoological Museum of the Institute of Animal System- atics and Ecology, Novosibirsk, Russia (SZMN), it became apparent that the new taxon from Sulawesi has several striking characters of the male genitalia that distinguish it from both €. transiens and C. wilemani. It should there- fore be described as a new species. 114 Vladimir V. DUBATOLOV & Jeremy D. HOLLOWAY: A new species of the Creatonotos transiens-group from Sulawesi However, a female specimen from Sulawesi was includ- ed in the type series of C. transiens, along with other fe- males from North India, Sylhet, and East Indies. A lecto- type of C. transiens has yet to be designated. HAMPSON (1901) treated a specimen from Sylhet as “type” of C. transiens, but neither syntype with these data (from the Sowerby collection) was rediscovered during our inves- tigations. So, there are two female syntype specimens from North India, Almorah, from the Stevens collection, and they belong to the nominotypical C. transiens subspecies sensu NAKAMURA (1976). Therefore, in the interests of sta- bility, one of these females from North India, Almorah (handwritten on the label as “N. India, Kmorah”), is here- by designated as the LECTOTYPE (Fig. 2). Similarly, the type series of Aloa isabellina Walker, 1855 included male and female specimens from Sylhet, North India and East Indies. These localities, taken at face value, are shared with transiens, but the concept of “East Indies’ is very impre- cise and may involve parts of southeast Asia and Indone- sia where other races occur; in fact, the type material so labelled for both taxa was from the Archdeacon Clerk col- lection, and the one syntype of transiens located from this source is labelled as from Moulmein in southern Burma (Myanmar). Therefore, in the interests of stability, we des- so] O-AFGHANISTAK SE] Sarghicttog m ES ei = 3 leg:6 Ebert Figs 1-9. ignate the Sylhet male specimen of isabellina as LECTO- TYPE. It is possible that North India and East Indies syn- types of transiens were also listed as syntypes of isabel- lina, given that only six syntypes of the taxa together have been located. The discovery of the new species from Sulawesi led us to review geographic variation in male genitalia for the whole C. transiens group. For this, we use the terminol- ogy of NAKAMURA (1976) for the ornamentation of the aedeagus vesica. First of all, it was found that the male genitalia of a C. r. vacillans (Walker, 1855) topotype specimen from Hong Kong do not differ from those of C. £. orientalis Naka- mura, 1976 from Thailand in the structure of the fused cor- nuti plate M1-M2-M3-M4-B of NAKAMURA (1976), the shape of which can vary significantly even in geograph- ically close populations; it differs from C. £. koni Miyake, 1909 from Taiwan and South Japan, which has the cor- nuti plate disrupted into several cornuti M1, M2, M3, M4. Moreover, two continental subspecies, the nominotypical C. 1. transiens and C. t. vacillans, differ only slightly from each other, the former from the Himalayas and India hav- Creatonotos, adult moths. 1. C. transiens albina, holotype, Afghanistan, Sarobi, 1100 m, 11.X 1961, G. Ebert leg. 2. C. transiens, lectotype and its labels, “N. India, Kmorah”. 3. C. £. transiens, India, E Sikkim, 20 km SE of Ganghtok, Phidim Rain Forest, h = 1600 m, VII 2002, O. Yu. Ammosov leg. 4. C. £. vacillans, Vietnam, Cao Bang, Mt. Pia Oak, h = 1700 m, III 2002, native collector leg. 5. C. 1. sundana, Indonesia, Bali, Tamblingan, VI 2004, native collector leg. 6. C. £. koni, Japan, Okinawa, Na- ha, P. Schmidt leg. 7. C. wilemani, Philippines, Negros Is., Mt. Canlaon, native collector leg. 8-9. C. kishidai, Indonesia, Sulawe- si, Tondano, VII 1988, native collector leg. 8. Holotype. 9. Paratype. 2. Female. 1, 3-8. Males. Bonner zoologische Beiträge 55 (2006) 115 10 11 E 5 j a oa 3 UE j / 14 we » m _ Figs 10-15. Male genitalia of Creatonotos. 10-12. C. t. transiens, India, E Sikkim, 20 km SE of Ganghtok, Phidim Rain Forest, h=1600 m, VII 2002, O. Yu. Ammosov leg. 13-15. C. t. albina, Afghanistan, Sarobi, 14.X 1961, G. Ebert leg. 10, 13. General view. 11, 14. Aedeagus, left side. 12, 15. Aedeagus, right side. ing a more uneven cornuti plate M1-M2-M3-M4-B. So, the list of the C. transiens-group taxa is as follows: Creatonotos transiens (Walker, 1855) (Figs 1-6) South-East Asia from India to China, South Japan, Bor- neo and Lombok. Records of C. transiens from the Philip- pines in HOLLOWAY (1976, 1988) are likely to have aris- en from misplaced specimens of C. wilemani in the se- ries of C. transiens in BMNH; these were recognised as misidentified subsequently, and one is the Philippines specimen included amongst those listed by WALKER (1856) in his redescription of vacillans (see below). A fur- ther search of supplementary drawers has not yielded any Philippines specimens that are definitely C. transiens, so this record must be treated as suspect. 116 Vladimir V. DuBATOLOV & Jeremy D. HOLLOWAY: A new species of the Creatonotos transiens-group from Sulawesi 19,20 i mm i = E 21 Figs 16-21. Male genitalia of Creatonotos. 16-18. C. 1. vacillans, Hong Kong, Kan Sui Chan, 31.XII 2005, J. Lee leg. 19-21. C. ¢. vacillans, Thailand, Chanthaburi Province, Khao-Khitehakut National Park, by light at headwaters, 6.1 2006, O. E. Kosterin leg. 16, 19. General view. 17, 20. Aedeagus, left side. 18, 21. Aedeagus, right side. C. t. albina (Daniel, 1971); Ann. Naturh. Mus. Wien 75: 654, t. 1, fig. 5 (Phissana transiens albina); type locali- ty: “[O-Afghanistan] ... Sarobi” [Paktika: Sarowbi] (Fig. 1): Afghanistan: Paktika, Nangarhar, Nuristan, Badakhshan (DANIEL 1971). Remarks. According to the original description (DANIEL 1971) and the figure of the holotype (Fig. 1), this sub- species has the palest whitish-grey wings in males (forewing being noticeably lighter than hindwings) and white wings in females. Male genitalia were stated by Daniel to be identical with the nominotypical subspecies; however, they are only similar (Figs 13-15). Cornuti patch M1-M2-M3-M4-B of right side of vesica is massive, reg- ular in shape; cornuti patch O is large, like in the nomino- typical subspecies. C. £. transiens (Walker, 1855); List Specimens lepid. In- sects Colln. Br. Mus. 3: 675 (Spilosoma transiens); type locality: North India, Almorah, designated here from the lectotype (Figs 2-3). Bonner zoologische Beitráge 55 (2006) 117 25 26 | a on es: So a 24 wu T 27 Figs 22-27. Male genitalia of Creatonotos. 22-24. C. t. sundana, Indonesia, Mentawai Isl., Siberut Isl., Bojakan, IX 2004, na- tive collector leg. 25-27. C. 1. koni, Japan, Okinawa, Taha, P. Schmidt leg. 22, 25. General view. 23, 26. Aedeagus, left side. 24, 27. Aedeagus, right side. =Aloa isabellina Walker, 1855; List Specimens lepid. In- sects Colln. Br. Mus. 3: 705—706; type locality: Sylhet, designated here from the lectotype. =Creatonotos ananthakrishanani Kirti et Kaleka, 1999, syn. nov.; Entomon 24 (2): 137-140, figs 1-11; type lo- cality: “Uttar Pradesh: Mussorrie”. India, North Pakistan (a photo of a male specimen from the Indus Canyon, 50 km E of Peshawar in Mr. V. Gurko’s collection (Tchernovtsy, Ukraine) has been studied). Remarks. C. ananthakrishanani Kırti et Kaleka, 1999 was described from the North-West Himalayas, Mussorrie, on the basis of differences in wing venation from C. transiens and misinterpreted minute differences in male genitalia. Moreover, the authors mistakenly considered there to be two cornuti groups in C. transiens and three groups in C. ananthakrishanani (Fig. 36), but they never everted vesi- cas in both taxa. Presence of three cornuti groups is char- acteristic for the whole C. transiens-complex. The specif- ic status of C. ananthakrishanani is therefore unjustified. The nominotypical subspecies vary from the darkest 118 Vladimir V. DUBATOLOV & Jeremy D. HoLLoway: A new species of the Creatonotos transiens-group from Sulawesi 31 1 mm Figs 28-34. Male genitalia of Creatonotos. 28-30. C. wilemani, Philippines, Negros Is., Mt. Canlaon, native collector leg. 31— 33. C. kishidai, holotype, Indonesia, Sulawesi, Tondano, VII 1988, native collector leg. 34. — C. kishidai, paratype, Indonesia, Ce- lebes [Sulawesi], Loda, Paloe, 4000”, V 1937, J. P. A. Kalis (Zoologisches Forschungsinstitut und Museum Alexander Koenig, Bonn, Germany). 28, 31. General view. 29, 32, 34. Aedeagus, left side. 30, 33. Aedeagus, right side. brown forewings in the specimens from Sikkim to the pale brown forewings in the specimens from North India and North Pakistan, whilst in male genitalia (Figs 10-12, 35: 2) it ıs similar to the next subspecies, differing in the ir- regular shape of elongate cornuti patch M1-M2-M3-M4- B of right side of vesica; cornuti patch O is large. C. t. vacillans (Walker, 1855); List Specimens lepid. In- sects Colln. Br. Mus. 3: 685 (Amphissa vacillans); type locality: “Hong Kong” (Fig. 4). =Creatonotos transiens orientalis Nakamura, 1976, syn. nov., Tyó to Ga 27 (3): 116, fig. 1 (3), 3 (5); type locali- ty: “Fang, Thai” [Thailand]. China: Heilongjiang (Tili, Pingdin Mt., SZMN; accord- ing to the general species distribution, this outlying north- ern locality needs confirmation), Shanxi, Shaanxi, Shan- dong, Anhui, Henan, Jiangsu, Zhejiang, Fujiang, Jiangxi, Hubei, Hunan, Guangdong, Hainan, Guanxi, Guizhou, Sichuan, Yunnan (FANG 2000), East and South Tibet (FANG Bonner zoologische Beiträge 55 (2006) 119 1982, 1987); Indochina, Peninsular Malaysia, Sumatra, Borneo. Remarks. This subspecies is paler than the nominotypi- cal one (taking into account specimens from Sikkim), cor- nuti patch M1-M2-M3-M4-B on the vesica right side is not massive, equal in width, but varies ın length, cornuti patch O is also large (Figs 16-21, 35: 5). The original de- scription of vacillans was based on a single (therefore holotype) male. However, WALKER (1856: 1702) later de- scribed Aloa vacillans, referring to three specimens: a “typical” female from N. India; var. beta, a male of wile- mani from the Philippines; var. gamma, the holotype Hong Kong male of the original vacillans. This may be a fur- ther indication that Walker did on occasion “recycle” his type material! C. t. sundana Nakamura, 1976, Tyó to Ga 27 (3): 116, fig. 1 (2), 3 (4); type locality: “Mégamenden, Java” (Fig. 5). Indonesia: Siberut (Bojakan, SZMN), Java, Bali (Tamblin- gan, SZMN), east to Lombok. Remark. This island subspecies does not differ from the typical subspecies in wing colouration, but it has reduced cornuti patches M1-M2-M3-M4-B and O on vesica (Figs 22-24, 35: 4). C. t. koni Miyake, 1909, stat. rev.; Bull. Coll. Agric. Univ. Tokyo 8 (2): 169 (Creatonotus Koni); type locality: “Mt. Arisan in Formosa” (Fig. 6). Taiwan, Japan: Yonagumi, Iriomote, Ishigaki, Miyako, Okinawa, Okinoerabu, Amami, Yaku (NAKAMURA 1976; INOUE 1982). Remarks. According to NAKAMURA (1976) this subspecies is characterized by the very light rosy tone of head, pata- gia, tegulae, thorax and forewings; cornuti patch M1-M2- M3-M4 is divided into separate small cornuti groups, while cornuti patch O is not reduced (Figs 25-27, 35: 3). Creatonotos wilemani (Rothschild, 1933) Am. & Mag. Nat Hist. (10) 11: 183-184 (Creatonotus wilemani), type locality: “Palali, 2000 ft., Benguet, Klondyke, 800 ft., Benguet, Luzon, ..., Kolambugan, Lanao Plains, Mindanao, ...; Luzon...” (Fig. 7). =Creatonotos philippinensis Nakamura, 1976, Tyo to Ga 27 (3): 113-114, fig. 1 (4), 3 (6); type locality: “Subasta, Mindanao Is.” Philippines: from Luzon to Mindanao. Remarks. The species is characterized by absence of black dots at forewing discocellular veins, presence of the largest peniculi or finger-like processes at base of the transtilla, which is wide and the most strongly sclerotized in the whole group (Figs 28-30, 35: 6). Creatonotos kishidai Dubatolov & Holloway, sp. nov. (Figs 8-9) Material. Holotype — male, Indonesia, Sulawesi, Tondano, VII 1988, native collector leg., received from Y. Kishida. Preserved in SZMN coll. (http://szmn.sbras.ru). Paratypes: 1 9, the same data (SZMN); 2 ' (BMNH genitalia slides 2313, 2314), 1 2 (BMNH slide 2311), Koelawi, Paloe [Palu], 3100”, W. Celebes, March 1937, J. P. A. Kalis (BMNH); 1 & (BMNH slide 2312), Loda, Paloe [Palu], W. Celebes, 4000”, May 1937, J. P. A. Kalis (BMNH); 3 JO, the same data (Zoologisches Forschungsinsmuseum Alexander Koenig, Bonn, Germany); 1 9 (syntype of tran- siens), Celebes [Menado], [Madame Ida] Pfeifer coll. (BMNH). There are large numbers of specimens of C. kishidai in BMNH from the localities listed for the paratypes, but we have restricted these to dissected spec- imens and to the syntype of transiens. Description. Male forewing length 21.0-22.5 mm, wing expanse 41.0-46 mm. Wings dark grey, the forewing with lighter costal margin and veins on the external part of the wing. There are three or four black dots around the dis- cocellular veins, two inside the cell being larger. Head and thorax whitish, abdomen dorsally yellow with one dorsal and two lateral rows of black spots; ventrally 1t is grey. Male genitalia (Figs 31-34). Uncus elongate, horn-like. Valvae narrow, elongate, slightly curved inwards, with two distal processes: the apex and a lateral spur. Juxta longer than wide, its apical lateral sides covered with small spines. Transtilla at both bases with distinct finger-like processes (peniculi). Aedeagus straight, widened at apex, where it is enveloped by a sclerotized spine-bearing band (hypertrophied cornuti patch O); this band is sometimes disrupted ventrally (Fig. 34). Vesica with two elongate sclerotized cornuti patches M and B, between them there is one more small round sclerotization B on the opposite side of the vesica. Female forewing. Length 22.0-23.0 mm, wing expanse 43.0-49.0 mm, the syntype of transiens being at the up- per boundary of this range. Forewings light grey with lighter veins in external part of the wing. There are four black dots around the discocellular veins, two inside the cell being larger; the external anterior one is the weakest. Hindwings whitish with dark scales on veins; those on the cubital stem are darkest. 120 Vladimir V. DUBATOLOV & Jeremy D. HOLLOWAY: A new species of the Creatonotos transiens-group from Sulawesi 3» IN Figs 35-36. 35. Vesica structure of the Creatonotos species, by Nakamura (1976). 1. C. gangis, 2. C. t. transiens, 3. C. t. koni, 4. C. t. sundana, 5. C. t. orientalis ( = vacillans), 6. C. t. philippinense ( = wilemani). O, M1, M2, M3, M4, B — cornuti patches. 36. Aedeagus structure (vesica only slightly everted) of C. ananthakrishanani, from original description by Kirti & KALEKA (1999). AED — aedeagus, CRN — cornuti, VES — vesica. Remarks. The presence of a sclerotized spine-bearing band, which envelops the aedeagus apex, distinguishes the new species from both C. transiens and C. wilemani, where it is absent. Moreover, these two species lack the spining of the juxta characteristic of the new species. The presence of small peniculi (finger-like processes) at the bases of the transtilla is shared with C. wilemani (Figs 28, 31), though this species has them larger on a larger and more sclerotized transtilla. The valves also differ across the three species, extending beyond the uncus in €. tran- siens, but falling short of it in C. kishidai and C. wilemani; the lateral spur 1s short and in the most basal position in C. transiens, but nearer the apex of the valve and longer in C. kishidai, where distinct bilateral asymmetry occurs, the right valve being longer, with the apical process longer beyond the lateral spur. In €. wilemani the lateral spur is small and very close to the apex. Acknowledgements. Authors are grateful to Mr. Yasunori Kishi- da (Tokyo, Japan) for important material of Creatonotos species, to Mr. M.R. Honey (London, U.K.) for much help with discovering type material in the BMNH collection, to Mr. J. Lee (Hong Kong) for the topotype male specimen of C. ¢. vacillans, to Mr. V. O. Gurko (Tshernovtsy, Ukraine), Dr. D. V. Logunov (Manchester Museum, Manchester, England) and Dr. O. E. Kos- terin (Novosibirsk, Russia) for additional material of C. tran- siens, to Dr. S. Yu. Sinev and Dr. A. Yu. Matov (St.-Petersburg, Russia) for a loan of the C .1 .koni male specimen from the Zo- ological Institute collection, to Dr. W. Speidel, (Witt collection, Munich, Germany) and Mr. U. Buchsbaum (Munich, Germany) for the photograph of the holotype of C. 1. albina, deposited in the Zoological State’s Collection of Bavaria, Munich, Germany and a loan of two paratypes of C. £. albina from the ZSM, to Dr. D. 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Arctiidae. 1: 638-659; 2: 74-79, 136-137, 334-342, pl. 154-162, 346-348 in: INOUE, H., Sut, S., Kuroko, H., MORIUTI, S., KAWABE, A. Moths of Japan 1: 1-967, 2: 1-554, pl. 1-392. Kodansha, Tokyo. INOUE, H. 1988. Three new species and some synonymic notes on the Arctiidae from Japan, Taiwan and the Philippines. Tyó to Ga 39 (2): 99-118. Kirti, J. S. & KALEKA, A. S. 1999. Reporting of a new species under genus Creatonotos Húbner (Arctiinae: Arctiidae: Lep- idoptera) from India. Entomon 24 (2): 135-141. MIYAKE, T. 1909. A revision of the Arctianae of Japan. Bulletin of College of Agriculture, Tokyo Imperial University 8: 153-174. NAKAMURA, M. 1976. Notes on the geographical variations of Creatonotos transiens Walker with descriptions of new species and two new subspecies (Lepidoptera: Arctiidae). Tyó to Ga 27 (3): 111-117. ROTHSCHILD, L. 1933. New species and subspecies of Arctiinae. The Annals and Magazine of Natural History (10) 11: 167-194. WALKER, F. 1855. List of the specimens of lepidopterous insects in the collection of the British Museum 3: 583-775. London. WALKER, F. 1856. List of the specimens of lepidopterous insects in the collection of the British Museum 7: 1509-1808. Lon- don. Author”s addresses: Vladimir V. Dubatolov (correspon- ding author), Siberian Zoological Museum of the Institute of Animal Systematics and Ecology, SB RAS, Frunze street, 11, Novosibirsk, 91, 630091, Russia. E-Mail: vvdu- batwonline.nsk.su Jeremy D. Holloway, Department of Entomology, The Natural History Museum, Cromwell Road, London SW7 SBD, U.K. E-Mail: j.holloway(@nhn.ac.uk. Received: 21.11.2006 Revised and accepted: 18.05.2007 Corresponding editor: D. Stiining “ =, Bonner zoologische Beiträge Band 55 (2006) Heft 2 Seiten 123-150 i Bonn, Juli 2007 Herpetofauna Kakamegensis — The amphibians and reptiles of Kakamega Forest, western Kenya / Philipp WAGNER & Wolfgang BOHME Bonn, Germany Abstract. We present an annotated checklist of the herpetofauna of Kakamega Forest with comments on the biology and systematics of the taxa. Twenty-five amphibian, one turtle, 22 lizard and 36 snake species are recorded from within the forest and its immediate environment. We discuss the generalized zoogeography of the forest and distribution pattern of the taxa comment on the protection of the forest. Analysis of the reptile species composition shows Kakamega Forest to be similar to the Guinea-Congolian rainforest and is considered the easternmost remnant of this forest block. Kakamega forest has a high diversity value for Kenya and represents a diversity hotspot on a national scale. Two species, Lygodac- tvlus gutturalis and Psammophis phillipsi, are recorded in Kenya for the first time. Several other first records and the description of a new species (Agamidae: Agama finchi) were published already separately. Keywords. East Africa, Kenya, Kakamega Forest, herpetological survey, checklist, national diversity hotspot. 1. INTRODUCTION In Africa tropical rainforests extend from southern Sene- gal in the west to the coastal forests of Kenya and Tanza- nia in the east (COLLINS 1992). The East African rainforests belong to different biogeographical clades. Apart from the mountain forests and inselbergs, which are not clearly as- signable, there are three important forest types: the coastal forests and the Eastern Arc mountain forests, both with a high degree of endemism, and the easternmost outliers of the Guinea-Congolian rain forest. The Eastern Arc Moun- tains were already the object of several biological stud- ies and have recently been reviewed in respect to their her- petofauna by HOWELL (1993). The value of the coastal for- est biodiversity was underestimated for a long time. Re- cently the herpetofauna of the Kenyan part of the coastal forest was surveyed by the Kifcon project (KIFCON 1995). Only the forests associated with the Guinea-Congolian for- est (e. g. Budongo, Bwamba, Kibale, Bwindi, Mbira and Mt. Elgon in Uganda) are at least partly explored. Her- petological inventories have been made for the Bwindi (DREWES & VINDUM 1991) and Kibale Forests (VONESH 2001). All together these forests harbour an estimated number of 333 amphibian (DUELLMAN 1993), 105 snake (HUGHES 1983), 95 lizard, 16 turtle and 3 crocodilian species (BAUER 1993). This total herpetofaunal species richness is more than 550. Considering the fact that zoo- logical research in Africa is becoming more and more dif- ficult because of the political instabilities in numerous countries, this number is certainly too low. Because trop- ical forests in Africa are seriously threatened numerous undescribed species may become extinct before their dis covery and scientific descriptions. Uganda, for example has lost 86 % of its original forest in the past two decades and the remaining parts are isolated fragments (VONESH 2001). Kakamega Forest in western Kenya is a similar isolated fragment. This small forest is the easternmost fragment of the equatorial rain forest system (CLAUSNITZER 2005; DREWES 1976; HAMILTON 1976; KOHLER 2004; KOKWARO 1988; ScHioTz 1976; VONESH 2001; WAGNER et al. sub- mitted; ZIMMERMANN 1972) and has recently been sur- veyed in regard to its faunal communities by the ‘BIOTA East Africa Project’ and especially to its amphibian fau- na by e. g. SCHICK et al. (2005) and LOTTERs et al. (2006). The herpetogeographical relationships to other fragments of the equatorial rain forest and other tropical forests have recently been discussed by WAGNER et al. (subm.). Despite belonging to Guinea-Congolian forest, Kakamega Forest contains also numerous Afromontane elements (MUTAN- GAH et al. 1992; BENNUN & NJOROGE 1999) in its flora and fauna. Therefore, the forest has a very large diversity and zoogeographical value which has been shown by several authors for the different species groups (e. g. Odonata: CLAUSNITZER 1999 & 2005; Amphibia: SCHICK et al. 2005; Reptilia: this paper; Aves: ZIMMERMANN 1972) and a lot of species are not found elsewhere in Kenya. Additional- ly, most of the remaining closed canopy forest within the country is found in western Kenya (Wass 1995). On a na- tional scale, the forest can be considered as a diversity hotspot and needs efficient protection. On the pan-African 124 Philipp WAGNER & Wolfgang BÖHME: The amphibians and reptiles of Kakamega Forest, western Kenya Fig. 1. The main areas of investigation of BIOTA East Africa in the Kakamega Forest in the background a subset of a Land- sat 7 (ETM+) scene from 5 Feb 2001, contrast-enhanced band combination 5/4/3 for within-forest differentiation but printed in black and white. (Courtesy to G. SCHAAB of BIOTA — E02). scale, Kakamega forest 1s not considered as a hot-spot be- cause in comparison with e. g. the Mt. Nlonako in Cameroon (99 amphibian [HERRMANN et al. 2005a] and 89 reptile species, [HERRMANN et al. 2005b]) the diversi- ty 1s comparatively low. Compared with other vertebrate groups, East African am- phibians and reptiles are rather poorly studied and insuf- ficiently known. In order to provide conservationists da- ta for defining priorities for conservation It is necessary to obtain basic information on the diversity and commu- nity of forest amphibians and reptiles. Amphibians have been intensively studied by several au- thors within the ‘Biota East Africa Project’ (e. g. SCHICK et al. 2005; LOTTERS et al. 2004; LOTTERS et al. 2006; KOH- LER et al. 2006) whereas reptiles have been surveyed on- ly superficially in the Kaimosi fragment by LOVERIDGE (1935, 1936) and in the main forest by DREWES (1976). Subsequently no further reports on Kakamega reptiles have been published apart from the mentioning of single voucher specimens as e. g. in SPAWLS et al. (2002) and publications arising from this study (BOHME et al. 2005; KOHLER et al. 2004; WAGNER & SCHMITZ 2006; WAGNER et al. subm.). The aim of the present paper is to provide an overview of the herpetofauna of this forest as basic information for conservationists and wildlife biologists and to highlight the importance of the Kakamega Forest because of its im- pact and value on the biodiversity of Kenya. 2. DESCRIPTION OF THE STUDY AREA Kakamega Forest is situated in the Kakamega District near Kakamega town in the Western Province of Kenya. The forest extends from 0%10” and 0°21 N to 34%47” and 34°58’E, covering an area of 240 km?, of which only 44.55 km? are protected by law (MITCHELL 2004). The forest altitude varies between 1500 and 1700 m a.s.l. (above sea level) averaging 1650 m a.s.l. The forest be- comes part of the stratified landscape of the East African Rift Valley, situated 150 km to the east. The annual pre- cipitation ranges from 1500 to 2300 mm. The annual av- erage temperature is 27 °C at daytime and 15 °C at night. Two important rivers traverse the forest: the Isiukhu Riv- er in the north and the Yala River in the south. Both have their source in the Nandi Escarpment and drain into the nearby Lake Victoria. The forest block itself is surround- ed by several forest fragments (e. g. Kisere, Malaba, Kaimosi), which differ in size, in the degree of destruc- tion and their conservation status. The most important of these are the Kisere in the north and the Kaimosi fragment in the south. Kisere is protected as a National Reserve whereas Kaimosi is unprotected but well known histori- cally from several collections made by A. LOVERIDGE and from the three herpetological taxa described by him from Kaimosi and named after this forest: Agama kaimosae, Ty- phlops kaimosae and Dendroaspis jamesoni kaimosae. The eastern border of the forest is the 2200 m high Nan- di Escarpment with its Northern Nandi and Southern Nan- di forest. Both are considered to be montane forests be- cause of the occurrence of the tree fern Cyathea manni- ana as an indicator species of this forest type. The forests were contiguous with the Kakamega forest system until recently. MITCHELL (2004) pointed out that the North Nan- di forest was not connected with Kakamega forest in the 20th century although there was “dense forest” in the 1960's between South Nandi forest and Kakamega Forest. Bonner zoologische Beiträge 55 (2006) 125 Collection sites mentioned in this paper are as follows: the ‘Buyangu area’ is the name of the northern part of the Kakamega National Reserve (“primary-like” forest), with the Buyangu Village on his northern margin. Salazar Cir- cuit is an old plantation within the National Reserve, which is now secondary forest dominated by guava. Udo's Campsite is located within the northern part of the Nation- al Reserve and is the home of the Biota field camp. Isecheno is the low protected southern part of the forest. Rondo Retreat Centre is a small hotel within this area. 3. MATERIAL AND METHODS The material presented in this paper was partly collected during a three-month herpetological mission by the sen- ¡or author, which was carried out between March and June 2003. It was completed by some older voucher material from Kakamega Forest in the ZFMK collection, collect- ed by H.W. HERRMANN, D. MODRY and P. NECAS. Mate- rial from Kakamega Forest is also part of the collections of CAS, MHNG, NHMW, NMK, USNM, but this mate- rial was only partly analysed by the authors. Relevant lit- erature data was also evaluated. During the 2003 mission 170 reptiles were collected. Amphibians were not the main emphasis of the study but also collected and compared with the species list presented by SCHICK et al. (2005). Specimens of the study were fixed in 98 % ethanol and subsequently transferred to 70 % ethanol. For final dep- osition, they were equally partitioned between the NMK and ZFMK collections. The main forest and the forest fragments were walked during both day and night and specimens, mostly arboreal, were caught dominantly by visual encounter surveys along transects and opportunis- tic searches. In addition, Y-shaped drift-fences with pit- falls were used for terrestrial species. Catching success was low; only Adolfus africanus (Lacertidae) and amphib- ians were caught using this method. The roads were pa- trolled for snakes and the human inhabitants of the sur- rounding villages were recruited to help with collecting. The individual species accounts include the following parts: Specimens examined: gives a list of the material from museums collections examined by the authors; 4d- ditional specimens: refer to material known from other col- lections and not examined by the authors; Key references: lists publications with more detailed information on species of Kakamega forest; Remarks: gives information about the collected specimens and taxonomic statements. We follow mostly the classification and taxonomic con- clusions of FROST et al. (2006), however there are sever- al taxonomic conclusions in their amphibian tree of live tat we can not fathom and in our opinion require addition- al evidence and study. Collection codens: BIOTA= Biomonitoring Transect Analysis in Africa; BMNH = The Natural History Muse- um (British Museum [Natural History]), London, England; CAS= California Academy of Science, San Francisco, USA; MCZ= Museum of comparative Zoology, Harvard University, Cambridge/Massachusetts, USA; MHNG= Muséum d’histoire naturelle, Geneva, Switzerland; NHMW= Naturhistorisches Museum Wien, Vienna, Aus- tria, NMK= National Museums of Kenya, Nairobi, Kenya; PW= field number of the senior author; USNM= National Museum of Natural History, Smithsonian Insti- tution, Washington D.C., USA; ZFMK= Zoologisches Forschungsmuseum Alexander Koenig, Bonn, Germany. 4. RESULTS 4.1. Checklist of the herpetofauna of Kakamega Forest Amphibia Pipidae Gray, 1825 Xenopus victorianus Ahl, 1924 1924 Xenopus victorianus Ahl, Mitt. Zool. Mus. Berlin, 11: 270. Specimens examined. ZFMK 81733-735, 81940. Additional specimens. NMK A/3874/1, A/3935, A/3944, A/4025/1-2, A/4062/1-8, A/4163. Key references: SCHICK et al. 2005. Remarks: This species was often found in drift fence buckets in the Buyangu area near a small pond within the forest (ZFMK 81940). It was also found at Rondo Retreat in the southern part of the forest (ZFMK 81735). The vouchers are assigned to Xenopus victorianus in SCHICK et al. (2005) and to Xenopus sp. in LÖTTERS et al. (2006). Bufonidae Gray, 1825 Amietophrynus kisoloensis (Loveridge, 1932) 1932 Bufo regularis kisoloensis Loveridge, Occas. Pap. Boston Soc. Nat. Hist., 8: 52. Specimens examined. ZFMK 81727-730, 81943. Additional specimens. NMK A/104-106, A/108, A/1072/1- 2, A/1293/1-2, A/1318/1-2, A/1648/1-3, A/2046/1-4, A/3055/1-3, A/3104/1-3, A/3153/1-9, A/3563/1-2, A/3750/1-3, A/3813, A/3850/2, A/3851/1-3, A/3943. Remarks: ZFMK 81727-728 were collected in the Buyangu area. ZFMK 81943 was found in the Malaba fragment of the forest. ZFMK 81729 and A/3055/1-3 are from Rondo Retreat Centre, NMK A/3104/1-3 from Isecheno forest camp both located in the southern part of 126 Philipp WAGNER & Wolfgang BÖHME: The amphibians and reptiles of Kakamega Forest, western Kenya the forest. NMK A/3850/2 was collected at the Isiukhu falls in the Buyangu area. Amietophrynus maculatus (Hallowell, 1854) 1854 Bufo maculatus Hallowell, Proc. Acad. Nat. Sci. Philadelphia, 7: 101. Specimens examined. ZFMK 77458, 81723-726, 81941- 942. Additional specimens. NMK A/1194, A/3850/1,3-10. Remarks: ZFMK 81941-942 were collected within Buyangu Village along a road under stones next to a stream. NMK A/3850/5 was collected at the Istukhu falls in the Buyangu area. NMK A/1194 was from the Malava forest fragment. Ranidae Rafinesque, 1814 Hoplobatrachus occipitalis (Günther, 1859) 1859 Rana occipitalis Günther, Arch. Naturgesch., 24: 320. Specimens examined. NMK A/3938. Key references: SCHICK et al. 2005. Remarks: The voucher was collected in a swamp in the Buyangu area. Specimens, both adults and tadpoles, were found in an old swimming pool of the Serena Island Lodge in Kakamega town and were documented by a voucher specimen (NMK uncatalogued) and photographs. Fig. 2. Hoplobatrachus occipitalis from Kakamega town. Photo by Jórn KÓHLER. Phrynobatrachus graueri (Nieden, 1911) 1911 Arthroleptis graueri Nieden, Sitzungsber. Ges. Naturforsch. Freun- de Berlin, 1910: 441. Specimens examined. None. Additional specimens. NMK A/198, A/3105/2. Key references: LÖTTERS et al. 2006. Remarks: The specimen NMK A/198 was collected on a bridge near Kakamega town. NMK A/3105/2 was col- lected at the Kalunga glade at the Kakamega Forest. Phrynobatrachus aff. mababiensis FitzSimons, 1932 1932 Phrynobatrachus mababiensis FitzSimons, Ann. Transvaal Mus., 15: 40. Specimens examined. None. Remarks: This species has only been recorded from lit- erature (SCHICK et al. 2005) and ıs currently under inves- tigation by SCHICK et al. (2005) and other colleagues. Phrynobatrachus natalensis (Smith, 1849) 1849 Stenorhynchus natalensis Smith, Ill. Zool. S. Afr., 3 (Appendix): 24. Specimens examined. ZFMK 81742-43. Additional specimens. NMK A/3105/1, A/3863/1-4, A/3931, A/3932. Remarks: NMK A/3105/1 was from the Kalunga glade within the Kakamega forest area. The series NMK A/3863 and the ZFMK vouchers were from the Buyangu area. De- tails on the other vouchers are unknown. Phrynobatrachus aff. minutus (Boulenger, 1895) 1895 Arthroleptis minutus Boulenger, Proc. Zool. Soc. London, 1895: 539. Specimens examined. None Additional specimens. NMK A/3924/1-6, A/4310. Key references: LÖTTERS et al. 2006. Remarks: Details on the vouchers are unknown. NMK A/3924/1,2,5 are on permanent loan to the ZFMK. This taxon was not recorded by SCHICK et al. (2005). Afrana angolensis (Bocage, 1866) 1866 Rana angolensis Bocage, J. Sci. Math. Phys. Nat., Lisboa, 1: 73. Specimens examined. NMK A/100/1-8, A/101, A/102/1- 5 Additional specimens. NMK A/1294/1, A/1314/1-8, A/1649/1-3, A/3639, A/3937/1-2, A/4239. Key references: LÖTTERS et al. 2006. Remarks: NMK A/4239 was found along the Salazar Bonner zoologische Beitráge 55 (2006) 127 road. The series NMK A/1314/1-8 and A/1649 were col- lected at the Ikuywa River. Details on the other specimens are unknown. Amnirana cf. albolabris (Hallowell, 1856) 1856 Hyla albolabris Hallowell, Proc. Acad. Nat. Sci. Philadelphia, 8: 153. Specimens examined. None. Additional specimens. NMK A/196/1-2, A/1966. Key references: SCHICK et al. 2005; LÖTTERS et al. 2006. Remarks: The specimens NMK A/196/1-2 came from Kakamega town, near the Forest Department Pump House. Ptychadena anchietae (Bocage, 1867) 1867 Rana anchietae Bocage, Proc. Zool. Soc. London, 1867: 843. Specimens examined. NMK A/3845. Additional specimens. A/4212, A/4216, A/4220/1-2, A/4224, A/4226/1-2, A/4234/1-3. Remarks: NMK A/4212 was collected in a small puddle near the Buyangu view point; NMK A/4220/1-2 were col- lected in amplexus on the 28.1V.2004 from the same pud- dle. NMK A/4216 was found in short grass habitat next to a small puddle within the Salazar secondary forest. NMK A/4224 was collected on new Buyangu Campsite. NMK A/4234/1 was collected near the Keep office, A/4234/2-3 near the Isecheno Primary school. Details on the other specimens are unknown. Ptychadena porosissima (Steindachner, 1867) 1867 Rana porosissima Steindachner, Reise Österreichischen Fregatte Novara, Zool.: 18. Specimens examined. None. Additional specimens. NMK A/3107/1-3, A/3574, A/4222. Key references: SCHICK et al. 2005; LÖTTERS et al. 2006. Remarks: NMK A/3107/1-3 were collected at Kalunga glade. NMK A/4222 was found calling on wet mud near a water edge in the Buyangu area. Details of the other specimen are unknown. Ptychadena taenioscelis Laurent, 1954 1954 Ptychadena taenioscelis Laurent, Ann. Mus. R. Congo Belge, Ter- vuren, Zool., 34: 25. Specimens examined. None. Additional specimens. NMK A/3955/1-2, A/4213. Key references: LÖTTERS et al. 2006. Remarks: NMK A/4213 was found in a small swamp at the Buyangu area. Details on the other voucher are un- known. This species was recorded for the first time for Kenya by LÓTTERS et al. (2006) and at present is only known to occur in Kakamega Forest within Kenya. Ptychadena aff. mascareniensis (Duméril & Bibron, 1841) 1841 Rana mascariensis Duméril & Bibron, Erp. Gen., 8: 350. Specimens examined. ZFMK 81944. Additional specimens. NMK A/3572, A/3840/4-5, A/3856, A/4214, A/4217/1-2, A/4221, A/4223, A/4227/1-10, A/4229/1-2, A/4232/1-4, A/4235/1-9. Key references: VENCES et al. 2004; LÖTTERS et al. 2006. Remarks: P mascareniensis was often found on the shore of the Isıukhu River in its small discharging streams and in the Buyangu area. NMK A/4227/1-3 was found near the Keep Office at Isecheno, NMK A/4227/4-10 in a tem- porary swamp at Isecheno. This taxon was listed by LÖT- TERS et al. (2006) as Ptychadena aff. mascareniensis. Ptychadena oxyrhynchus (Smith, 1849) 1849 Rana oxyrhynchus Smith, Ill. Zool. S. Afr., 3(Part 28): pl. 77, fig 2. Specimens examined. None. Additional specimens. NMK A/103/1-2, A/3846, A/4211, A/4215/1-4, A/4218, A/4219/1-3, A/4225/1-2, A/4228/1- 3, A/4230/1-2, A/4231/1-2, A/4233/1-4, A/4236/1-2. Key references: LÓTTERS et al. 2006. Remarks: NMK A/4225/1-2, A/4228/1-3 and A/4236/1- 2 were collected in a pond on the new Buyangu Camp- site, NMK A/4231/1-2 was found there in amplexus on the 31. V. 2004 and the female laid eggs over night. NMK A/4233/1-4 were found at Isecheno in the southern part of the forest. Details on the other specimens are unknown. This taxon was listed by LÖTTERS et al. (2006) as Pty- chadena aff. oxyrhynchus. Hyperoliidae Laurent, 1943 Afrixalus osorioi (Ferreira, 1906) 1906 Rappia osorioi Ferreira, J. Sci. Math. Phys. Nat., Lisboa, Ser. 2, 72 162; Specimens examined. NMK A/3927/2. 128 Philipp WAGNER & Wolfgang BÖHME: The amphibians and reptiles of Kakamega Forest, western Kenya Fig. 3. Afrixalus osorioi from Kakamega Forest. Photo by Jörn KÖHLER. Additional specimens. A/3928, A/4017, A/4316/1-7. Key references: KÖHLER et al. 2005. Remarks: NMK A/3927/2 is on permanent loan to the ZFMK. NMK A/4017 and the series A/4316 were collect- ed at a pond within the Buyangu area. One specimen was additionally sighted in the South Nandi forest. This species was recorded for the first time for Kenya by KOHLER et al. (2005) and at present only occurs in the Kakamega For- est complex within Kenya. Afrixalus quadrivittatus (Werger, 1908) 1908 *1907” Megalixalus leptosomus leptosomus Werger, Sitzungsber. Akad. Wiss. Wien, Phys. Math. Naturwiss. Kl., 116: 1900. Specimens examined. NMK A/3933/2. Additional specimens. NMK A/3933/1, A/4317/1-7. Key references: KÓHLER et al. 2005. Remarks: NMK A/3933/2 is on permanent loan to the ZFMK and was collected at a swamp in the Buyangu area. The series NMK A/4317 was collected at the pond of the Buyangu area. Hyperolius acuticeps Ahl, 1931 1931 Hyperolius acuticeps Ahl, Das Tierreich, 55: 282. Specimens examined. ZFMK 77616, 81749-750. Additional specimens. NMK A/3922/2. Key references: LÓTTERS et al. 2006. Remarks: ZFMK 81749-750 were from the Buyangu area. Details on the other vouchers are unknown. Hyperolius kivuensis Ahl, 1931 1931 Hyperolius kivuensis Ahl, Das Tierreich, 55: 280. Specimens examined. ZFMK 81745-746, 82183-184. Additional specimens. NMK A/3032/1-16, A/3103, A/3579, A/3709, A/3710, A/3748/1-4, A/3825/1-6, A/3864/4, A/3867/1-5, A/3953, A/4011/1-2, A/4026/1-2, A/4065. Key references: SCHICK et al. 2005; LÖTTERS et al. 2006. Remarks: NMK A/3103, A/3867, A/4011, A/4026 and ZFMK 81745-746 were from the Buyangu area. NMK A/4065 was collected at the BIOTA Campsite. Details on the other vouchers are unknown. Fig. 4. Hyperolius lateralis from Kakamega Forest. Photo by Jörn KÖHLER. Hyperolius lateralis Laurent, 1940 1940 Hyperolius lateralis Laurent, Rev. Zool. Bot. Afr., 34: 1. Specimens examined. ZFMK 81747-748. Additional specimens. NMK A/2075/5-6, A/2075/8-10, A/3925/1-4, A/3936. Remarks: NMK A/3925/1 is on permanent loan to ZFMK. ZFMK 81747-748 were from the Buyangu area. NMK A/3936 was from Rondo Retreat Centre in the southern part of the forest. Details on the remaining vouch- ers are unknown. Within Kenya, this taxon is currently on- ly recorded from the Kakamega Forest. Hyperolius viridiflavus (Duméril & Bibron, 1841) 1841 Eucnemis viridi-flavus Duméril & Bibron, Erp. Gen., 8: 528. Specimens examined. ZFMK 77426, 81950. Additional specimens. NMK A/3056/1-3, A/3578, A/3866/5-11, A/668/1-4, A/1193/1-2, A/1444/1-5, A/3749, A/3809/1-5, A/3954/1-2, A/4013, A/4027/1-3, A/4066. Bonner zoologische Beiträge 55 Remarks: Details on the vouchers are unknown, but most are from the Buyangu area. A. viridiflavus is the most common frog inside the forest. Specimens were observed at different ponds and also clearings for example, the Bio- TA field camp, where several specimens were sitting in- side the lavatory. NMK A/1193/1-2 were from Malava for- est. Fig. 5. Hyperolius cinnamomeoventris from Kakamega Forest. Photo by Jórn KÓHLER. Hyperolius cinnamomeoventris Bocage, 1866 1866 Hyperolius cinnamomeo-ventris Bocage, J. Sci. Math. Phys. Nat., Lisboa, 1: 75. Specimens examined. ZEMK 77431-432, 81744. Additional specimens. NMK A/2095/1, A/3858/1-2, A/3918/1-2, A/4012. Key references: SCHICK et al. 2005; LÖTTERS et al. 2006. Remarks: NMK A/3858, A/4012 and ZFMK 81744 are from the Buyangu area. This taxon was reported by LÖT- TERS et al. (2006) as Hyperolius aff. cinnamomeoventris. Fig. 6. Kassina senegalensis from Kakamega Forest. Photo by Jórn KÓHLER. (2006) 129 Kassina senegalensis (Duméril & Bibron, 1841) 1841 Cystignathus senegalensis Duméril & Bibron Erp. Gen., 8: 418. Specimens examined. ZFMK 81741, 81946-949. Additional specimens. NMK A/1158, A/1317, A/3711, A/3920. Remarks: All vouchers were from the Buyangu area and some were collected with a drift fence next to a pond. One specimen was collected on the Buyangu Hill outside the forest under stones. ic Fig. 7. Leptopelis mackayi from Kakamega Forest. Photo by Arne SCHIOTZ. Arthroleptidae Mivart, 1869 Leptopelis mackayi Kohler, Bwong, Schick, Veith & Lótters, 2006 2006 Leptopelis mackayi Kohler, Bwong, Schick, Veith & Lotters, Her- petological Journal 16: 183-189. Specimens examined. ZFMK 83304-305 (paratypes), ZFMK 83306. Additional specimens. NMK A/3057/1 (holotype), NMK A/1407/1-3 (paratypes), NMK A/3072/1. Key references: KÖHLER et al. 2006. Remarks: This species was recently described by KÖH- LER et al. (2006) and is only known from Kakamega For- est and its vicinity and inhabits so far only forest habitats (also secondary and disturbed forest). It represents the sis- ter taxon of the West African L. modestus and was referred to this species in the past by ScHıoTz (1975, 1999). Leptopelis aff. bocagii (Günther, 1865) 1865 “1864” Cystignathus bocagii Günther, Proc. Zool. Soc. London, 1864: 481. Specimens examined. None. Additional specimens. UZM R/074960-2. 130 Bonner zoologische Beitráge 55 (2006) Fig. 8. Leptopelis aff. bocagii undescribed form from Kakame- ga Forest. Photo by Arne SCHIOTZ. Key references: SCHIOTZ 1975. Remarks: ScHioTz (1975) collected two males and one female (UZM R/074960-2). These vouchers were taken from a savannah-like clearing near the Forest Station in Kakamega Forest. One male was sitting in the bush when calling. See ScHıoTz (1975) for more details on the call. L. aft. bocagii is an undescribed species, very close in mor- phology to L. bocagii. Only a few specimens are known, all from Kakamega (A. SCHIOTZ, pers. comm.). Reptilia Agamidae Spix, 1825 Acanthocercus atricollis (Smith, 1849) 1849 Agama atricollis Smith, Illustrations of the Zoology of South Africa. 3 (Reptiles). Specimens examined. NMK L/2655, 2660/2-3; ZFMK 81952-963. Additional specimens. CAS 122731-122739. Key references: KLAUSEWITZ 1957. Fig. 9. Acanthocercus atricollis from Kakamega Forest. Pho- to by Philipp WAGNER. Remarks: The debate of the taxonomic status of this species is still ongoing. Many authors (e. g. BOULENGER 1896, KLAUSEWITZ 1957, LOVERIDGE 1957) have discussed differences or similarities between this taxon and Acan- thocercus cyanogaster RÚPPELL 1835. SPAWLS et al. (2002), LARGEN & SPAWLS (2006) and our own morphol- ogy studies of the two species support KLAUSEWITZ (1957) who regarded them as two distinct species. The reported distribution of both taxa is unclear because of the men- tioned taxonomic problems. Despite the works of KLAUSEWITZ (1954, 1957), a new re- view of the Acanthocercus- species complex is needed. The review will be a part of the PhD thesis of the senior author. The diagnoses of the subspecies of A. atricollis by KLAUSEWITZ (1957) are not adequate. Therefore, the ma- terial is only preliminarily assigned to the subspecies ugandaensis because Kakamega Forest is geographical- ly closer to the area of this subspecies than to minuta, ac- cording to KLAUSEWITZ’s (1957) map: A. a. minuta imhab- its Ethiopia and eastern Kenya, while A. atricollis ugan- daensis occurs within Uganda and western Kenya. Ea << MO [Eco Fig. 10. Agama kaimosae from Ngoromosi/ Nandi escarpment. Photo by Alexander BURMANN. Bonner zoologische Beitráge 55 (2006) 131 This agama subspecies was found in all areas surround- ing Kakamega Forest. It reaches the highest density in small villages and Ranger Stations and lives there on the clay huts, in syntopy with Trachylepis striata and Adol- fus jacksoni. It also occurs in hedges close to trees. It was not found inside the forest and also not in plantations and on forest edges. Agama kaimosae Loveridge, 1935 1935 Agama agama kaimosae Loveridge, Bull. Mus. Comp. zool., Har- vard 79:10. Specimens examined. ZFMK 83658-660; NMK L/2715/1,3,4; MCZ 40136- 40150. Key references: BURMANN 2006, WAGNER et al. 2007. Remarks: After its synonymisation by LOVERIDGE (1936), this taxon was regarded as a synonym of Agama cau- dospinosa. But further investigations (BURMANN 2006, Wagner et al. 2007) have shown that Agama kaimosae is a valid species. The taxon is a SW Kenyan/N Tanzanian endemic (see map in WAGNER et al. 2007). The type lo- cality is a rocky highland three miles west of the Friends’ Africa Mission Station at Kaimosi. However no specimens were found there, neither by the senior author in 2003 nor by A. BURMANN in 2005. Our specimens were collected in Ngoromosi in the Nandi escarpment. Chamaeleonidae Gray, 1825 Chamaeleo gracilis Hallowell, 1842 1842 Chamaeleo gracilis Hallowell, J. Acad. Nat. Sci. Philadelphia 8: 324. Specimens examined. NMK L/2203/1, NMK L/2653/1-2; ZFMK 82055-056. Remarks: The species was not found in Kakamega For- est directly but in forest edge areas of the South Nandi For- est, located slightly east of Kakamega Forest. These forests were united with Kakamega until recently (MITCHELL 2004), but in contrast to Kakamega the Nandi Escarpment has. montane vegetation. The specimens were found in bushes next to the forest, on the western slopes of the Nan- di escarpment. Chamaeleo hoehnelii Steindachner, 1891 1891 Chamael(ejo höhnelii Steindachner, Anz. Akad. Wiss. Wien 28: 141. Specimens examined. NMK L/252. Remarks: This voucher was collected in Kakamega town, so it is possible that the species also occurs in the sur- rounding area of Kakamega Forest. Fig. 11. Chamaeleo ellioti from Kakamega Forest. Photo by Philipp WAGNER. Chamaeleo laevigatus Gray, 1863 1863 Chamaeleo laevigatus Gray, Proc. Zool. Soc. London 1863: 95. Specimens examined. NMK L/1588. Remarks: The only specimen of this taxon was found 1981 by MADSEN six kilometres north of Kakamega town. Further data was not available. Chamaeleo ellioti Gúnther, 1895 1895 Chamaeleo ellioti Gúnther, Ann. Mag. Nat. Hist., London 15: 524, pl. 21, fig. A. Specimens examined. NMK L/2480, L/1227, L/1273, L/2647, L/2653/1-2, L/2652/1-2, L/2658; ZFMK 54025, 68494-97, 70835, 81974-80, 82057-58. Additional specimens. CAS 147912, 153247. Remarks: The species has a wide distribution in the Kakamega area and specimens were found in several habi- tats in and around the forest. In the year of the fieldwork 132 Philipp WAGNER & Wolfgang BÖHME: The amphibians and reptiles of Kakamega Forest, western Kenya Fig. 12. Rhampholeon boulengeri from Kakamega Forest. Pho- to by Philipp WAGNER. of the senior author, C. ellioti was one of the species with the highest observed density. Only two years later 1t was very difficult to find them (P. HITA-GARCIA pers. comm.). This was also corroborated by SpawLs et al. (2002) and SCHMIDT et al. (2000) who also indicate that C. ellioti has large population density dynamics. This chameleon was found in the surrounding areas of Kakamega and Nandi forests, and also inside the forest, on the shore of the Istukhu River. Habitats were as follows: grassland, banana and guava plantations, gardens, hedges, bushes, small trees, secondary and riverside forest. Rhampholeon boulengeri Steindachner, 1911 1911 Rhampholeon boulengeri Steindachner, Anz. Akad. Wiss., Wien, 48: 178. Specimens examined. NMK L/249, L/7270, L/2651, ; ZFMK 54021-24, 77456, 81999. Remarks: R. boulengeri ıs a typical dweller of the East African rain forests from the eastern parts of the DR Con- go in the west to Kenya in the east. Here it is, apart from Kakamega and Nandi forests, also found in the Cheran- gani Mts. (SPAWLS et al. 2002). Within Kakamega Forest the pygmy chameleon was found in the northern part of the main fragment and in the Kisere fragment. Gekkonidae Oppel, 1811 Cnemaspis africana (Werner, 1895) 1895 Gymnodactvlus africanus Werner, Verh. zool.-bot. Ges. Wien 45: 190, pl. V, fig. 5. Specimens examined. NMK L/1987; ZFMK 82022. Remarks: This taxon is endemic to eastern Africa and is only distributed in Kenya and Tanzania. Next to Kakamega Forest, the lizard is only known from Taita and Shimba Hills and Athiplain. ZFMK 82022 was collected in a pitfall trap near the Buyangu Hill. This suggests that C. africana is not a strict tree dweller but also inhabits leaf litter during its search for food. Fig. 13. Cnemaspis elgonensis from Kakamega Forest. Photo by Philipp WAGNER. Cnemaspis elgonensis Loveridge, 1935 1935 Cnemaspis elgonensis Loveridge, Proc. Zool. Soc. London 1935: 820. Specimens examined. NMK L/1129, L/2263; ZFMK 82023. Additional specimens. USNM 158923. Remarks: C. elgonensis is endemic to Uganda and Kenya and has a very restricted distribution area from the Mt. El- gon range to Kakamega in the south. Nothing is known about its biology but similarities with other species of the genus can be anticipated. ZFMK 82023 was found at dawn on the lavatory of Udo’s Campsite in the northern part of the forest. Hemidactylus mabouia (Moreau de Jonnes, 1818) 1818 Gekko mabouia Moreau de Jonnes, Bull. Scient. Soc. Philomath. Paris, sér 3,5: 138. Specimens examined. NMK L/2648/1-2, L/2659/1, L/2659/3-4; ZFMK 70833-34, 81982-86. Remarks: This species inhabits nearly the entire sub-Sa- haran Africa. It has been transported by humans to South America (DIRKSEN 1995), Florida, USA (POWELL et al. 1998), Madagascar (GLaw & VENCES 1994) and to Madeira, Portugal (JESUS et al. 2002). In Kakamega it in- habits the houses of the villages around the forest and al- so of Kakamega town. No specimens were found outside of human habitations. Bonner zoologische Beitráge 55 (2006) 133 Fig. 14. Lygodactylus gutturalis from Kakamega Forest. Pho- to by Jörn KÖHLER. Lygodactylus gutturalis (Bocage, 1873) 1873 Hemidactylus gutturalis Bocage, J. Sci. math. phys. nat. Lisboa 4: 211. Specimens examined. NMK L/2464; ZFMK 81987. Remarks: This is the first record of the species for Kenya. The geographically closest record is from the Ugandan side of Mt. Elgon. L. gutturalis is a typical species of the equatorial rain forest and is distributed from Senegal in the west to Kenya in the east. As it is the case in several other forest species with the same distribution pattern (e. g. Lepidothyris fernandi species complex: WAGNER et al. subm.), the East African populations might prove to be a new taxon. NMK L/2464 was found inside a tent on Udo's Campside. Scincidae Oppel, 1811 Eumecia anchietae Bocage, 1870 1870 Eumecia anchitae Bocage, J. Sci. Lisboa 3: 66-68. Specimens examined. NMK L/110/1+2, L/2657, L/2669; ZFMK 75069, 76044, 81981. Remarks: This species was found in several habitats, e. g. urban areas inside the villages and Guava dominated secondary structures. ZFMK 81981 was found on a tree (Strychnaceae: Strychnus cf. usambarensis), which was very slanting and shaggy with ferns and mosses, so this sighting was presumable an exception of this normally ground-dwelling species. Feylinia currori Gray, 1845 1845 Feylinia currori Gray, Catalogue of the lizards of the British Mu- seum, p. 129. Specimens examined. NMK L/2662; ZFMK 81998. Key references: WAGNER & SCHMITZ 2006. Remarks: These specimens were the first records of this species and genus in Kenya (WAGNER & SCHMITZ 2006). They were found crossing a road (ZFMK 81998) and in leaf litter (NMK L/2662). A morphological comparison between east and west African populations results in no geographic directed differences (WAGNER & SCHMITZ 2006). se = E de Fig. 15. Feylinia currori from Kakamega Forest. Photo by Phil- ipp WAGNER. Lepidothyris aff. fernandi (Burton, 1836) 1836 Tiliquia fernandi Burton, Proc. Zool. Soc. London 1836: 62. Specimens examined. NMK L/2147. Key references: WAGNER et al. (subm.). Remarks: This secretive skink is probably the rarest rep- tile of the forest. It is only recorded by one voucher spec- imen and from one sighting (S. SCHICK, pers. comm.) in 2002. This typical equatorial rain forest species complex is distributed from Sierra Leone in the west to Kenya in the east and is currently portioned into several taxa (WAG- NER et al. subm.). The East African populations inhabit the forests of the eastern DR Congo to Kenya, southwards to Zambia and south-westwards to Congo and Angola. The single specimen was found in a pitfall trap next to Buyan- gu Hill. Also the visual record was on a forest path with- in this area. Afroablepharus wahlbergi (Smith, 1849) 1849 Cryptoblepharus wahlbergi Smith, Illus. zool. S. Africa, 3, App. p.10. Specimens examined. None. Additional specimens. MCZ 41601-10, 41614-7. Key references: LOVERIDGE 1936. Remarks: The vouchers were collected by LOVERIDGE (1936) at Kaimosi area and he remarked that most of the collected female specimens were pregnant in February. 134 Philipp WAGNER £ Wolfgang BÖHME: The amphibians and reptiles of Kakamega Forest, western Kenya Trachylepis maculilabris (Gray, 1845) 1845 Euprepis maculilabris Gray, Catalogue of the lizards of the British Museum. Specimens examined. None. Additional specimens. CAS 122720, 122723; USNM 49203. Key references: DREWES 1976. Remarks: The material from CAS was collected by DREWES (1976) in Kaimosi area. Since then no other vouchers were collected. Trachylepis megalura (Peters, 1878) 1878 Euprepis megalurus Peters, Monatsber. Akad. Wiss. Berlin, p. 204. Specimens examined. NMK L/1915/1-2; ZFMK 81997. Additional specimens. CAS 122728; USNM 49066-68, 49199. Remarks: Only a single specimen was found during the survey 2003 on a slope of the Liranda Hill in the south- ern part of the forest. This habitat is dominated by grass- land with sporadic trees. The only other reptile species recorded on this hill in rocky areas was 7. quinquetaeni- ata. DREWES (1976) found the taxon also on clearings in the Buyangu arca. Trachylepis quinquetaeniata (Lichtenstein, 1823) 1823 Scincus quinquetaeniatus Lichtenstein, Verzeichnis der Dubletten, Berlin, p. 103. Specimens examined. NMK L/2650/2-3, 2650/5, 2656/2- 3; ZFMK 81988-93, 81995-96, 82060-61. Additional specimens. CAS 122709-719, 122721-722. Remarks: The species shows a disjunct distribution with- in the Kakamega area. It is only found in rocky areas of the Buyangu and Liranda Hill as well as on a small hill near Kisere. No specimens were found on houses, bridges or other human buildings as 1t was described by SPAWLS et al. (2002). Also DREWES (1976) found the speci- mens on exposed rocks. FINK (2003) has shown that the diet is dominated by isopod species, but also molluses were found. Trachylepis striata (Peters, 1844) 1844 Tropidolepisma striatum Peters, Berl. Bekanntmach. Geeignet. Ver- handl. Königl.-Preuss. Akad. Wiss. Berlin 1844: 32-37. Specimens examined. NMK L/2654/2, 2654/5; ZFMK 70825-30, 82002-06, 82011-21. Additional specimens. CAS 122724-727; USNM 49069- 71, 49207-16, 49389, 49393. Remarks: 7. striata is one of the species with the high- est density in Kakamega area and was found everywhere outside the forest or wooded areas. But meanwhile the species has reached the Biota Camp, located on a small clearing of Udo’s Campside inside the forest. Here and on the houses of the near villages it is sympatric with 4dol- fus jacksoni (Lacertidae) and only on the houses also with Acanthocercus atricollis (Agamidae). In contrast to the da- ta given by RAZZETTI & Msuya (2002) and SPAWLs et al. (2002), T. striata was never found on trees or in planta- tions. The diet analysed by FINK (2003) is dominated by Coleoptera and also consists of other winged insects, collembolans, spiders, nematodes and molluscs. Fig. 16. Adolfus africanus from Kakamega Forest. Photo by Philipp WAGNER. Lacertidae Oppel, 1811 Adolfus africanus (Boulenger, 1906) 1906 Algiroides africanus Boulenger, Proc. Zool. Soc. London 2: 570- 512: Specimens examined. NMK L/2661/2; ZFMK 77457, 81205-07. Key references: KÖHLER et al. 2004. Remarks: These specimens were the first record of the species for Kenya (KÖHLER et al. 2004). As a typical in- habitant ofthe equatorial rainforest, A. africanus has a dis- junct distribution from Cameroon in the West to Kenya in the East, but taxonomic comparisons have shown that there are no geographic directed differences between the populations (KÓHLER et al. 2004) and no subspecies are recognizable. Most of the vouchers were collected in a pit- fall trap near the Buyangu Hill. ZFMK 81207 was collect- ed by hand in a secondary guava forest, the Salazar Cir- cuit. ZFMK 77457 was collected in the southern part of the forest near Isecheno by W. FREUND and J. KÓHLER in 2002. Bonner zoologische Beitráge 55 (2006) 135 Fig. 17. Adolfus jacksoni from Kakamega Forest. Photo by Jórn KÖHLER. Adolfus jacksoni (Boulenger, 1899) 1899 Lacerta jacksoni Boulenger, Proc. Zool. Soc. London 1: 96-98. Specimens examined. PW 04 & PW 05 (now part of the NMK collection); ZFMK 70831-32, 81964-73. Additional specimens. CAS 122729-30, 141566, 147904. Key references: SPAWLS & ROTICH 1997. Remarks: 4. jacksoni was recorded for Kakamega For- est by SPAWLS & ROTICH (1997) for the first time. Most of the specimens were collected in the Biota Camp and in forest surrounding villages. No specimens were found in gardens or small plantations but they were sighted on trees within maize and cane fields. Only one specimen was sighted near to the forest inside the Salazar Circuit. In con- trast to the data provided by RAZZETTI & Msuya (2002), A. jacksoni was rarely seen climbing on trees. The diet consists mainly of Isoptera, Orthoptera and Lepidoptera, but also of other arthropods. Cordylidae Mertens, 1937 Chamaesaura anguina (Linnaeus, 1758) 1758 Chamaesaura tenuior Linnaeus, Systema Naturae. 10th edition. Specimens examined. NMK L/2020. Remarks: The specimen was collected in 1992 probably in the southern part of the forest. Further data are not avail- able. Because of the distribution this specimen was as- signed by use to the subspecies tenuior. Varanidae Hardwicke & Gray, 1827 Varanus aff. niloticus (Linnaeus, 1766) 1766 Lacerta nilotica Linnaeus, Syst. Nat. Ed. 12 (1): 369. Specimens examined. None. Remarks: This species is known only from literature and sightings (LOVERIDGE 1936; MERTENS 1942). The details of the specimens seen are unknown. Specimens were sighted (by the first author and by J. KÓHLER, pers. comm.) on the shore of the Isiukhu River near Buyangu village. It remains to be shown whether the Kakamega population belongs to V. niloticus or V. ornatus. The fact that Kakamega Forest is a remnant of the equatorial rain for- est makes the existence of Y. ornatus possible. The near- est locality documented by a voucher specimen is Kisumu (NMK L/2476), but this is a typical habitat for V. niloticus. Also ANALO (2003) refers to the occurrence of Varanus niloticus in the Kakamega area and reported that the skin is used for traditional drums. Typhlopidae Merrem, 1820 Typhlops angolensis (Bocage, 1866) 1866 Onychocephalus angolensis Bocage, Jorn. Sci. Math. Phys. Nat. Lisboa 1: 46, 65. Specimens examined. None. Remarks: This taxon is known only from literature (LOVERIDGE 1935) and further data were not available. Fig. 18. Typhlops lineolatus from Kakamega Forest. Photo by Mike DoBIEY. Typhlops lineolatus Jan, 1864 1864 Typhlops lineolatus Jan, Iconogr. Gén. Ophid. 1(9. livr.): 5. Specimens examined. PW 157 £ 162 (now part of the NMK collection); ZFMK 73283, 82051-52. Remarks: Most of the vouchers were collected after rain- fall on Udo's Campsite. One specimen (ZFMK 82051) was from Buyangu village and collected in a grassy area on a rainy day. 136 Philipp WAGNER & Wolfgang BÖHME: The amphibians and reptiles of Kakamega Forest, western Kenya Colubridae Oppel, 1811 Crotaphopeltis hotamboeia (Laurenti, 1768) 1768 Coronella hotamboeia LAURENTI, Syn. Rept.: 58. Specimens examined. NMK O/2691 Remarks: Further data on the NMK specimen is not avail- able, but one specimen was sighted at Salazar Circuit. Dasypeltis atra Sternfeld, 1912 1912 Dasypeltis scabra var. atra Sternfeld, IV. Zool. II Lfg. Reptilia, in: Schubotz: Wiss. Ergeb. Deut. Zentr. Afr. Exp: 197-272. Specimens examined. MHNG 1.262.072-075; NMK S/2576; ZFMK 77459, 82054. Additional specimens. CAS 142248. Remarks: SPAwLs et al. (2002) noted that completely black specimens have been collected east of the Rift Val- ley. So, ZFMK 82054 is probably the first melanistic voucher from west of the Rift Valley. However, ZFMK 77459 demonstrates sympatric occurrence of the light colour phase with the melanistic one. Dasypeltis scabra (Linnaeus, 1758) 1758 Coluber scaber Linnaeus, Syst. Nat. Ed. 10(1): 824, Specimens examined. ZEMK 75070. Additional specimens. USNM 49376. Remarks: Further data are not available, but specimens were sighted at Salazar Circuit. Dispholidus typus Smith, 1829 1829 Bucephalus typus Smith, Zool. Journ. 4: 433-444. Specimens examined. NMK S/986 Remarks: This specimen is assigned to the subspecies kivuensis, because of its distribution from west of the Rift Valley in Kenya to Rwanda in the east and Zambia in the south. Further data are not available, but one specimen was sighted by the senior author in secondary forest at the Salazar Circuit. Hapsidophrys lineatus Fischer, 1856 1856 Hapsidophrys lineatus Fischer, Abhandl. Nat. Ver. Hamburg 3 (4): 81-116. Specimens examined. NMK O/2715; ZFMK 82053. Fig. 19. Dispholidus typus from Kakamega Forest. Photo by Mike DoBIEY. Additional specimens. CAS 147910-11, 147913, 150987, 193222; Remarks: This species which is typical for the equatori- al rainforest, is distributed from Guinea-Bissau in the west to Kenya in the east and taxonomic analyses did not show geographic directed differences between the populations. ZFMK 82053 was found basking on a bush next to a small pond in the Buyangu area. Lamprophis fuliginosus (Boie, 1827) 1827 Lycodon fuliginosus Boie, Isis van Oken 20, col. 551. Specimens examined. NMK S/3981/1-2; ZFMK 82037-43. Additional specimens. CAS 122743, 141529. Remarks: All vouchers were collected inside the houses or gardens of Buyangu Village. Lycophidion capense (Smith, 1831) 1831 Lydodon capensis Smith, S. Afr. Quart. J.. 1: 18. Specimens examined. None. Additional specimens. CAS 122741. Remarks: Further data are not available. The specimen is catalogued at CAS as L. c. capense BOULENGER, 1893. Lycophidion depressirostre Laurent, 1968 1968 Lycophidion depressirostre Laurent, Bull. Mus. Comp. Zool., Harv. 136 (12): 472. Specimens examined. None. Additional specimens. USNM 49388. Remarks: Further data are not available. Bonner zoologische Lycophidion ornatum Parker, 1936 1936 Lycophidion ornatum Parker, Novit. Zool. 40: 122. Specimens examined. ZFMK 75071, 82044. Additional specimens. BMNH 1962.819; MCZ 40471-73. Remarks: L. ornatum has a wide distribution from Nige- ria in the West to Mt Kenya in the East. ZFMK 82044 was found in twilight inside the forest. Mehelya capensis (Smith, 1847) 1847 Heterolepis capensis Smith, Illustrations of the zoology of South Africa, Reptilia. Specimens examined. None. Additional specimens. CAS 150988. Remarks: This voucher is catalogued at CAS as M. c. sa- vorgnani MOCQUARD, 1887. Further data are not available. Natriciteres olivacea (Peters, 1854) 1854 Coronella olivacea Peters, Monatsber. Kónigl. Akad. Wiss. Berlin 1854: 614-628. Specimens examined. ZFMK 82035. Remarks: N. olivacea inhabits water bodies in bushland and savannah regions. In contrast to the described habi- tats (e. g. SPAWLS et al. 2002; MARAIS 2004), our speci- men was collected basking on dense vegetation on the shore of the Isiukhu inside the northern part of the forest. But there is another specimen collected inside a forest by ULLENBRUCH (2003) in Benin. Philothamnus battersbyi Loveridge, 1951 1951 Philothamnus irregularis battersbyi Loveridge, Bull. Mus. Comp. zool. Harvard 106: 51. Specimens examined. NMK S/3986, S/3992; ZFMK 82048. Additional specimens. CAS 150978-980, 153223. Remarks: Philothamnus is probably one of the most dif- ficult, for the taxonomists, reptile genera in Africa. But P. battersbyi 1s relatively easy to identify by the uniform green colouration, two supralabials entering the eye, a di- vided anal scale, 15 midbody scale rows and no keels on the subcaudal scales. It can only be confused with the like- wise uniform green P angolensis. Most vouchers collect- ed within the study were found in Buyangu Village bask- ing on small bushes. One was found killed on the road. Beitrage 55 (2006) Fig. 20. Philothamnus battersbyi from Kakamega Forest. Pho- to by Philipp WAGNER. Philothamnus carinatus (Andersson, 1901) 1901 Chlorophis carinatus Andersson, Bihang Till. K. Sv. Vet.- Akad. Handl. 27 (5): 9. Specimens examined. None. Remarks: This taxon is only recorded from literature (HUGHES 1985; SPAWLS et al. 2002). Further data are not available. Philothamnus heterolepidotus (Günther, 1863) 1863 Ahaetulla heterolepidota Gúnther, Ann. Mag. Nat. Hist. 11 (3): 286. Specimens examined. NMK S/65-66, S/68, S/120-123. Remarks: The vouchers were not collected in the area close to Kakamega Forest but on the shores of the Yala River in the Siaya District. The Yala also crosses the south- ern part of the Kakamega Forest. In view of this, the oc- currence of the species in the Kakamega area is likely. Philothamnus hoplogaster (GUNTHER, 1863) 1863 Ahaetulla hoplogaster GUNTHER, Ann. Mag. Nat. Hist. 11 (3): 284. Specimens examined. None. Remarks: This taxon is only recorded from literature (HUGHES 1985; SPAWLS et al. 2002). Further data are not available. Philothamnus nitidus Günther, 1863 1863 Ahaetulla nitida Günther, Ann. Mag. Nat. Hist. (3) 11: 283-287. Specimens examined. NMK S/67, S/69. 138 Philipp WAGNER & Wolfgang BÖHME: The amphibians and reptiles of Kakamega Forest, western Kenya Remarks: The vouchers were not collected in the area close to Kakamega Forest but on the shores of the Yala River in the Siaya District. The river crosses the southern part of the Kakamega Forest (see P heterolepidotus above). After closer examination, NMK S/67 & S/69 were assigned to the subspecies P. n. loveridgei Laurent, 1960. Psammophis mossambicus Peters, 1882 1882 Psammophis mossambicus Peters, Reise nach Mossambique, p. 122. Specimens examined. NMK S/2316, S/2319; ZFMK 82049. Key references: SPAWLS et al. 2002. Remarks: Spawıs et al. (2002) placed the eastern popu- lations of P. phillipsi and P. sibilans in the synonymy of P. mossambicus. Examination of the type material of P mossambicus support this decision but it has to be kept in mind that the earlier recognized P. phillipsi is a rela- tively small sized rainforest species with an entire anal scale (STERNFELD 1907; LOVERIDGE 1940; VILLIERS 1975; TAYLOR & WEYER 1958; DouceT 1963; CHIPPAUX 2001). The voucher ZFMK 82049 was found inside Buyangu Vil- lage, basking on a small bush. Psammophis phillipsi (Hallowell, 1844) 1844 Coluber phillipsi Hallowell, Proc. Acad. Nat. Sci. Philadelphia 1844: 169. Specimens examined. NMK 0/3603; ZFMK 82050. Key references: CHIPPAUX 2001. Remarks: These vouchers represent the first record of the species for Kenya. For the taxonomic assignment see P. mossambicus. SPAWLS et al. (2002) placed the eastern pop- ulations of P. phillipsi and P. sibilans in the synonymy of Fig. 21. Psammophis phillipsi from Kakamega Forest. Photo by Jörn KÖHLER. P. mossambicus. ZFMK 82050 was found 100 m away from a pond inside the primary forest of the Buyangu area. The second voucher was collected on the Buyangu Hill, a natural clearing inside the forest. Psammophis rukwae Broadley, 1966 1966 Psammophis sibilans rukwae Broadley, Arnoldia Rhod. 2 (36): 3. Specimens examined. None. Remarks: This taxon is only known from literature (SPAWLS et al. 2002). Further data are not available. Fig. 22. Thrasops aethiopissa from Kakamega Forest. Photo by Philipp WAGNER. Thrasops aethiopissa (Günther, 1862) 1862 Rhamniophis aethiopissa Günther, Ann. Mag. nat. Hist. (3) 9: 124- 132. Specimens examined. NMK 0/3563; ZFMK 76045, 77290, 82032. Additional specimens. CAS 147909, 152794. Remarks: 7. aethiopissa is a typical rain forest species whose East African populations are assigned to the sub- species 7. a. elgonensis (LOVERIDGE, 1929) which is on- ly known from the Mt. Elgon and Kakamega forests in Kenya. ZFMK 82032 was collected at daytime basking on a 20 cm high bush on a clearance of the Buyangu area. When threatened it displays typical defence behaviour similar to the boomslang. Thrasops jacksonii Günther, 1895 1895 Thrasops jacksonii Günther, Ann. Mag. Nat. Hist. 15 (6): 523-529. Specimens examined. MHNG 1.375.038-040, 042; ZFMK 66275-76, 68516. Additional specimens. CAS 122295, 152795. Remarks: Details on the vouchers are unknown., Bonner zoologische Beiträge 55 (2006) 139 Toxicodryas blandingii (Hallowell, 1844) 1844 Dipsas blandingii Hallowell, Proc. Acad. Nat. Sci. Philad. 1844: 170. Specimens examined. MHNG 1.356.058. Additional specimens. CAS 150981-82. Remarks: In Kenya it is only known from Kakamega For- est and the Serem area. Further data are not available. Toxicodryas pulverulenta (Fischer, 1856) 1856 Dipsas pulverulenta Fischer, Abhandl. Nat. Ver. Hamburg 3 (4): 81-116. Specimens examined. None. Additional specimens. CAS 122742. Remarks: Further data are not available.. Atractaspididae Günther, 1858 Polemon christyi (Boulenger, 1903) 1903 Miodon christyi Boulenger, Ann. Nat. Hist. (7) xii : 354. Specimens examined. None. Additional specimens. CAS 147905. Remarks: Details on thıs voucher are unknown. Elapidae Boie, 1827 Dendroaspis jamesoni Traill, 1843 1843 Elaps jamesoni Traill, Edinburgh new. phil. J., 34 (67): 54. Specimens examined. MHNG 1327.046-48; ZFMK 82036. Additional specimens. CAS 122298-99. Key references: LOVERIDGE 1936. Fig. 23. Dendroaspis jamesoni kaimosae from Kakamega Forest. Photo by Philipp WAGNER. Remarks: D. jamesoni is distributed from Ghana in the west to Kenya in the east and southwards to Angola. In Kenya, the subspecies D. j. kaimosae Loveridge, 1936 (with the type locality Kaimosi) is hitherto recorded from Kenya by vouchers from Kakamega Forest, and a sight- ing from Lolgorien, Mara Escarpment (SPAWLS ct al. 2002). During the course of this study we found a vouch- er specimen (NHMW 28109) collected from Nakuru. ZFMK 82036 was found on the shore of the Isiukhu Riv- er in the northern part of the forest, basking on a horizon- tal branch at about midday. Elapsoidea loveridgei (Parker, 1949) 1949 Elapsoidea sundevalli loveridgei Parker, Zool. Verh. Leiden 6: 1- 115. Specimens examined. MHNG 1328.010; ZFMK 82001. Additional specimens. CAS 152796. Remarks: ZFMK 82001 was found in the morning after a rainy night; it was crossing the forest road near to the Buyangu gate. After examination of the vouchers they were assigned to the subspecies EF. /. multicincta (LAUREN- TI, 1956) however the nominate form E. /. loveridgei al- so occurs in Kenya east of the Rift Valley. Naja melanoleuca Hallowell, 1857 1857 Naja haie var. melanoleuca Hallowell, Proc. Acad. Nat. Sci. Philadelphia 1857: 61. Specimens examined. NMK S/3980, S/3985; ZFMK 82045-47. Additional specimens. CAS 122749, 122758. Remarks: N. melanoleuca 1s the most common venomous snake in the Kakamega area. Snakes were found in sev- eral habitats and areas, e. g. inside the forest: near the Isıukhu Falls, near the Buyangu Gate, south of Udo's Campside, Yala River, Kaimosi fragment and Malava frag- ment; outside the forest (mostly juveniles) within Buyan- gu Village, Isecheno and Salazar Circuit (both sightings). Pseudohaje goldii (Boulenger, 1895) 1895 Naia goldii Boulenger, Ann. Mag. Nat. Hist. London 16 (6): 34. Specimens examined. NMK S/3366 Remarks: This species has a wide but disjunct distribu- tion from Togo (HUGHES & BARRY 1969) in the west to Kenya in the east and southwards to Angola and Namib- ia (Ost et al. 1988). In Kenya it is only known from Kakamega Forest but the occurrence in the Mt. Elgon forests is possible. Further data are not available. 140 Philipp WAGNER & Wolfgang BÖHME: The amphibians and reptiles of Kakamega Forest, western Kenya Fig. 24. Atheris hispida from Kakamega Forest. Photo by Phil- ipp WAGNER. Viperidae LAURENTI, 1768 Atheris hispida LAURENT, 1955 1955 Atheris hispidus LAURENT, Rev. Zool. Bot. Afr. 51: 138. Specimens examined. MHNG 2236.23; NMK 0/1648; NMZB-UM 5317; ZFMK 66374, 82024-26. Additional specimens. CAS 141750, 147906, 147908. Remarks: This bush viper is only known in Kenya from the Kakamega Forest, where it is sympatric with 4. squamigera. SPAWLS et al. (2002) stated that A. hispida in- habits higher and drier bushes than 4. squamigera which does not seem to be the case in Kakamega Forest, where A. hispida was found mostly in low elevation of a maxi- mum of 80 cm. SPAWLS et al. (2002) also reported that the taxon was never collected in Kakamega since the early 1990's, so that this record is the rediscovery of the species there. Most vouchers were collected in the northern part of the forest. Only CAS 122747 was found in the Kaimosi fragment. Atheris squamigera (Hallowell, 1854) 1854 Echis squamigera Hallowell, Proc. Acad. Nat. Sci. Philadelphia 1854: 193. Specimens examined. PW 158 £ 163 (now part of the NMK collection); ZFMK 64337-42, 81214, 82027, 82029. Additional specimens. CAS 122744-46, 122748, 147902- 03, 148629, 150983-84, 153468; NMZB-UM 5393, 6518-19. Remarks: A. squamigera is a common viper inside Kakamega Forest, where it occurs sympatrically with 4. hispida. Most vouchers were collected in the northern part of the forest. Only DREWES collected two specimens 1969 near Kaimosi in the southern part of the forest. A. squamigera has a disjunct distribution from Ghana (HUGHES & BARRY 1969) in the west to Kenya in the east, southwards to Angola and Tanzania. Additionally studies may demonstrate differences between the populations. In Kenya A. squamigera is, with two exceptions, only known from Kakamega Forest. Two records are from outside the forest: one specimen from Chemilil and one sighting from the Soit Ololol Escarpment (SPAWLs et al. 2002). Speci- mens collected in this study were exclusively found near water bodies in the Buyangu area. They were found by hunting by torchlight and basking on small bushes or in leaf litter at daytime. Fig. 25. Atheris squamigera from Kakamega Forest. Photo by Jórn KÓHLER. Bitis gabonica (Dumeril, Bibron & Dumeril, 1854) 1854 Echidna gabonica Dumeril, Bibron & Dumeril, Erpét. Gén. 7: 1428. Specimens examined. NMK S/2904 Remarks: Details on this record are unknown, but sev- eral sightings of the taxon are reported from the grassland near the entrance of Udo's Campsite and from the Salazar Circuit. In Kenya this taxon is only known from the Kakamega and Nandi areas. Bitis nasicornis (Shaw, 1802) 1802 Coluber nasicornis Shaw, Nat. Misc. 3, pl. 94. Specimens examined. NMK S/3978; ZFMK 82030-31. Additional specimens. CAS 150989, 150990. Remarks: Like the previous species also B. nasicornis was often found and sighted on the roads of the Salazar Circuit. Also this taxon occurs within Kenya only at the Kakamega and Nandi areas. Two vouchers (ZFMK 82030 and 82031) were found near the Isiukhu Falls. The stom- ach of ZFMK 82031 contained (Muridae: Lophuromys lat- iceps) as prey item. Bonner zoologische Beiträge 55 (2006) 14] Causus lichtensteini (Jan, 1859) 1859 Aspideplaps lichtensteini Jan, Rev. & Mag. Zool. 1859: 511. Specimens examined. NMK S/2499; ZFMK 82033-34. Additional specimens. CAS 154579. Remarks: This forest species has a wide distribution from Cote d’Ivoire in the west (RÓDEL & MAHSBERG 2000) to Kenya in the east and southwards to Zambia (BROADLEY et al. 2003). In Kenya this taxon is only known from the Kakamega and Nandi areas. All vouchers of the study are juveniles and were collected after rainfall in the evening in the grassland near Udo's Campsite and in the morning at the forest border and in Guava bushland in the Buyan- gu area. Causus resimus (Peters, 1862) 1862 Heterophis resimus Peters, Monatsber. Akad. Wiss. Berlin 1862: 271-279. Specimens examined. None. 4.2. KEY TO THE SPECIES AMPHIBIANS 1 - Tongue absent. - Tongue present. 2 - Upper jaw toothless - Upper jaw with teeth. 3 - Fine dorsal skin ridge along midline. - Dorsal skin ridge absent. 4 - Last phalanx of fingers out of alignment. - Last phalanx of fingers not out of alignment. 5 - Parotoid glands distinct and raised. - Parotoid glands indistinct and flattened. 6 - Heels (tarsus) black with thin white border in males. - Heels lacking this pattern in males. 7 - Pupil horizontal to round. - Pupil vertical. 8 - Vomerine teeth absent. - Vomerine teeth present. 9 - Thin pale marking from the lower back around a dark rectangular patch anteriorly. - This particular pattern absent. Remarks: This taxon is only recorded from literature (PrT- MAN 1974). Further data are not available. Testudines Pelomedusidae Cope, 1868 Pelomedusa subrufa (Lacépede, 1788) 1788 Testudo subrufa Lacépede, Hist. nat. Quadrup. Ovip. Synops. Method. 1: 173, pl. 12. Specimens examined. NMK C/53; ZFMK 81951. Remarks: Interestingly this typical savannah species was also found inside the forest. Several adult specimens were found inside a pond in the Buyangu area. Juveniles were found in smallest water bodies inside and outside the for- est e. g. in small temporary stream on the Buyangu Hill. But P. subrufa was also found in the surrounding area, in fish and garden ponds. Specimens were also sighted on the shores of the Isiukhu River, but not at the Yala River. PIPIDAE Xenopus victorianus 2 BUFONIDAE (5) 3 ARTHROLEPTIDAE (6) 4 HYPEROLUDAE (7) RANIDAE (10) Amietophrynus kisoloensis Amietophrynus maculatus Leptopelis mackayi Leptopelis aff. bocagii Hyperolius 8 Afrixalus (9) Kassina Kassina senegalensis Afrixalus osorioi Afrixalus quadrivittatus 142 Philipp WAGNER & Wolfgang BÖHME: The amphibians and reptiles of Kakamega Forest, western Kenya 10 - Vomerine teeth absent. - Vomerine teeth present. 11 - Vomerine projections between internal nostrils. - Vomerine projections abutting onto anterior margins of internal nostrils 12 - Transverse skin groove behind the eyes. - Transverse skin groove absent. 13 - Golden to brownish band from snout to vent. - Band absent. 14 - Pale triangle on snout. - Pale triangle absent. 15 - Distance from nostril to snout greater than internarial distance. - Distance from nostril to snout not more than internarial distance. 16 - 2 to 2 1/3 phalanges of fourth toe free of web. - 2.5 phalanges or more on fourth toe free of web. 17 - Back of thigh spotted or mottled. - Back of thigh with and dark longitudinal bands. 18 - Tympanum visible. - Tympanum not visible. 19 - Discs on fingers absent. - Discs on fingers very small, only swellings. TERRAPINS 1 Only terrapin in Kakamega area LIZARD FAMILIES 1 - Head dorsally covered with granular, small and irregular scales. - Head dorsally covered with large scales. 2 - Eyelids absent. - Eyelids present. 3 - Head much wider than neck, with clusters of spiny scales around the ear; tongue short and broad. - Head only slightly wider than neck; tongue long and slender. 4 - Tongue very long and telescopic; digits fused together. - Tongue long and forked; digits separate. 3 - Dorsal and ventral scales similar, mostly smooth. - Ventral scales rectangular, larger than dorsals. 6 - Lateral granular fold present. - Lateral granular fold absent. Phrynobatrachus (18) 11 12 Ptychadena (14) Hoplobatrachus occipitalis 13 Amnirana albolabris Afrana angolensis 15 16 Ptychadena oxyrhynchus Ptychadena anchietae Ptychadena mascariensis 17, Ptychadena porossima Ptychadena taenioscelis 19 Phrynobatrachus aff. mababiensis Phrynobatrachus natalensis Phrynobatrachus graueri Pelomedusa subrufa 2 5 GEKKONIDAE 3 AGAMIDAE 4 CHAMAELEONIDAE VARANIDAE SCINCIDAE 6 LACERTIDAE CORDYLIDAE Chamaesaura anguina Bonner zoologische Beiträge 55 (2006) GEKKONIDAE 1 - Pupil round. - Pupil vertical. 2 - Claws on digits except thumb present. - Toes with a distinctive angle at the last or last two joints. 3 - Subcaudals with a continous median row; 9-12 preanal pores. - Subcaudals with a discontinous median row; 6-8 preanal pores. SCINCIDAE 1 - Eye covered by skin. - Eye visible. 2 - Lower eyelid with a large transparent disc. - Lower eyelid without a transparent disc. 3 - 2-3 digits on forelimb, 3 on hindlimb. - 4 or 5 digits on both limbs. 4 - Supranasals present, broadly in contact. - Supranasals absent, 1f present widely separated. 5 - Scales on feet usually non-spinose and smooth. - Scales on feet keeled and spinose. 6 - Midbody scale rows number 24-26. - Midbody scale rows number 28 or more. 7 - Midbody scale rows number 32-42; 5 black-bordered longitudinal stripes, blue tail in juveniles. - Midbody scale rows number 30-38, without such stripes. LACERTIDAE 1 - Midbody scale rows number 18-24. - Midbody scale rows number more than 35. AGAMIDAE 1 - Interparietal scale not larger than the adjoining head scales. - Interparietal scale larger than the adjoining head scales. CHAMAELEONIDAE 1 - Tail short and non-prehensile. - Tail long and prehensile. 2 - Body scalation heterogeneous. - Body scalation homogeneous. 3 - Occipital dermal lobes absent. - Occipital dermal lobes present. 4 - Long gular crest; parietal crest very high. - Low crest; one or two black coloured throat grooves. 2 Hemidactylus mabouia Lygodactylus gutturalis 3 Cnemaspis africana Cnemaspis elgonensis Feylinia currori 2 3 3 Eumecia anchietae 4 Lepidothyris aft. fernandi Afroablepharus wahlbergii 6 Trachylepis striata Trachylepis megalura 7 Trachylepis quinquetaeniata Trachylepis maculilabris Adolfus africanus Adolfus jacksoni Acanthocercus atricollis Agama kaimosae (>) Rhampholeon boulengeri 2 4 3 Chamaeleo laevigatus Chamaeleo gracilis Chamaeleo hoehnelii Chamaeleo ellioti 144 Philipp WAGNER & Wolfgang BÖHME: The amphibians and reptiles of Kakamega Forest, western Kenya SNAKE FAMILIES 1 - Eye covered by skin. - Eye not covered by skin. 2 - No poison fangs in the front upper jaw, pupil usually round. - Poison fangs in the front upper jaw present 3 - Poison fangs relatively short, immobile. - Poison fangs mobile or folding. 4 - Eye large, head dorsum covered with small scales (except Causus). - Eye tiny, head dorsum with large symmetrical scales. TYPHLOPIDAE 1 - Second supralabial overlapping preocular scale. - Second supralabial overlapping ocular scale. ELAPIDAE 1 - Midbody scales rows number 13, always with broad bands. - Midbody scale rows number more than 13. 2 - 3 preoculars scales, head long and narrow. - | or 2 preocular scales. 3 - Midbody scale rows number 15 (Rarely 17). - Midbody scale rows number 19. VIPERIDAE 1 - Pupil round, 9 large symmetrical scales on top of the head. - Pupil vertical, many scales on top of the head. 2 - Subcaudal scales paired. - Subcaudal scales single. 3 - Long horns on the snout of the adults. - No or only short horns on the snout. 4 - Scales strongly lanceolate on the head and front part of the body. - Scales not lanceolate. un - Subcaudal scales single. - Subcaudal scales paired. COLUBRIDAE 1 - No venom-delivery fangs in the upper jaw. - One or more pairs of venom-delivery fangs in the upper jaw. 2 - Pupil vertical. - Pupil round. 3 - Ventrals number 141-183. - Ventrals number 195-270. 4 - Midbody scale rows number 19. - Midbody scale rows number 21-25. TYPHLOPIDAE 2 COLUBRIDAE 3 ELAPIDAE 4 VIPERIDAE ATRACTASPIDIDAE Typhlops angolensis Typhlops lineolatus Elapsoidea loveridgei 2 Dendroaspis jamesoni Pseudohaje goldii Naja melanoleuca NN un 4 Bitis nasicornis Bitis gabonica Atheris hispida Atheris squamigera Causus lichtensteini Causus resimus [o 2) 5 Crotaphopeltis hotamboeia 4 Toxicodryas blandingii Toxicodryas pulverulenta 10 11 12 13 14 15 16 17 18 19 20 21 22 Bonner zoologische Beitráge 55 (2006) - Dorsal scales keeled. - Dorsal scales smooth. - First 5 infralabials usually in contact with the anterior sublinguals. - First 4 infralabials usually in contact with the anterior sublinguals. - Anal scale divided. - Anal scale single. - Dorsal scales smooth. - Dorsal scales keeled. - - Nostril (naris) located with a divided or semi-divided nasal. - Nostril (naris) located with an entire nasal. - Anal scale single. - Anal scale divided. - Midbody scale rows number 13-15, reducing to 11 posteriorly. - Midbody scale rows number 15-21, if 15 not reducing posteriorly. - A pair of enlarged occipital scales. - No enlarged occipital scales. - Anal scale single. - Anal scale divided. - Subcaudal scales rounded or angular, not keeled. - Subcaudal scales sharply angular, keeled. - Ventral scales 168-194, no concealed spots on dorsal scales. - Ventral scales 138-179, cryptic pattern with spots on dorsal scales. - Subcaudal scales 60—104, flanks often blue. - Subcaudal scales 88-128, no blue on the body. - Subcaudal scales 130-155. - Subcaudal scales less than 130. - Midbody scale rows number 15, not reduced before the vent. - Midbody scale rows number 15, reduced before the vent. - Ventral scales 153-173. - Ventral scales 170-221. - Midbody scale rows number 15. - Midbody scale rows number 21-27. - Ventral scales 193-244. - Ventral scales 150-175. - Fewer than 80 pattern cycles between nape and base of tail. - More than 80 pattern cycles between nape and base of tail or uniform black. Dispholidus typus 6 Psammophis rukwae 7 Psammophis mossambicus Psammophis phillipsi 9 20 10 18 Lamprophis fuliginosus 11 12 17 Thrasops aethiopissa 13 Philothamnus carinatus 14 15 Philothamnus nitidus Philothamnus heterolepidotus 16 Philothamnus hoplogaster Philothamnus battersbyi Thrasops jacksoni Natriciteres olivacea Lycophidion ornatum 19 Lycophidion depressirostre Lycophidion capense 21 22 Mehelya capensis Hapsidophrys lineatus Dasypeltis scabra Dasypeltis atra 146 Philipp WAGNER & Wolfgang BÖHME: The amphibians and reptiles of Kakamega Forest, western Kenya 5. DISCUSSION The composition of the herpetological community is typ- ical for a forest fragment. It includes typical forest species but also some ubiquitous and bushland species. From the 22 lizard species recorded, only seven are true forest species, and for five species Kakamega Forest is the on- ly Kenyan locality. The situation for snakes ıs nearly the same: from 26 species recorded, 14 are forest dwellers; of these, 12 again have their only Kenyan occurrence here. In addition, three ubiquitous species are also confined to Kenya to the Kakamega area. The species richness of snakes in this forest and its environs is remarkable on an African scale (for comparative species numbers of African snake communities see BÖHME (1993), HERRMANN et al. (2005b) and RÖDEL & MAHSBERG (2000). The rich- ness of forest snake species also argues for a relatıvely in- tact status of the main forest fragment which seems to be still relatively undisturbed, enabling survival for many for- est species. However, the remaining smaller fragments of Kakamega Forest have drastically reduced species richness as com- pared to the main fragment. From Malaba, the least ın- tact fragment, only two forest species, viz. the toad 4mi- etophrynus kisoloensis and the forest cobra Naja melanoleuca have been recorded. The Kisere fragment is known to have only one additional forest species, viz. Rampholeon boulengeri, but more forest species might be expected in this protected forest relict. The Kaimosi frag- ment, finally, is home for two bushviper (Atheris) species and the green mamba (Dendoaspis jamesoni kaimosae). This drastic decrease in numbers of forest-dwelling species underlines the need of effective conservation measures in order to stop any further deforestation and forest fragmen- tation. BENNUN & NJOROGE (1999) characterised Kakamega For- est as a mid-altitude tropical rainforest. This view is sup- ported by the occurrence of the (sub)montane tree fern Cy- athea manniana which is currently found only in the Nan- di Escarpment forests which lie a few hundred meters higher than Kakamega Forest. The same can be conclud- ed from the altitudinal distribution of the forest reptiles: of 21 forest species, 18 have a mid-altitudinal distribution, but this group is dominated by 11 species assignable to lowland forests. The assignment to a lowland rather than to a montane forest is also corroborated by the bird fau- na. BENNUN & NJOROGE (1999) found 194 forest species of which 40 were typical members of the Guineo-Congo- lian forest block. Of 134 typical Congo Basin bird species as defined by CHAPIN (1932), 37 occur in Kakamega For- est. This amount increases to 57 out of 125 when the species from lowland secondary forests and forest clear- ings are also taken into account. Studies of Kakamega’s Lepidoptera yielded similar results (CARCASSON 1964), and also the reptiles support the view of a mid-altitude rainforest dominated by lowland species. Several authors regarded Kakamega Forest as the eastern- most outlier of the Guinea-Congolian rainforest block (ZIMMERMANN 1972; DREWES 1976; HAMILTON 1976; ScHIOTZ 1976; KOKWARO 1988; VONESH 2001; KOHLER 2004; CLAUSNITZER 2005; SCHICK et al. 2005). The last mentioned authors have analysed the distribution of the 24 amphibian species recorded so far from Kakamega For- est. They assigned eight species to their distribution pat- tern type “East African Highland”, seven were widely dis- tributed forms including other parts of the equatorial rain- forests and even parts of southern Africa. Five species show a typical “arid corridor” distribution (see POYNTON 1995; P. WAGNER, unpubl. data), and only four species (Amnirana albolabris, Afrixalus osorioi, A. quadrivitta- tus, Hyperolius cinnamomeoventris) exhibit a typical Guinea-Congolian distribution pattern. In this regard, the amphibian fauna of Kakamega gives a less clear picture of the geographic assignment of this forest as compared with the reptilian fauna. Comparing the latter communi- ty with that of other African rainforests (WAGNER et al. submitted. ), it turns out that it shares many more species with the Bwindi and Kibale Forests in Uganda, and even with the far distant Ziama Forest in Guinea, West Africa than e. g. with the Kenyan coastal Arabuko-Sokoke For- est or even the Eastern Arc montane forests. The analy- sis of Kakamega's reptile fauna, therefore, clearly argues for a Guinea-Congolian assignment of the relictual Kakamega Forest. Acknowledgments. We are grateful to Jörn Köhler, the ex-co- ordinator of the BIOTA project, who made the study on the rep- tiles of Kakamega Forest possible. We thank Andreas Schmitz and Scott M. Moody for their critical and precious review. Gu- drun Schaab, Arne Schiotz, Jórn Kóhler, Stefan Lótters, Alexan- der Burmann, Mike Dobiey & Chimaira publishing contribute pictures of relevant species from the forest. We thank S. Schick, A. Schiotz, S. Lötters, P. Hita Garcia & W. Freund for contribut- img unpublished data and/or material. Special thanks to Claudia Fink for joining the senior author in the field and to Sylvester Shirandula and Caleb Analo for their grand help. Also special thanks to Patrick Malonza and Vincent Muchai (both NMK) for the fruitful co-operation. Very special thanks from the senior author to Steve Spawls for discussions on East African reptiles and his answers to so many questions. The senior author is indebted to his father, Rúdiger Wagner, for his paternal support of this study. We also thank the following sponsors: Deutsche Telekom, Fu- ji-Films, Henkel KGaA, Kodak Films, Siemens, Unilever, Var- ta AG and Volkswagen who supported the study in different ways. The study benefited from the support of the BIOLOG-BIOTA pro- gram of the German Federal Ministry of Education and Research (BMBF). We are indebted to the Kenyan partners, the National Museums of Kenya (NMK) and the Kenya Wildlife Service (K WS) who kindly issued facilities and permissions to carry out work in Kakamega Forest. Bonner zoologische Beiträge 55 (2006) 147 REFERENCES ANALO, C. I. 2003. Adventures of Kakamega Forest. BIOTA- East Africa E12, 35 pp. Bauer, A. 1993. African-South American relationships: A per- spective from the Reptilia. Pp. 245-288 in GOLDBLATT, P. (ed.) Biological Relationships Between Africa and South America. Yale University Press, New Haven, Connecticut. BENNUN, L. & NJOROGE, P. 1999. Important Bird areas in Kenya. 318 pp., East Africa Natural History Society, Nairobi. BÖHME, W. 1993. Mission d’études herpétologique dans les fo- réts de Ziama et Diécké Guinée forestiere. 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WAGNER, P., SCHMITZ, A., PAUWELS, O.S.G & BOHME, W. subm. Geographic variation in Mochlus fernandi BURTON, 1836 (Squamata: Scincidae), with the description of two new taxa and two revalidations. Zootaxa Wass, P. (editor) 1995. Kenya’s Indigenous Forests: Status, Man- agement and Conservation. IUCN, Gland & Cambridge. ZIMMERMANN, D. 1972. The avifauna of the Kakamega Forest, western Kenya, including a bird population study. Bulletin of the American Museum of Natural History 149: 257-339. Authors’ address: Philipp WAGNER (corresponding author), Wolfgang BOHME, Zoologisches Forschungsmuseum Alexander Koenig, Adenauerallee 160, D-53113 Bonn, Germany; E-Mail: philipp.wagner.zfmk@uni-bonn.de; w.boehme.zfmk@uni-bonn.de Received: 27.06.2007 Accepted: 03.07.2007 Corresponding editor: M. Schmitt Bonner zoologische Beiträge 55 (2006) APPENDIX Ecological characteristics of the 58 reptiles recorded within the area of the Kakamega Forest Taxon Abun2 Locb Diete Habd Acte Micro! Testudinae Pelomedusa subrufa U 2 LA,B W,L D Aq Sauria, Gekkonidae Cnemaspis africana I 2 I FI N A Cnemaspis elgonensis I 2 I FI N A Hemidactylus mabouia C 1 I F N A,AP Lygodactylus gutturalis R 2,5 I FE DN A Agamidae Acanthocercus atricollis € | I ASF D A, AP Agama kaimosae R 7 I L D Chamaeleonidae Chamaeleo ellioti C 1,2,3,4,6,7, I A,FE, D AL 10,11,13 FFI Chamaeleo gracilis I 10 I L D AL Chamaeleo laevigatus I 12 I L D AL Rhampholeon boulengeri U 2, 16 I FI, FE D L, AL Scincidae Afroablepharus wahlbergii R 9,12 I A,EL D L,T Eumecia anchietae U 1,3 I L, ¡E D E Feylinia currori S 2 I FE ? B Lepidothyris aff. fernandi R 2 I FE,. FI L, T Trachylepis maculilabris R 7 I A,FE D LT Trachylepis megalura R 6,7 I A, FE D L, T Trachylepis quinquetaeniata € 13 I L D T Trachylepis striata € 1,5 I A, FE, F D L, T Lacertidae Adolfus africanus I 2 I FI, FE D A, L Adolfus jacksoni € 1,5 I BE, A, E D T; E Cordylidae Chamaesaura anguina R 14 I L D UN Varanidae Varanus niloticus R 9 M,Ga,F, A, EL D I: A,B,l Serpentes, Typhlopidae Typhlops angolensis R 7 I A, FE N B Typhlops lineolatus U 25 I E, FE N B Colubridae Crotaphopeltis hotamboeia R 14 A FE, A N T, Aq Dasypeltis atra U 2,14 B FI, FL,L N TA Dasypeltis scabra U 7 B El, EL, L;F N T, A Dispholidus typus R A,Ro,B,E FE, LF D A, AL, AH Hapsidophrys lineatus I 2,14 A FE, El DN? A Lamprophis fuliginosus € Ro A, F N T Lycophidion capense R 7 E,S L,A N T Lycophidion depressirostre R 7 E,S L N T Lycophidion ornatum I 2 E FI, FE N E Mehelya capensis R 14 E,S FEEL N T Natriciteres olivacea R 4 A,F FE, A DN T, Aq Philothamnus battersbyi U 1,14 A,F E-W E D AL Philothamnus carinatus R 1 A FI, FE DN? A Philothamnus heterolepidotus R 15 A L D A Philothamnus hoplogaster R 14, 10 A,F L, W D A Philothamnus nitidus R 15, 14 A FE, FL D A Psammophis mossambicus I 14,1 S,E,A,B,Ro L,F,A D TT Psammophis phillipsi I 2 M,E,A,B FE D A Psammophis rukwae R 7,8 E,S LEA D AR Thrasops aethiopissa U 2 A FI, FE DN? A Alte 0-1600 0-2000 1200-2200 0-2500 700-2000 0-2400 1800 1500-2800 0-1600 1000-1500 1400-2000 0-2200 600-2100 0-2300 1500-3000 200-1600 0-2300 580-1200 450-3000 1200-3000 0-1600 0-2500 1000-2800 0-2600 0-2200 700-1800 0-2400 0-2400 700-2700 0-2000 0-2200 1300-1800 0-2300 600-2000 0-1800 medium 0-2500 medium medium medium 149 Distr." SSA CE SSA MA SSA SSA SSA WE 150 Philipp WAGNER & Wolfgang BÖHME: The amphibians and reptiles of Kakamega Forest, western Kenya Taxon Abun* Loch Diet“ Habd Act Micro! Alte Distr.h Thrasops jacksonii U 2,14 M,E,B,A FE, FL A DN A 600-2400 CE Toxicodryas blandingii R 14 B,E,A,M FE; EI DN A 700-2200 WE Toxicodryas pulverulenta R 7 Ro,E Fl, FE N A medium WE Atractaspididae Polemon christyi R 14 E FE, FI, A N? B 600-1700 CE Elapidae Dendroaspis jamesoni U 14,2 Ro,B FE, F],A N A,T 600-2200 GE Elapsoidea loveridgei I 14,2 S,E,A,Ro FE, L, F N T 600-2200 CE Naja melanoleuca E 14,11 A,F,S,E,M,B FE, FI DN T 0-2500 SSA Pseudohaje goldii I 14 A,F Fl N A 600-1700 WE Viperidae Causus lichtensteini U 14,5 A FI, FE DN T 500-2100 WE Causus resimus R 7 A FE, E, E DN iE 0-1800 WE Atheris hispida U 2,14 Ga,A,Ro FE, W N A 900-2400 CE Atheris squamigera U 2,4,14,13,7 E,S,A,M FL FE N A 700-1700 WE Bitis nasicornis U 4,3,14 M Fl, FE; N T, AL 600-2400 CE Bitis gabonica I N T 0-2100 CE 14 M FE, L Remarks: a - Abundance, based on the number of records: C = very common; U = common, often sighted within the study and many vouchers collected; I = rare, only seen once or twice, only single vouchers; R = rare, only from literature or single vouchers; E= proposed occurrence. b - Localities of vouchers, literature records and sightings: 1= Buyangu Village: 2= Buyangu forest; 3= Salazar Circuit; 4= Isiukhu River; 5= Udo's Campside; 6= Liranda Hill; 7=Kaimosi; 8= reported by Spawls er al. 2002; 9= reported by Loveridge; 10= Nandis; 11= Malava; 12= vicinity of Kakamega Forest; 13 = Buyangu clearing; 14= Kakamega Forest in general; 15= Yala River; 16= Kisere. c - Diet, A= amphibians, I= insects, Ga= gastropods, F= fishes, E= lizards, S= snakes, B= birds, M= mammals, Ro = rodents. d - Habitats, FI= forest; FE= clearings or forest edges; A= agricultural land; F= Farmland, residential area and gardens; W= waters or shores; L= Bushland and open landscape. e - Activity, D = day active; N = night active; DN = diurnal. f- Microhabitat, Aq= aquatic; B= burrowing; L= leaf litter; T= terrestrial; A= arboricol; AL= arboricol in low vegetation; AH= arboricol in trees; AP= anthropophilous. - Altitude in metres. - Distribution, East to west distribution in Africa: key references: Pitman (1974), Spawls er al. (2002), Broadley & Howell (1991), Uetz (2002) and Chippaux (2001): CE= central to eastern Africa; CS= central to southern Africa; EA= East Africa; K= Kenya; MA= middle of Africa; NA= northern Africa; NEA= north-eastern Africa; OS= East- to South Africa; SSA= Sub-Saharan Africa; WE= West- to East Africa. go So h Bonner zoologische Beiträge Band 55 (2006) Heft 2 Seiten 151-157 Bonn, Juli 2007 A new Meligethes of the M. pubescens species-group from South Africa (Coleoptera, Nitidulidae, Meligethinae) Paolo AubIsIo & Alessio DE BIASE Rome, Italy Abstract. A new South African species (from Western Cape Province) of the recently revised Meligethes pubescens spe- cies-group is described as M. colonnellii n. sp., an updated key to the identification of the known species of the group is provided, and new data are added on the geographic distribution of the recently described M. hermanniae Audisio, Kirk- Spriggs & Kirejtshuk. Keywords. Coleoptera, Nitidulidae, Meligethes, new species, South Africa, Hermannia, Sterculiaceae. JE INTRODUCTION Recent years have seen the description of a number of new taxa within the Meligethes pubescens species-group from Southern Africa (SPORNRAFT & AUDISIO 1995; KIREJTSHUK 1996; Aupisıo et al. 1998), and a revision of the group was recently published (Aupisıo et al. 1998). This paper mainly deals with the description of a new South African species more closely related to Meligethes confertus Reitter, 1872, and to M. aurivestis Audisio, Kirk- Spriggs & Kirejtshuk, 1998, and includes an updated key to the identification of the known species of the group: both form part of a coming revision of the Southern African species of the subfamily Meligethinae as a whole (P. AUDISIO, in preparation). 2. MATERIALS The new species described herein was collected during two recent fieldtrips to South Africa (April-May, and Oc- tober, 2005) organised by the Dept. of Animal and Hu- man Biology of the Rome University “La Sapienza”; the studied material is mostly preserved in P.Audisio’s collec- tion, Rome (CAR), in the Albany Museum, Grahamstown (AMG) and in the Transvaal Museum, Pretoria (TMSA). Other comparative material has been borrowed for study from several South African and Namibian institutions, in- cluding the National Collection of Insects, Plant Protec- tion Institute, Pretoria (SANC), the South African Muse- um, Cape Town (SAMC), the National Museum, Bloem- fontein (NMBS), the Department of Entomology and Ne- matology, University of Stellenbosch (USSA), and the National Museum of Namibia, Windhoek (NMNW ); much other (mostly unnamed) material, and the type material of all previously described species have kindly been made available for study by various European institutions, such as the Natural History Museum, London (BMNH), the Museum National d Histoire Naturelle, Paris (MHNP), the Zoological Museum of the University, Lund (MZLU), the Museum fir Naturkunde, Humboldt University, Berlin (ZMHB), the Museum Alexander Koenig, Bonn (MAKB), the Zoological Museum of the Russian Academy of Sci- ences, St. Petersburg (ZMAS), the Zoologischen Staat- sammlung München (ZSSM) and the Museo Civico di Zo- ologia, Rome (MCZR). 3. DESCRIPTION OF THE NEW SPECIES Meligethes colonnellii n. sp. (Figs 1-3) Diagnosis. Medium-sized (length 2.8-2.9 mm), black species with orange brown legs and antennae (outer edges of all tibiae, and antennal scapes and clubs, slightly dark- er), with tarsal claws strongly toothed. In general appear- ance (Fig. 3) similar to M. aurivestis Audisio, Kirk-Sprig- gs & Kirejtshuk, 1998 (Aupisio et al. 1998), but slightly narrower and less convex, without distinct metallic cop- pery reflections on pronotum and elytra, with finer and shorter silvery-golden dorsal pubescence, and with amply distinct male genitalia. M. confertus Reitter, 1872, recent- ly re-described by Spornraft & Kirejtshuk (1993 ) and by AUDISIO et al. (1998), shows less developed elytral trans- verse strigosity, wider and more convex body shape, and again amply distinct male genitalia. M. translatus Grou- velle, 1913 (REITTER 1872; GROUVELLE 1913), recently re- described by SPORNRAFT & KIREJTSHUK (1993 ) and by Aupisıo et al. (1998), shows less developed elytral trans- 152 Paolo AUDISIO & Alessio DE BIASE: A new Meligethes of the M. pubescens species-group from South Africa | Figs 1-2. Tegmen and aedeagus (dorsal aspect) of Meligethes colonnellii n.sp. (male holotype from Western Cape Province, 20 km S Oudtshoorn). Scale bar = 0.20 mm. verse strigosity, longer golden pubescence, and again am- ply distinct male genitalia. M. pubescens Reitter, 1872 eventually shows strongly modified posterior tibiae in males, much less developed elytral transverse strigosity, and different male genitalia. Description. Male. Length 2.76 mm; breadth (at widest point of elytra) 1.30 mm. Moderately elongate, oval, con- vex (Fig. 3); black, with moderately developed silver-gold- en pubescence. Legs orange brown, except for dorsal out- er edges of tibiae, which are slightly darker, nut-brown. Antennae orange brown, with scape and club slightly dark- ci Head with dorsal punctures as large as eye facets, rather strongly impressed, separated by less than one diameter, surface between them smooth; front margin of clypeus completely and very distinctly bordered, widely and shal- lowly emarginate with obtusely rounded side angles. Fron- to-geneal grooves very distinct and complete. Antennae of normal size (Fig. 3), third antennal segment slender, as long as second one; antennal club medium-sized. Pronotum 1.54 times as wide as long (Fig. 3), broadest at posterior third, narrower anteriorly; sides very narrowly bordered, not explanate; posterior angles obtuse but dis- tinct. Posterior base not sinuate on either side of scutel- lum; punctures as on head, surface between them smooth and shining. Scutellum medium-sized, finely and sparsely punctate in posterior half; surface showing a faint trace of microscop- ic reticulation. Elytra 1.08 times as long as wide, broad- est at basal second fifth, scarcely wider than pronotum (1.07 times); shoulders feebly raised, humeral striae al- most absent; elytral punctures as on head and pronotum, but slightly longer and closer, each joined to its lateral neighbours by rather strongly raised aciculation, so giv- ing rise to a markedly distinct transrugosity, surface be- tween them smooth. Ventral surface black, with fine short pubescence. Proster- nal antennal furrows strongly marked. Prosternal process rather long and narrow in the middle (here scarcely as wide as antennal club), with much wider and flatly rounded Bonner zoologische Beiträge 55 (2006) 15 O A SS Don SI Fig. 3. Habitus (dorsal aspect) of Meligethes colonnellii n.sp. (male holotype from Western Cape Province, 20 km S Oudts- hoorn). Scale bar = | mm. apex, 1.4 times as wide as antennal club; punctures near- ly as on head; surface smooth and shining. Mesosternum with hind edge straight. Metasternum rather flat (punctures and the spaces between as on head and pronotum), with a couple of small and barely distinct tubercles at its ante- rior two-fifths, and with a shallow longitudinal impression on its posterior two-thirds, slightly widened posteriorly. The caudal marginal line of the hind coxal cavity follows closely its posterior edge, turning back just before its out- er end. Last visible abdominal sternite with its hind edge slightly and widely curving forward medially, here with a flattened, scarcely raised, but markedly shining tuber- cle. Anterior tibiae (Fig. 3) shaped nearly as in M. translatus (Fig. 10 in Aupisio et al. 1998), with their outer edges fine- ly crenulate from basal third, then with a subapical group of 4-5 progressively larger and slightly sharper teeth, sep- arated by one smaller tooth from the narrow subtarsal plate, but the distal teeth are of reduced size and less sharp, if compared with those of M. translatus, M. aurivestis, and M. confertus; anterior tarsi as wide as antennal club; in- termediate and posterior tibiae with their inner edges nar- rowly bordered; posterior tibiae relatively narrow, their in- Ww ner edges not sinuate (Fig. 3); all tarsal claws strongly toothed. Genitalia. Tegmen (Fig. 1) with apex of paramera obtuse- ly rounded, scarcely pubescent, and with median excision relatively small, U-shaped; aedeagus elongate (Fig. 2), strongly narrowed from distal sixth, with narrowly trun- cate apex. Female. Metasternum rather flat (punctures as in male holotype), without tubercles at its anterior two-fifths, and without longitudinal impression on its posterior two-thirds. Ovipositor shaped as in M. confertus Reitter and M. trans- latus Grouvelle (Fig 46 in Audisio et al., 1998), moder- ately sclerotized. Type material:- Holotype, 7, Republic of South Africa: Western Cape Province, Little Karoo, 20 Km S Oudt- shoorn (R328), road between Oudtshoorn and the Robin- son Pass, 400 m, 339.49”.55”S, 22°.07’.13”E, 6.v.2005, P. Audisio leg., on Asparagus sp. (Liliaceae) (AMG). Paratypes, | 9, 1 female; South Africa: Western Cape, Swartberg Mts, Seweweekspoort, NW Calitzdorp, 700 m, 33°25’ .16°S, 21%.23”,50”E, 14.x.2005, P. Audisio, M. Biondi & A. De Biase leg., by sweeping on riverine veg- etation, | Y (CAR); Western Cape Province, Langeberg Mts, road between Ashton and Montagu, Kogmanskloof, 180 m, 33°.49°.26”S, 20°.05’.25”E, 9.x.2005, P. Audisio, M. Biondi & A. De Biase leg., by sweeping on riverine vegetation, | female (CAR). Comparative notes. M. colonnellii n. sp. is rather close- ly related to M. aurivestis Audisio, Kirk-Spriggs & Kire- jtshuk, M. confertus Reitter and M. translatus Grouvelle, from which it differs by the characters given in the diag- nosis and in the following keys. Easily recognizable also from other related members of the genus, such as M. pu- bescens Reitter and M. hermanniae Audisio, Kirk-Sprig- gs & Kirejtshuk, by the not sinuate inner edges of the pos- terior tibiae, and the strongly differing male genitalia. Biological notes. The type specimen of the new species was collected in late Autumn (May) by sweeping in dis- turbed fynbos from flowers of an unidentified Asparagus sp. (Liliaceae) with white flowers, which 1s clearly not its true larval host-plant. The above listed paratypes have been both collected by sweeping on riverine herbaceous vegetation. Larvae very likely develop in late Summer/Au- tumn on certain hygrophilous species of the genus Her- mannia (Sterculiaceae; VERDOORN 1980; ARNOLD & DE Wer 1993), given that all its closely related species are associated with members of this botanical genus (AUDI- sio et al. 1998). The holotype and the two paratypes were collected in company of several specimens of the com- mon and widespread Meligethes pubescens Reitter. 154 Paolo Aupisio & Alessio DE BIASE: A new Meligethes of the M. pubescens species-group from South Africa Figs 4. Known geographic distribution of Meligethes colonnellii n.sp. (squares), M. hermanniae Audisio, Kirk-Spriggs & Kirejts- huk (circles), and of M. aurivestis Audisio, Kirk-Spriggs & Kirejtshuk (triangles). Geographic distribution. So far only known from the above mentioned localities in the Western Cape Province (Fig. 4); this species 1s probably more widely distributed throughout the mountain areas bordering the Little Karoo and the south eastern edge of the Great Karoo. Etymology. This species is so named from our colleague and friend Enzo Colonnelli (Rome), renowned specialist in Curculionidae Ceutorhynchinae, in whose company the first known specimen of the new species was collected by the senior author. 4. ADDITIONAL NOTES Meligethes hermanniae Audisio, Kirk-Spriggs & Kirejt- shuk, 1998 Examined material: South Africa, Eastern Cape Province, 10 km W Kareedouw, 600 m, 33°.55°.41”S, 24°.14°.32”E, 5.v.2005, E. Colonnelli leg., on flowering bush of Protea sp. (Proteaceae), 1 Y (CAR). This species, at larval stage associated with Hermannia salviifolia L. fil. (Sterculiaceae), was known so far only from two coastal localities of the eastern areas of the West- ern Cape Province (AUDISIO et al. 1998; fig. 4). The above reported new locality significatively extends eastwards and inwards the geographic range of this species. The men- tioned male specimen from Kareedouw area is peculiar- ly large, reaching 3.0 mm in length. 5. UPDATED KEY TO THE SPECIES OF THE MELIGETHES PUBESCENS SPECIES-GROUP Refer to Figs 1-71 of the Aupisio et al. (1998) revision (indicated in italic) and to Figs 1-5 of the present paper (in- dicated in bold). 1 (2) Fronto-geneal and periocular grooves on head absent.(as in fig: 53) aces ne vaaae acne sds 685m eee 3 Bonner zoologische Beiträge 55 (2006) 155 2 (1) Fronto-geneal and periocular grooves on head distinct (as in fig. 34) ........oooooooooooooooooomoooo.. 9 3 (4) Tarsal claws with a well-formed, large and sharp tooth at base (as in Fig. 55). Male genitalia with aedeagus pe- culiarly long and narrow (Figs 30-32); ovipositor peculiarly small and unsclerotized (Fig. 51). Anterior tibiae with minute and irregular teeth on their distal outer edges (Fig. 16). Eastern Cape Province, Natal, southern Mpumalanga (= Eastern: Transvaal) (Fig: 63) . 2.0: 2.220.220 82222 15. marshalli Grouvelle 4 (3) Tarsal claws with a virtually undeveloped tooth at base (as in fig. 57), or with a blunt tooth at base (as in fig. 56). Male genitalia with aedeagus considerably less elongate (Figs 43-44, 68-71), ovipositor larger (Fig. 50), valer Et (isa) itn re Se Rede ae OS aes ERE Ee ia 5 (8) Tarsal claws with a blunt tooth at base (as in fig. 56). Anterior tibiae with a series of 5/10 sharp teeth placed in their distal half or two-thirds, an intermediate one and the penultimate usually being the largest (as in fig. 13). Body usually brown to blackish, with yellowish to orange legs and antennae, sparse and more or less conspic- uous silvery, or golden pubescence, and elytra paler, or with a large reddish discal spot .................- 6 6 (7) Dorsal and ventral pubescence usually silvery, more conspicuous, each hair being moderately long, but of un- usually large diameter. Male genitalia smaller, shaped as in figs 70-71. Ovipositor as in the following species (Fig. 50). Northern and eastern Namibia, Botswana, northern parts of South Africa (Fig. 64) RR Be ee Roce a mow 12. rufofuscus Audisio, Kirk-Spriggs & Kirejtshuk 7 (6) Dorsal and ventral pubescence golden, less conspicuous, each hair being moderately long and fine. Male geni- talia larger, shaped as in figs 68-69. Ovipositor as in fig. 50. Central and southern Namibia, Namaqualand (Fig. 4) oc. oon ah BRC io es o a ele 13. eremita Audisio, Kirk-Spriggs & Kirejtshuk 8 (5) Tarsal claws lacking a distinct tooth at base (as in fig. 57). Anterior tibiae with a series of small, irregular and usually blunt teeth on their distal outer edges (Fig. 17). Body unicolorous blackish-brown, with legs and anten- nae brown to blackish. Male genitalia as in figs 43—44. Ovipositor as in fig. 52. NE Eastern Cape Province, south- ennaksywarzulu-Natal (Fig. 61)... 2002... 200er 14. argentarius Audisio, Kirk-Spriggs & Kirejtshuk 9 (10) Tarsal claws simple, lacking a distinct tooth at base (as in Fig. 57) ........oooooooomoonommmmmmmoo... 11 10 (9) Tarsal claws with a well-formed, large and sharp tooth at base (as in Fig. 55) 2... 0... cece 19 11 (12) Last segment of the antennal club peculiarly large (Fig. 2/). Body peculiarly large, short and convex (Fig. 65). Anterior tibiae as in fig. 18. Male genitalia as in figs 66-67. Northern Transvaal (Fig. 62). 11. massivus Audisio, Kirk-Spriggs & Kirejtshuk 12 (11) Last segment of the antennal club much smaller, and normally shaped (Figs I-5) ................... 13 13 (14) Dorsal pubescence golden or silvery, scarcely long and dense, never obscuring upper surface. Outer edges of anterior tibiae with a series of 7/12 sharp and usually irregular teeth, never regularly increasing in size from the imst@torthe penultimate (Figs 14=15)..(TheM. fuerschi-complex) 2.2 .0s242060420+ 2100 RR 15 14 (13) Dorsal pubescence golden or silvery, peculiarly long and dense, partially obscuring upper surface (in unrubbed specimens). Outer edges of anterior tibiae with a series of 7/10 more or less sharp teeth placed in their distal three-fifths, usually slightly and regularly increasing in size from the first to the penultimate, which as a rule, is thedlarsestiimigsJ1—12)) (ihe M: plimbeus-Complex) nn: a A 17 15 (16) Elytra strongly transversely strigose. Male genitalia with a very small and shallow distal excision on tegmen (Fig. 26) and moderately pointed aedeagus (Fig. 27). Ovipositor slightly more pointed (Fig. 49). Southern parts of the Western Cape Province, eastern Free State (Fig. 62) ....10. pecten Audisio, Kirk-Spriggs & Kirejtshuk 16 (15) Elytra less strongly transversely strigose. Male genitalia with much deeper and narrow distal excision on tegmen (Fig. 28), and subtruncate aedeagal apex (Fig. 29). Ovipositor slightly less pointed (Fig. 48). Eastern parts of 156 Paolo Aupisio & Alessio DE BIASE: A new Meligethes of the M. pubescens species-group from South Africa the Eastern Cape Province and of the Free State, Kwa Zulu-Natal, and Mpumalanga (Fig. 62). Sn in Bee a Se aa Re wie pri ee LEN 9. fuerschi Spornraft & Audisio 17 (18) Metasternum flat, not impressed in both sexes. Male genitalia much shorter (Figs 35-36). Ovipositor as in fig. 47. Western parts of the Western Cape Province (Fig: 59) „u... 2:00, 2.1.00 8. plumbeus Reitter 18 (17) Metasternum shallowly impressed in males. Male genitalia much longer and differently shaped (Figs 33-34). Ovipositor as in M. plumbeus (fig. 47). Western parts of the Western Cape Province, southern Namibia (Fig. 61) Sa BAD ota aes road dre a 7. namakwaensis Audisio, Kirk-Spriggs & Kirejtshuk 19 (20) Inner edges of posterior tibiae markedly sinuate at distal third in both sexes (as in fig. 19). (The M. pubes- CERS-COMPIEX) asirios A EA A 21 20 (19) Inner edges of posterior tibiae not sinuate at distal third in both sexes (as in fig. 20 and in fig. 3). (The M. CONFEFIUS-COMPIEX) 52... en Hu ee en aa ee a SO 23 21 (22) Male genitalia much longer and larger, with median distal excision of tegmen more rounded basally (Figs 24-25). Body on average larger (length: 2.3-3.0 mm). Eastern coastal and subcoastal areas of the Western Cape Province and of the western portion of the Eastern Cape Province (Fig. 4) ae ee En as 2. hermanniae Audisio, Kirk-Spriggs & Kirejtshuk 22 (21) Male genitalia much shorter and smaller, with median distal excision of tegmen V-shaped basally (Figs 22-23). Body on average smaller (length: 2.2-2.8 mm). Southern parts of the Western Cape Province and SW parts of the Basten Cape Province (FE: 59) ae ae ee SR 1. pubescens Reitter 23 (24) Pronotum more distinctly narrower than elytra (Fig. 4); pronotum and elytra partially covered by usually gold- en pubescence. Elytra without, or with only a faint trace of transverse strigosity. Male genitalia with distal apex of tegmen with deep and wide V-shaped excision (Figs 37-38). Subdesertic inner parts of the Western Cape Province and extreme West of the Northern Cape Province (Fig. 60) ............... 3. translatus Grouvelle 24 (23) Pronotum nearly as wide as elytra (as in fig. 3 and 3); pronotum and elytra partially covered by usually sil- very, silvery-golden, or olivaceous pubescence. Elytra with distinct traces of, or with strong, transverse strigos- ity. Male genitalia differently shaped (Figs 39-42; Figs 1-2) .........ooooooomocoorrorrrnc een 25 25 (26) Usually only with trace of elytral transverse strigosity. Male genitalia differently shaped, with distally wide- ly truncate median lobe of aedeagus, but without distally subtruncate tegmen (Figs 39-40). Body lacking a faint coppery reflection. Southern parts of the Western Cape Province and SW parts of the Eastern Cape Province (ETE-OON nan ae a pared be Na eats ANT 4. confertus Reitter 26 (25) Elytra with strong transverse strigosity. Male genitalia with very peculiarly shaped and distally truncate tegmen (Fig. 41), or with median lobe of the aedeagus much more narrowly pointed at the apex (Fig. 2) .........27 27 (28) Body with a faint coppery reflection, especially dorsally. Dorsal pubescence silvery-olivaceous, longer and more developed. Male genitalia with much more widely truncate apex of the median lobe of aedeagus, com- bined with a subtruncate apex of tegmen, with short U-shaped median excision of the latter (Figs 41-42). Mon- tane areas of the SW Western Cape Province (Fig. 4) .......5. aurivestis Audisio, Kirk-Spriggs & Kirejtshuk 28 (27) Body without coppery reflection. Dorsal pubescence silvery-golden, finer and less developed. Male genitalia with much more narrowly truncate apex of the median lobe of aedeagus, combined with roundly obtuse apex of paramera, with short U-shaped median excision of tegmen (Figs 1-2). Montane areas of the SE Western Cape Province(Hie:4). aus Grey slaty hives wee are Sid edited ee es Sil 6. colonnellii n.sp. Bonner zoologische Beitráge 55 (2006) 157 Acknowledgements. Sincere thanks are due to numerous col- leagues, who enabled us to study the valuable material preserved in their respective institutions: the late friend S. Endródy-Youn- ga and the colleagues J. Harrison and R. Müller (TMSA), A. Kirk-Spriggs (AMG), B. Grobbelaar (SANC), H. Robertson and M.A. Cochrane (SAMC), J. Irish (NMBS), S. Louw (Department of Zoology, University of Bloemfontein), G.H. Giliomee (US- SA), E. Marais (NMNW); M. Schmitt (MAKB), R. Danielsson (ZMLU), M.D. Kerley (BMNH), N. Berti (MHNP), A.G. Kire- jtshuk (ZMSP), M. Jäch (NMW), B. Viklund (SMNH), M. Uh- lig (MHB), V. Vomero (MZR). We are also grateful to K. Sporn- raft (Penzberg; CSP and ZSSM), M.A. Bologna (Rome), M.Biondi (L’ Aquila, Italy), E. Colonnelli (Rome), and S. Zola (Milan), for having provided us with material collected during entomological expeditions to South Africa. This paper was supported by grants from Ministero del- l’Istruzione, dell’ Universita e della Ricerca (PRIN 2004057217 “Zoogeography of Mediterranean-southern African disjunct dis- tributions by a multimethod approach”) and from University of Rome “La Sapienza” (60 % funds ”Aspetti genetici e morfo- metrici della biodiversita animale in aree africane e medio-ori- entali a basso impatto antropico”). REFERENCES ARNOLD, J. H. & DE Wer, B. C. (1993): Plants of southern Africa: names and distribution. Memoirs of the Botanical Sur- vey of South Africa 62. National Botanical Institute, Preto- ria, 825 pp. AUDISIO, P., KIRK-SPRIGGS, A. & KIREJTSHUK, A. G. (1998): The Meligethes of the M. pubescens species-group from Southern Africa (Coleoptera: Nitidulidae, Meligethinae). En- tomologica scandinavica 29: 169-198. GROUVELLE, A. (1913): Famille des Nitidulidae. Notes syn- onymiques et rectifications a la nomenclature. Annales de la Société Entomologique de France 81 (1912): 387-400. KIREJTSHUK, A. G. (1996): Some results of Study on the Nitidul- idae from Namibia and Adjacent Territories. Part | (Coleoptera, Cucujoidea, Nitidulidae). Mitteilungen aus dem Zoologischen Museum in Berlin 72 (1): 21-52. REITTER, E. (1872): Die südafrikanischen Arten der Nitidulinen- Gattung Meligethes nach dem Materiale der Herren Chevro- lat, Dr. Fritsch und Anderer. Berliner Entomologische Zeitschrift 16: 241-264. SPORNRAFT , K. & Aupisio, P. (1995): Ein weiterer neuer Meligethes aus Südafrika (Coleoptera, Nitidulidae). Nachricht- enblatt der Bayerischen Entomologen 44 (3-4): 69-73. SPORNRAFT ,K. & KIREÍTSHUK, A. G. (1993): Über alte und neue südafrikanische Meligethes-Arten (Coleoptera, Nitidulidae). Mitteilungen der Münchener Entomologischen Gesellschaft 83: 47-75. VERDOORN, 1. C. (1980): Revision of Hermannia subgenus Her- mannia in southern Africa. Bothalia 13 (1-2): 1-63. Authors” address: Paolo AUDISIO (corresponding author, E-Mail: paolo.audisio@@uniromal.it) & Alessio DE BIASE: Dipartimento di Biologia Animale e dell’Uomo, Univer- sita di Roma “La Sapienza”, viale dell’Universitä 32, I- 00185, Rome, Italy. Received 24.06.2006 Accepted 13.03.2007 Corresponding editor: M. Schmitt 158 Buchbesprechung BRAUN, Monika & DIETERLEN, Fritz (Hrsg.): Die Säugetiere Ba- den-Wiirttembergs. (2003): Band 1: Allgemeiner Teil; Fledermäuse (Chiroptera). 688 S., 320 Farbphotos, 49 SW-Photos, 186 Diagramme & Zeich- nungen, 46 Verbreitungskarten, 55 Tabellen. Preis 49,90 € (2005): Band 2: Insektenfresser (Insectivora), Hasentiere (La- gomorpha), Nagetiere (Rodentia), Raubtiere (Carnivora), Paar- hufer (Artiodactyla). 704 S., 193 Farbphotos, 2 SW-Photos, 172 Diagramme & Zeichnungen, 47 Verbreitungskarten, 138 Tabel- len. 49,90 €. Eugen Ulmer Verlag, Stuttgart. Mit dem Erscheinen des 2. Bandes so relativ kurz nach dem des 1. liegt dieses eindrucksvolle Werk mit zusammen weit mehr als 1000 Seiten nun komplett vor. Grundlage seiner Veróffentlichung ist das langjáhrige und aus 6ffentlichen Mitteln gefórderte For- schungsprojekt „Wild lebende Säugetiere in Baden-Württem- berg“. Außer von den beiden Herausgebern enthalten die bei- den Bände Textbeiträge von jeweils 20 weiteren Autoren. Ent- scheidend für das Projekt und die Fülle der darın präsentierten Daten war sicherlich auch die ehrenamtliche Mitarbeit von über 1000 weiteren Personen, die Beobachtungsdaten und Hinweise zum Vorkommen von Säugetieren in Baden-Württemberg bei- getragen haben. Den ersten Teil des 1. Bandes mit nahezu 300 Seiten bildet der Allgemeine Teil mit u.a. den folgenden Kapiteln: Säugetiere (Kennzeichen, Verbreitung und Vielfalt); Zur Geschichte der Er- forschung der Säugetierfauna (allgemein und regional ın Würt- temberg und Baden); Naturräume ın Baden-Württemberg [im folgenden: B-W]; Methoden der Datenerhebung und Datenver- arbeitung; Gewólleanalyse; Die Säugetiere im Quartär von B- W; Einwanderung, Einbürgerung und (Wieder-) Ansiedlung von Säugetieren; Jagd; Domestizierte Säugetiere in B-W; Zoonosen wildlebender Säugetiere in B-W; Gefährdungsursachen; Schutz- möglichkeiten; Rote Liste der gefährdeten Säugetiere ın B-W. Er spannt damit einen wirklich weiten thematischen Bogen. Den anschließenden Speziellen Teil von nahezu 400 Seiten Umfang leitet ein allgemeines Kapitel über Säugetiere zu ,,Klassifikati- on, taxonomische Merkmale, Maße, Fachbezeichnungen u.a.“ ein, gefolgt von einer Einführung in die Ordnung Chiroptera, ei- nem Überblick über die heimischen Fledermäuse und einem Be- stimmungsschlüssel der mitteleuropäischen Arten nach äußeren Kennzeichen und Zahnmerkmalen. Ausführliche monographi- sche Abhandlungen zu 22 ın B-W vorkommenden Fledermaus- arten schließen sich an. Die Artkapitel sind jeweils nach einem weitgehend einheitlichen Schema gegliedert und behandeln fol- gende Themen: Namensgebung (Taxonomie und Geschichte des deutschen Trivialnamens); Beschreibung (äußere Merkmale, Ma- Be und Gewicht, Schädel und Gebiss [bei einander ähnlichen Ar- ten wird ausführlich auf Unterscheidungsmerkmale eingegan- gen]); Verbreitung (Artareal insgesamt, in Deutschland und spe- ziell in B-W); Lebensraum; Lebensweise (Sommerquartiere, Nahrungsökologie, Echoortung, Fortpflanzung, Wanderungen, Winterquartiere und Winterschlaf, Populationsdynamik); Arten- schutz. Am Beginn des 2. Bandes wird das Projekt, das Grund- lage der vorliegenden Veröffentlichung war, noch einmal kurz vorgestellt, gefolgt von einer erneuten kleinen Einführung ın Da- tenerhebung und Datendarstellung sowie zu den für Säugetie- ren spezifischen Maßangaben. Daran schließt sich auf mehr als 600 Seiten die Darstellung der übrigen in B-W vorkommenden Säugetierarten aus den Ordnungen der Insectivora (11 Arten) (der aktuelle Erkenntnisstand, wonach die Zusammenfassung der Igel mit Spitzmäusen und Maulwürfen in einer gemeinsamen Ord- nung keine natürliche Verwandtschaftsgruppe darstellt, wurde hier noch nicht berücksichtigt), Lagomorpha (3 Arten), Roden- tia (21 Arten), Carnıvora (14 Arten) und Artiodactyla (7 Arten) an. Auch in diesem Band gibt es zu allen Ordnungen jeweils kür- zere allgemeine Einführungskapitel (bei den Artiodactyla auch zu den Unterordnungen der Nichtwiederkáuer und der Wieder- käuer), ebenso zu den jeweiligen Familien (bei den Rodentia auch zu Unterfamilien). Der Aufbau der einzelnen Artkapitel folgt einem ähnlichen Schema wie in Band 1, es fehlen aber durchgehend die Ausführungen zur Namensgeschichte. Beson- ders eingehend wird die Lebensweise der einzelnen Arten mit Ausführungen zu Verhalten, Aktivität, Fortbewegung, Aktions- raum, Sinnen, Kommunikation, Lautäußerungen, Ernährung, Fortpflanzung, Jugendentwicklung, Populationsdynamik, natür- lichen Feinden, Parasiten und Krankheiten sowie — wo relevant — auch zur Bejagung und deren Auswirkungen dargestellt. Alle Beiträge sind in ihrer Darstellung allgemeiner Zusammen- hänge und der zu den einzelnen Arten präsentierten Fakten fach- lich fundiert und bereiten für den betroffenen geographischen Raum eine große Fülle interessanter Einzeldaten auf; diese wie- derum werden dann in sinnvoller Weise in einen größeren Zu- sammenhang gestellt. Auch in ihrer Ausstattung, ihrem Druck und der Zahl und Qualität der Abbildungen werden diese bei- den Bände hohen Ansprüchen gerecht. Bei einem so umfang- reichen Werk, an dem viele Autoren mitgewirkt haben, ıst es fast unvermeidlich, dass sich hier und da Uneinheitlichkeiten der Darstellung oder auch kleinere redaktionelle Fehler einschlei- chen. Um nur ein Beispiel zu nennen: In Band 2 sind ın Tabel- le 60 (S. 357) für die Felidae rezent 11 Gattungen und 36 Arten aufgeführt, der anschließende Text auf S. 358 nennt aber 18 Gat- tungen und 35 Arten. Als wenig nutzerfreundlich empfindet es der Rezensent, dass das vollständige Literaturverzeichnis am En- de des 2. Bandes nach den einzelnen darin behandelten Säuger- ordnungen unterteilt ist. So sucht man erst einmal nach der Sei- te, auf der die Literaturzitate für die jeweiligen Ordnungen be- ginnen und kann erst dann gezielt alphabetisch nach dem spe- ziellen Zitat suchen, an dem man interessiert ist. Im 1. Band ist dies besser gelöst; dort folgen dem Allgemeinen Teil und dem Speziellen Teil die jeweiligen Literaturverzeichnisse. Von solchen wirklich geringfügigen Kritikpunkten abgesehen ge- bührt den Förderern des Projekts, aus dem diese Veröffentlichung hervorgegangen ist, Dank und allen anderen daran beteiligten Personen, besonders natürlich den beiden Herausgebern, aus- drückliche Anerkennung für diese Leistung. Diese beiden Bän- de gehören ohne Einschränkung ın jede größere öffentliche Bi- bliothek, die von ernsthaft an unseren heimischen Säugetieren interessierten Personen genutzt wird. Wer beruflich mit diesen Tieren arbeitet, sollte sich dieses Werk auf jeden Fall anschaf- fen. Wenn man einmal vom viel umfangreicheren „Handbuch der Säugetiere Europas“ absieht (das primär für einen anderen Nutzerkreis gedacht ist), gibt es aktuell auf dem deutschsprachi- gen Buchmarkt keine auch nur annährend in Qualität und Infor- mationsfülle vergleichbare Veröffentlichung zur Biologie (im weitesten Sinne) der heimischen Säuger — und es besteht wohl auch wenig Aussicht, dass auf absehbare Zeit eine solche erschei- nen wird. Angesichts der bedauernswerten Entwicklung, dass die organismische Biologie im Unterricht an weiterführenden Schulen und im Grundstudium der Biologie immer weiter zu- rückgedrängt wird, kann man nur hoffen, dass diese beiden Bän- de in diesen Schulen und in den betreffenden Universitätsinsti- tuten angeschafft werden und eine möglichst intensive Nutzung erfahren. Gustav PETERS, Zoologisches Forschungsmuseum Alexander Koenig, Bonn Bonner zoologische Beiträge Band 55 (2006) Heft 2 | Seiten 159-162 Bonn, Juli 2007 Morphological differentiation of mainland Citril Finches, Carduelis [citrinella] citrinella and insular Corsican (Citril) Finches, Carduelis |citrinella| corsicanus Marc Imanuel FÖRSCHLER| 0) . 2 & Karl Heinz SIEBENROCK2 ! Institut für Vogelforschung, “Vogelwarte Helgoland”, Germany 2 Max Planck Institut für Ornithologie, Vogelwarte Radolfzell, Germany The Citril Finch 1s one of the few endemic bird species restricted to European mountains (Voous 1960; NEWTON 2003). Two allopatric forms may be distinguished. Insu- lar Corsican (Citril) Finches, Carduelis [citrinella] cor- sicanus, live exclusively on Corsica, Sardinia and some Tuscany islands (Capraia, Elba, Gorgona) (ARCAMONE 1993; BACCETTI & MARKI 1997; CRAMP & PERRINS 1994; THIBAULT 1983; THIBAULT & BONACCORSI 1999; MOLTO- NI 1975), whereas mainland Citril Finches, Carduelis [ci- trinella] citrinella, occur at higher elevations in the moun- tain ranges of central and south-western Europe (Alps, Black Forest, Vosges, Jura, Massif Central, Cevennes, Py- renees, Cantabrians and Sierras of central Spain), gene- rally above 800 m asl. (BACCETTI & MARKI 1997; CRAMP & PERRINS 1994; GLUTZ VON BLOTZHEIM & BAUER 1997; HOLZINGER 1997). There is an ongoing debate about the taxonomic status of Citril Finches and Corsican Finches. In spite of recently detected genetic differences (mitochondrial DNA differs by 2.7 %), the authors of these studies still regard the two forms as conspecific (PASQUET & THIBAULT 1997; THIBAULT & BONACCORSI 1999). In contrast, SANGSTER (2000) assigned species status to the Corsican Finch, re- ferring to the same genetic results. Based on his recom- mendations in combination with data on variations in plumage colouration (CRAMP & PERRINS 1994; PASQUET 1994) and vocalization (CHAPPUIS 1976; CRAMP & PER- RINS 1994; recently confirmed by FORSCHLER & KALKO 2007), the Association of the European Rarities Commit- tees (AERC) now treats the two forms as full species with their own evolutionary histories (SANGSTER et al. 2002). Furthermore the two species have been recently reclassi- fied within the genus Carduelis instead of being a mem- ber of Serinus (ARNAIZ-VILLENA et al. 1998, 1999). Both forms are therefore members of the superspecies Cardu- elis [citrinella] (BARTHEL & HELBIG 2005; Helbig 2005). Only little data has been published on evidence for dif- ferentiation in morphological character traits between the two forms. According to the available notes on morphom- etry, both species do not show large geographical differ- ences in body size (ALONSO & ARIZAGA 2006; BORRAS et al. 1998; BRANDL & BEZZEL 1989; CRAMP & PERRINS 1994; MARKI & BIBER 1975), with the exception that Cor- sican Finches seem to have generally slightly shorter wings than nominate Citril Finches (CRAMP & PERRINS 1994). Nominate birds in the Eastern Pyrenees appear to have longer wings than do nominate birds in the Alps, but evidence is weak and may be also due to variation between different measurers (BRANDL & BEZZEL 1989; BORRAS et al. 1998; MARKI & BIBER 1975). Adult male Citril Finch- es have an average population weight of 12.5 g in the East- ern Pyrenees (BorRAS et al. 1998), while adult male Cor- sican Finches are little bit lighter with 11.5 g in average (NICOLA BACCETTI, GILLES FAGGIO, SERGIO NISSARDI & PHILIPPE PERRET, pers. comm. ). We analysed 20 morphological character traits following the methods of LEISLER & WINKLER (1985, 1991, 2001), LEISLER et al. (1997) and KORNER-NIEVERGELT & LEISLER (2004). The following external features were measured. (1) wing length (the distance between carpal joint and tip of falltened wing); (2) tail length (from insertion of cen- tral pair of feathers to tip of longest rectrix); (3) fork of the tail (difference from longest to shortest inner rectrix); (4) wing width (the distance between carpal joint and the tip of the longest secondary); (5) primary projection (the distance between tip of 1% secondary and wing tip); (6) distal (the distance between tip of 10'h primary and wing tip); (7) alula (the distance between carpal joint and tip of the longest alula); (8) notch P21 (length of notch on in- ner web of second primary); (9) notch P3o (length of notch on outer web of third primary); (10) Kipp Index (distance of 1% secondary to wing tip x 100/ wing length see Kipp 160 Marc Imanuel FÓRSCHLER & Karl Heinz SIEBENROCK: Morphological differentiation of (Citril) Finches 1959) (11) bill length (from tip to skull); (12) bill width (at the rear end of nostrils); (13) bill height (maximum depth); (14) rictal bristles (the length of the vibrisse at the bill base) (15) tarsus length (from notch of the intertarsal joint to the lower edge of the last complete scale); (16) tarsus diameter | (lateral maximal diameter of tarsus): (17) tarsus diameter 2 (sagittal maximal diameter of the tar- sus; (18) hind toe (length of digit 1); (19) middle toe (length of digit 3); (20) hind claw (the cord from the tip to dege of the skin of nail); (21) middle claw (the chord of nail 3); (22) index of wing length/ tarsus length. In total we measured 23 individuals of mainland Citril Finches from the Northern Alps and 13 individuals of in- sular Corsican Finches from Corsica and Sardinia. For our study we used only adult males to guarantee comparabil- ity. All measurements were conducted by KARL HEINZ SIEBENROCK. Museum specimens from the following mu- seum collections were used exclusively: Zoologische Staatssammlung München (Munich), Zoologisches Mu- seum der Humboldt Universitat (Berlin) and Naturkunde- musem Rosenstein (Stuttgart). We found significant morphological differentiation be- tween mainland Citril Finches and insular Corsican Finch- es (see Table 1). Corsican Finches are generally smaller. Wings are considerably longer and more pointed (Kipp In- dex) in Citril Finches. Accordingly, significant variations may be also found between other wing measurements (sec- ondary, primary projection, distal, alula, notch P21, notch P30). Corsican Finches have a shorter tail and their tail is less forked. Furthermore Citril Finches possess significant- ly broader, higher and longer bills than Corsican Finch- es. No significant difference was found for the length of the rictal bristles. Citril Finches have longer and stonger tarsi, a longer middle toe and longer claws. However the length of the hind toe (maybe due to measurement errors) and the ratio of wing length to tarsus length are not sig- nificantly different. Climate is known to play an important role in geograph- ic size variation (JAMES 1970; JOHNSTON & SELEANDER 1973). Corsican Finches appear to inhabit generally areas of warmer clime than do Citril Finches (THIBAULT & Bonaccors! 1999) and may therefore have developed Table 1. Mean values of 20 morphological traits and 2 indices of Citril and Corsican Finches. Comparisons were conducted with t-tests (parametric distributed data sets) and Mann Whitney U tests (non-parametric distributed data sets). Citril Finch Corsican Finch Statistical comparison Sign-Level Flight apparatus (n=23) in mm (n=13) in mm Wing length 76.4+1.5 SD 72.5+2.2 SD t-test; t=6.569; df=34; p<0.001 ane Tail length 53.3+1.8 SD 50.4+2.2 SD t-test; t=4.336; df=34; ps0.001 en Fork of the tail -10.0+1.1 SD -7.2+1.1 SD MWU; T=94; ps0.001 tag Wing width 53.521,6 SD 51.0+1.6 SD MWU; T=126.0; p<0.001 ity Primary projection 22.6+1.2 SD 20,121.5 SD t-test; t=5.319; df=34; p<0.001 ae Distal 54.1+1.4 SD 50.3+2.1 SD MWU; T=118.5; ps0.001 es Alula 56.4+1.5 SD 52.8+2.3 SD MWU; T=119; ps0.001 nr Notch P3o 23.1+1.4 SD 21.0+1.2 SD t-test; 154.692; df=33; p<0.001 E Notch P21 19.8+0.8 SD 17./21.1 SD MWU; T=94; p<0.001 Eb Kipp index 29.6+1.3 SD 27821.9 5D t-test; t=3.439; df=34; p=0.002 an Feeding apparatus Bill length 12.2+0.6 SD 11.6+0.5 SD MWU; T=133.0; ps0.001 visi Bill width 5.3£0.2 SD 4.8+0.3 SD MWU; T=115.5; ps0.001 er Bill height 6.2+0.3 SD 5.6+0.3 SD t-test; t=6.432; df=34; p<0.001 EXE Rictal bristles 2.6+0.7 SD 2.4+0.7 SD t-test; t=4.950; df=34; p=0.420 ns Hind limb Tarsus length 14.4+0.6 SD 13.5+0.5 SD t-test; t=4.653; df=34; ps0.001 BA Tarsus diameter | 0.88+0.08 SD 0.82+0.06 SD t-test; t=2.278; df=34; p=0.034 E Tarsus diameter 2 1.34+0.13 SD 1.15+0.06 SD t-test; 154.950; df=34; ps0.001 hae Hind toe 6.5+0.4 SD 6.4+0.3 SD t-test; t=0.599; df=34; p=0.553 ns Middle toe 10.9+0.6 SD 10.2+0.6 SD t-test; t=3.240; df=34; p=0.003 at Hind claw 6.1+0.5 SD 5.6+0.5 SD t-test; t=2.926; df=34; p=0.008 + Middle claw 5.4+0.4 SD 4.7+0.4 SD t-test; t=5.534; df=34; p<0.001 cds Wing length/ tarsus index 5.3+0.2 SD 5.4+0.2 SD MWU; T=267; p=0.392 ns Bonner zoologische Beiträge 55 (2006) 161 smaller body size and body weight. However, this sup- posed climatic difference is essentially not the case for a majority of Corsican Finches, living on the highest moun- tain slopes of Corsica, where climatic conditions are quite comparable to the Alps or Pyrenees. Furthermore, meas- urements conducted by BORRAS et al. (1998) revealed no clear clinal variation in size from north to south in main- land Citril Finches as expected. They propose that other factors such as habitat or diet may be of larger importance. We suggest therefore that the morphological differentia- tion observed in our study may have evolved as a conse- quence of different habitat conditions, since Citril and Cor- sican Finches differ in several features of habitat choice (FORSCHLER & KALKO 2006b). While Citril Finches pre- fer semi-open conifer forests, especially pine woodland in higher mountains, Corsican Finches generally inhabit more open habitats dominated by Tree Heath, Erica ar- borea. It is highly probable that Citril Finches are adapt- ed to forage over longer distances (longer and more point- ed wings, longer tail) and probably also to perch on stronger branches and twigs of pine trees (stronger feet), whereas Corsican Finches appear to show rather adapta- tions to the Macchia thickets with Tree Heath or other scrub vegetation of Mediterranean islands (FORSCHLER & Kalko 2006b). As a consequence Corsican Finches may have developed smaller and less pointed wings and short- er tails due to the reduction of long distance migration (THIBAULT & BONACCORSI 1999), favouring more the ma- noeuvrability within the Mediterranean scrub vegetation. In this context, the difference in bill size might be ex- plained by variations in feeding ecology of the two species (FORSCHLER & KALKO 2006a) with Citril Finches proba- bly more adapted to larger seeds (mainly those from pines) and Corsican Finches more adapted to smaller herb seeds (e.g. Shepherd's Purse). Further analyses may reveal how the morphological differentiation in the locally isolated populations of these two allopatric forms has evolved in relation to distinct ecological conditions and adaptation processes. Acknowledgements. We thank Bernd Leisler, Renate van den Elzen, Michael Schmitt and two anonymous referees for their constructive comments. We thank the staff of the Zoologische Staatssammlung Miinchen (Munich), Zoologische Museum of the Humboldt University (Berlin) and the Naturkundemusem Rosenstein (Stuttgart) for the support of this study. REFERENCES ALONSO D. & ARIZAGA J. 2006. Biometrics of Citril Finch Ser- inus citrinella in the west Pyrenees and the influence of feath- er abrasion on biometric data. Ringing & Migration 23. 116-124. ARCAMONE, E. 1993. Venturone. P. 286 in: MESCHINI, E. & FRU- ais S. (eds., Atlante degli uccelli nidificanti in Italia. Institu- to per la fauna selvatica. ARNAIZ-VILLENA, A., ALVAREZ-TEJADO, M., RUIZ-DEL-VALLE, V., GARCIA-DE-LA-TORRE, C., VARELA, P., RECIO, MJ., FERRE, S. & MARTINEZ-LASO, J. 1998. Phylogeny and rapid Northern and Southern Hemisphere speciation of goldfinches during the Miocene and Pliocene Epochs. Cellular Molecular Life Scien- ce 54: 1031-1041. ARNAIZ-VILLENA, A., ALVAREZ- TEJADO, M., RUIZ-DEL-VALLE, V., GARCIA-DE-LA-TORRE, C., VARELA, P., RECIO, MJ., FERRE, S. & MARTINEZ-LASO, J. 1999. Rapid radiation of Canaries. Mol- ecular Biology Evolution 16: 2-11. BACCETTI, N. & MARKI, H. 1997. Citril Finch. P. 711 in: HAGE- MENER, W. J. M. & BLaAıR, M. J. (eds.) The EBCC Atlas of European Breeding Birds. Poyser. London. BARTHEL, P. H. & HELBIG, A. J. 2005. Artenliste der Vögel Deutschlands. Limicola 19: 89-111. BRANDL, R. & BEZZEL, E. 1989. Morphometrische Alters- und Geschlechtsunterschiede beim Zitronenzeisig Serinus citri- nella. Ornithologischer Beobachter 86:137-143. CHAPPUIS, C. 1976. Origine et Evolution des vocalisations de cer- tains oiseaux de Corse et des Baléares. Alauda 44: 475-495. CRAMP, S. & PERRINS, C. M. 1994. The birds of the western Pa- learctic. Vol. 8. Oxford: University Press. FORSCHLER, M. I. & KALKo, E. K. V. 2006a. Macrogeographic variations in food choice of mainland citril finches Carduelis [citrinella] citrinella versus insular corsican (citril) finches Carduelis [citrinella] corsicanus. Journal of Ornithology 147: 441-447. FORSCHLER, M. I. & KALKO, E. K. V. 2006b Breeding ecology of mainland Citril Finches Carduelis citrinella and insular Cor- sican Finches Carduelis [citrinella] corsicanus. Journal of Or- nithology 147: 553-564. FÓRSCHLER, M. I. & KaLKo, E. K. V. 2007. Geographical diffe- rentiation, acoustic adaptation and species boundaries in main- land citril finches and insular Corsican finches, superspecies Carduelis [citrinella]. Journal of Biogeography 34: published online under DOI 10.1111/j.1365—2699.2007.01722.x GLUTZ VON BLOTZHEIM, U. N. & BAUER, K. M. 1997. Handbuch der Vögel Mitteleuropas. 14. Aula-Verlag, Wiesbaden. HELBIG, A. J. 2005. Anmerkungen zur Systematik und Taxono- mie der Artenliste der Vögel Deutschlands. Limicola 19: 112— 128. HÖLZINGER, J. 1997. Die Vögel Baden-Württembergs. Band 3.2. Verlag Eugen Ulmer, Stuttgart. KORNER-NIEVERGELT, F. & LEISLER, B. 2004. Morphological con- vergence in conifer-dwelling passerines. Journal of Ornithol- ogy 145: 245-255. JAMES, F. C. 1970. Geographic size variation in birds and its re- lationship to climate. Ecology 51: 365-390. JOHNSTON, R. F. & SELEANDER R. K. 1973. Evolution in the hou- se sparrow. III. Variation in size and sexual dimorúhism in Europe and North and South America. American Naturalist 107: 373-390. Kipp, A. 1959. Der Handflügel-Index als flubiologisches Maß. Die Vogelwarte 20:77-86. LEISLER, B. & WINKLER, H. 1985. Ecomorphology. Current Or- nithology 132: 155-186. LEISLER, B. 8 WINKLER, H. 1991. Ergebnisse und Konzepte öko- morphologischer Untersuchungen an Vögeln. Journal of Or- nithology 132: 373-425. LEISLER, B. & WINKLER, H. 2001. Morphological convergence in papayrus dwelling passerines. Ostrich 15: 24-29. LEISLER, B., HEIDRICH, P., SCHULZE-HAGEN, K. & WINK, M (1997. Taxonomy and phylogeny of reed warblers (genus Acrocephalus) based on mtDNA sequences and morphology. Journal of Ornithology 138: 469-496. 162 Marc Imanuel FÓRSCHLER & Karl Heinz SIEBENROCK: Morphological differentiation of (Citril) Finches MARKI, H. & BIBER, O. 1975. Flügellänge und Gewichtsschwan- THIBAULT, J.-C. & BONACCORSI, G. 1999. The Birds of Corsica. kungen beim Zitronenzeisig Serinus c. citrinella vor und wäh- BOU Checklist 17. British Ornithologists’ Union. The Natur- rend des Herbstzuges. Jahresberichte Naturhistorisches Mu- al History Museum, Tring, Herts, UK. seum Bern 5: 153-164. Voous, K. H. 1960. Atlas of European birds. Nelson, London MOLTONI, E. 1975. L’avifauna dell'isola di Capraia (Arcipela- and Amsterdam. go toscano). Rivista Italiana Ornithologia 45: 97-217. WHITEHEAD, J. 1885. Ornithological notes from Corsica. Ibis 27: NEWTON, 1. 2003. The speciation and biogeography of birds. Aca- 39. demic Press, London. PASQUET, E. 1994. Venturon montagnard. In: YEATMAN-BERTHO- Authors’ addresses: Marc Imanuel FÖRSCHLER (corre- LET, D. (edt.) Nouvel atlas des oiseaux nicheurs de France sponding author), Institut für Vogelforschung, “Vogelwarte 1985-1989. Société ornithologique de France, Paris. Helgoland”, An der Vogelwarte 21, D-26386 Wil- PASQUET, E. AND THIBAULT, J. C. 1997. Genetical differences : © among mainland and insular forms in the Citril Finch Seri- helmshaven, Germany; E-Mail: marc.foerschler@ifv.ter- nus citrinella. Ibis 139: 679-684. ramare.de; Karl Heinz SIEBENROCK Max Planck Institut fúr SANGSTER, G. 2000. Genetic distance as a test of species bound- Ornithologie, Dept. Vogelwarte Radolfzell, D-78315 aries in the Citril Finch Serinus citrinella: a critique and tax- Radolfzell, Germany onomic reinterpretation. Ibis 142: 487-490. des G., Knox, A. E .. ÉS J.& nn u nn. Received: 18.11.2005 oe recommendations for uropean Birds. 1S 2 Revised: 22.02.2006 THIBAULT, J.-C. 1983. Les oiseaux de la Corse. Ajaccio, Parc Na- Accepted 02.03.2006 turel Regional de la Corse. Corresponding editor: R. van den Elzen Bonner zoologische Beiträge Band 55 (2006) Heft 2 Seiten 163-178 Bonn, Juli 2007 African Coleoptera Type Specimens collected by Thomas Wagner in the collection of the ZFMK Karl-Heinz LAMPE, Dirk ROHWEDDER & Carola SCHMIDT Zoologisches Forschungsmuseum Alexander Koenig, Bonn, Germany Abstract. The type specimens of African Coleoptera from material collected by Thomas Wagner mainly using insecticidal ca- nopy fogging technique and housed in the Zoologisches Forschungsmuseum Alexander Koenig (ZFMK), Bonn, are listed. The- re are 71 names recorded; of these 37 (52%) are represented by holotypes. Specific names are sorted by their respective family and both are listed alphabetically. Reference to the original description of each taxon is provided and sex and number of each type given as well as images of the name bearing primary types. Keywords. Type specimens, canopy fogging, Coleoptera, Africa. 1. INTRODUCTION Thomas Wagner’s material at the Zoologisches Forschungsmuseum (ZFMK) house African Coleoptera mainly collected in Rwanda, Kenya and Uganda from 1993 until today by insecticidal tree fogging. Voucher specimens were computerized within phase II of the Ger- man BIOTA East Africa Project (subproject BIOTA E15: “Capacity building in biodiversity information systems for insects and plants in East Africa”) which is financially sup- ported by the German Ministry of Science and Education (BMBF) and runs until May 2007. Data capture was done with the specimen based database BIODAT. By adapta- tion to the ABCD schema (Access to Biological Collec- tion Data) BIODAT became both a BIOCASE- (Biologi- cal Collection Access Service for Europe) and GBIF- (Global Biodiversity Information Infrastructure) provider. Therefore full information at the specimen level, and not only of type specimens, for selected ZFMK insect collec- tions will be commonly available on the World Wide Web (http://www. biodat.de). In case of type specimens images of the type itself and of the original labels are provided. The present catalogue covers 370 type specimens of 71 species of which 37 are represented by holotypes. In this publication the specific taxa of Coleoptera are grouped by families and arranged alphabetically. They are listed by species epitheta, author, year of publication, page number where species was described, original generic as- signment in brackets followed by the catalogue number/s, number/s sex and kind of type specimen. All primary types are listed with the original label information, supplement- ed by date and place collected information. Normally ‘named area’ information is used. If available the exact ‘gathering site’ information is indicated by geographical coordinates (Latitude/Longitude). Higher ranked geo- graphic information such as region, country and continent is added in a hierarchy. To avoid hypertrophic text length most of the secondary types are listed without original la- bel information. If the collection, however, harbours on- ly secondary types of a given taxon the original label in- formation is given for the first type specimen. Images of the primary types are provided. The following abbreviations were used in the lists: HT — holotype, PT — paratype, AT — “allotype” COLEOPTERA ANTHRIBIDAE cognatus Frieser, 1997: 12 (Mecocerus) COL 2006/188 19 PT Rwanda, Nyakabuye - Cyamudongo X.93 - Th. Wagner leg. /C.g.7 1 / Paratype / Mecocerus cognatus sp. n. - Paratypus Q - det. R. Frieser 1996 October 1993 Cyamudongo, Nyakabuye, Cyangugu, Rwanda, Africa 2°33°S, 28°59°E (gathering site: estimated value) marginalis Frieser, 1997: 19 (Triplodus) COL 2006/185 CHT (Fig. 1) Rwanda, 1700 m - Cyamudongo, X.93 - Th. Wagner leg. / C.g.4 3 / Holotype / Triplodus marginalis sp. n. — Holotypus 0° - det. R. Frieser 1996 October 1993 Cyamudongo, Nyakabuye, Cyangugu, Rwanda, Africa 2°33°S, 2859 E (gathering site: estimated value) 164 Karl-Heinz LAMPE, Dirk ROHWEDDER & Carola SCHMIDT: African Coleoptera Types collected by Th. Wagner mm 3 Al Figs 1-9. Triplodus marginalis Frieser, 1997 (1) & HT, Uncifer nubilus Frieser, 1997 (2) @ HT, Uncifer proprius Frieser, 1997 (3) 7 HT, Monolepta alwineae Wagner, 2002 (4) Y HT, Monolepta budongoensis Wagner, 2002 (5) Y HT, Monolepta comoeen- sis Wagner, 2000 (6) & HT, Monolepta congoensis Wagner, 2002 (7) Y HT; Monolepta constantini Wagner, 2002 (8) 9 HT, Bo- noesioides jacobyi Freund & Wagner, 2003 (9) & HT: Habitus in dorsal view. 8 7 A Bonner zoologische Beitráge 55 (2006) 165 COL 2006/187 October 1993 Cyamudongo, Nyakabuye, Cyangugu, Rwanda, Africa 2°33°S, 28°59 E (gathering site: estimated value) COL 2006/186 10 PT October 1993 Karengera, Cyangugu, Rwanda, Africa 2°31'S, 29°02°E (gathering site: estimated value) 19 AT muehlei Frieser, 1997: 15 (Uncifer) COL 2006/175 109 PT Uganda, District Masindi - Budongo Forest n. Sonso - 1°45°N, 31°35°W 11.-20-VII.95 - Th. Wagner leg. / T.n.15 1 / Paratype / Uncifer muehlei sp. n. - Paratypus 0’ - det. R. Frieser 1996 11.-20. July 1995 Sonso River (Budongo Forest), Masindi, Uganda, Africa 1°45°N, 31°35°E (exact gathering site information; error on original label!) nubilus Frieser, 1997: 15-16 (Uncifer) COL 2006/176 O HT (Fig. 2) Uganda, District Masindi - Budongo Forest n: Sonso - 1°45°N, 31°35°W 1.-10.VILOS - Th. Wagner leg. / Holotype / Uncifer nubilus sp. n. - Holotypus O' - det. R. Frieser 1996 01.-10. July 1995 Sonso River (Budongo Forest), Masindi, Uganda, Africa 1°45°N, 31935'E (exact gathering site information; error on original label!) COL 2006/177 19 AT 21.-30. July 1995 COL 2006/178 IS PT 11.-20. July 1995 Sonso River (Budongo Forest), Masindi, Uganda, Africa 1°45°N, 31°35°E (exact gathering site information; error on original label!) proprius Frieser, 1997: 16 (Uncifer) COL 2006/179 Sd HT (Fig. 3) Uganda, District Masindi - Budongo Forest n. Sonso - 1°45°N, 31935" W 21.-30.V11.95 - Th. Wagner leg. / T.n.11/2 / Holotype / Uncifer proprius sp. n. - Holotypus - det. R. Frieser 1996 21.-30. July 1995 Sonso River (Budongo Forest), Masindi, Uganda, Africa 1°45°N, 31935'E (exact gathering site information; error on original label!) COL 2006/180 19 AT COL 2006/181, 184 20 PT 11.-20. July 1995 COL 2006/183 109 PT COL 2006/182 19 PT 21.-30. July 1995 Sonso River (Budongo Forest), Masindi, Uganda, Africa 1°45°N, 31°35°E (exact gathering site information; error on original label!) CERAMBYCIDAE wagneri Adlbauer, 2006: 20-21 (Stathmodera) Holotype @ - in: National Museums of Kenya (NMK), Nairobi COL 2006/688 1 PT Kenya, Kakamega F., - 0222 'N/34950'E, 1800 m, - 7-11.11.99 Th. Wagner leg. / Tn.33 1 / PARATYPUS STATHMODERA - wagneri - ADLBAUER, 2006 7.-11. Februay 1999 Kakamega Forest, Western, Kenya, Africa 0%17'N, 34953 "E (exact gathering site information; impre- cise data on original label!) COL 2006/689 1 PT January 2002 COL 2006/696 1 PT October 2002 Isiukhu River, Kakamega Forest, Western, Kenya, Africa 0°20°N, 34°53°E (exact gathering site information; impre- cise data on original label!) COL 2006/690-691 2 PT January-Februay 2002 COL 2006/692 - 695 4 PT October 2002 COL 2006/701 1 PT January 2003 Busumbuli River, Kakamega Forest, Western, Kenya, Africa 0219, 34°52°E (exact gathering site information; impre- cise data on original label!) COL 2006/697 - 698 2 PT October 2002 COL 2006/702 1 PT January 2003 Colobus Trail, Kakamega Forest, Western, Kenya, Africa 0221'N, 34°51’E (exact gathering site information; impre- cise data on original label!) COL 2006/699 1PT October 2002 Kisere Forest, Kakamega Forest, Western, Kenya, Africa 0224'N, 34754 'E (site information) COL 2006/700 1PT October 2002 Yala River, Kakamega Forest, Western, Kenya, Africa 0°12°N, 34°53°E (exact gathering site information; impre- cise data on original label!) CHRYSOMELIDAE alwineae Wagner, 2002: 35 (Monolepta) COL 2007/156 Co HT (Fig. 4) Ivory Coast, Comoe - NP, 8,75N / 3,80W - VI.-VIL.1997, C. Mody leg. / KC 35 .1C 111.15 / Holotypus / Monolepta - alwineae sp. nov. - Th. Wagner det. 02 / AfriGa - speci- men ID - 1270 - specimen data - documented - 23.111.2006 June-July 1997 Komoe, Parc National de la, Zanzan, Cote d’Ivoire, Africa 8°45°N, 3948 W (site information) 166 Karl-Heinz LAMPE, Dirk ROHWEDDER Carola SCHMIDT: African Coleoptera Types collected by Th. Wagner budongoensis Wagner, 2002: 29 (Monolepta) COL 2007/157 oO HT (Fig. 5) Uganda, District Masindi - Budongo Forest n. Sonso - 1°45°N, 31°35°E; - 21.-30.V 11.95 - Th. Wagner leg. / R.a.22 1 / Holotypus / Monolepta - budongoensis - Wag-ner / AfriGa - specimen ID - 1266 - specimen data - documented - 23.11.2006 21.-30. July 1995 Sonso River (Budongo Forest), Masindi, Uganda, Africa 1°45°N, 31°35°E (site information) COL 2006/158-159 2 PT 5.-12, Februay 1997 Semliki Forest, Bundibugyo, Western, Uganda, Africa 0°48°N, 30208 E (site information) clarae Wagner, 2000a: 36 (Monolepta) COL 2007/160 109 PT Congo Belge, P.N.G. - Miss. H. De Saeger - Ndelele/2, 6- VI-1952 - H. De Saeger. 3589 / Paratypus / Monolepta clarae - Wagner, 1999 - Th. Wagner det. 6. June 1952 Ndelele (mts.), Garamba, Parc National de la, Haut-Congo, Democratic Republic of the Congo, Africa 4%22'N, 2947'E (gathering site: estimated value) COL 2006/161 19 PT 20. June 1952 PFSK.5, Garamba, Parc National de la, Haut-Congo, Demo- cratic Republic of the Congo, Africa 4°10°N, 29°30°E (gathering site: estimated value) COL 2006/162 19 PT 22. June 1952 Utukuru (river), Garamba, Pare National de la, Haut-Congo, Democratic Republic of the Congo, Africa 4°19°N, 29°52°E (gathering site: estimated value) clasnaumanni Wagner, 2001b: 62 (Monolepta) COL 2007/163 1 PT LR.S.A.C.-Mus.Congo - Kivu : Terr. Masisi, Lacs - Moko- to, 1800 m. - B.82 N. Leleup VI-1959 / Bior. N° 81 - Hu- mus - en foret / Paratype - Monolepta - clausnaumanni June 1959 Lacs Mokoto (lake), Masisi, Nord-Kivu, Democratic Republic of the Congo, Africa 1216'S, 29°00°E (gathering site: estimated value) comoeensis Wagner, 2000b: 235 (Monolepta) COL 2007/164 CHT (Fig. 6) Ivory Coast, Comoe - NP, 8,75N / 3,80W - 1X.1999, C. Mody leg. / Holotypus / Monolepta - comoeensis - Wagner / AfriGa - specimen ID - 1296 - specimen data - document- ed - 23.111.2006 September 1999 Komoe, Parc National de la, Zanzan, Cote d'Ivoire, Africa 8°45°N, 3948 W (site information) COL 2006/165-190 September 1999 Komoe, Parc National de la, Zanzan, Cote d'Ivoire, Africa 8°45 N, 348 W (site information) 26 PT congoensis Wagner, 2002: 33 (Monolepta) COL 2007/192 CHT (Fig. 7) Uganda, Semliki Forest - 0°48°N, 30°8°E; 5.-12.11.97 - Th. Wagner leg. / C.a. S4 1 / Holotypus / Monolepta - congoensis - Wagner / AfriGa - specimen ID - 1267 - specimen data - documented - 23.111.2006 5.-12. Februay 1997 Semliki Forest, Bundibugyo, Western, Uganda, Africa 0°48°N, 30%08'E (site information) constantini Wagner, 2000a: 34 (Monolepta) COL 2007/193 CHT (Fig. 8) Uganda, Dtr. Mbale - Budadiri, 1240 m, 1°10°N, 34°20°E - 21.1X.1997 / Holotypus / Monolepta - constantini - Wagner, 2000 - Holotypus / AfriGa - specimen ID - 1289 - specimen data - documented - 23.111.2006 21. September 1997 Budadiri, Sironko, Northern, Uganda, Africa 1210'N, 34°20°E (site information) COL 2006/194 19 PT 18. June 1952 Iso, Garamba, Pare National de la, Haut-Congo, Democrat- Th. Wagner leg. ic Republic of the Congo, Africa 4°10°N, 29°30°E (gathering site: estimated value) COL 2006/195 Io PT 20. April 1952 Embe (mts.), Garamba, Pare National de la, Haut-Congo, De- mocratic Republic of the Congo, Africa 4%40'N, 29°31°E (gathering site: estimated value) decellei Wagner, 2002: 34 (Monolepta) COL 2007/225 1 PT Coll. Mus. Tervuren - Cote d'Ivoire: Mouyassue, - 30 km, E.d’Aboisso - J. Decelle VI.1961 / Récolté sur - cacaoyer / Paratypus / Monolepta - decellei Wagner June 1961 Mouyassue, Aboisso, Sud-Comoé, Cote d'Ivoire, Africa COL 2006/226 1*PT 20. April 1952 Embe (mts.), Garamba, Parc National de la, Haut-Congo, De- mocratic Republic of the Congo, Africa 4°40'N, 2931 'E (gathering site: estimated value) ephippiatoides Wagner, 2001a: 206 (Monolepta) COL 2007/227-229 3:PT SÜDAFRIKA, NE-Prov. - Tshipise, Honet Nature - Reserve Berg b. Camp - 22°37°S, 30°10°E, 320 m - NN, 23.- 25.X1.1996 - leg. M. Hartmann / Paratypus / Monolepta - ephippiatoides - Wagner 23.-25. November 1996 Bonner zoologische Beitráge 55 (2006) 167 Tshipise, Transvaal, Northern Province, South Africa, Africa 22°37°S, 30°10°E (site information) gerstaeckeri Wagner, 2001a: 200 (Monolepta) COL 2007/196-197 20' PT Musee du Congo - Kapiri - IX-1912 - Miss. Agric. / Paraty- pus / Monolepta - gerstaeckeri - Wagner September 1912 Kapiri, Katanga, Democratic Republic of the Congo, Africa 10°18°S, 26°10°E (gathering site: estimated value) COL 2006/198 1 PT Kundelungus, Katanga, Democratic Republic of the Congo, Africa 10°00°S, 28°00°E (gathering site: estimated value) hartmanni Wagner, 2000b: 233 (Monolepta) COL 2007/199 1 PT Shilouvane - “C ?” habina - Zoutpansberg - H. Junod / un- lesbare Etikett / Musee du Congo - Transvall: - Zoutpansberg - Coll. Clavareau / R. Det. - L - 3678 / Paratypus / Monolep- ta - hartmanni Wagner / coll. - Th. Wagner Zoutpansberg(= Soutpansberg), Transvaal, Northern Province, South Africa, Africa 22758 S, 29°45°E (gathering site: estimated value) COL 2006/200 10 PT 19. December 1972 Macheke, Mashonaland East, Zimbabwe, Africa 18%09'S, 31°S1°E (gathering site: estimated value) ivorensis Wagner, 2001b: 55 (Monolepta) COL 2007/201 10 PT Coll. Mus. Tervuren - Cote d'Ivoire: Heremankono, - au sud de Divo - J. Decelle VIII.1961 / Paratypus / Monolepta - ivorensis - Th. Wagner August 1961 Heremankono, Forest, Divo, Sud-Bandama, Cote d'Ivoire, Africa 549 'S, 5°00°W (gathering site: estimated value) COL 2006/202-204 3 PT June 1961 Mouyassue, Aboisso, Sud-Comoé, Cote d'Ivoire, Africa Jacobyi Freund & Wagner, 2003: 1953 (Bonesioides) COL 2007/191 O HT (Fig. 9) Uganda, District Masindi - Budongo Forest n. Sonso - 1°45°N, 31°35°E; - 19.-30.V1.95 - Th. Wagner leg. / Holo- typus - Bonesioides - jacobyi / Bonesioides - jacobyi - Fre- und & Wagner 2000 / AfriGa - specimen ID - 1642 - spec- imen data - documented - 23.11.2006 19.-30. June 1995 Sonso River (Budongo Forest), Masindi, Uganda, Africa 1°45°N, 31°35°E (site information) kakamegaensis Middelhauve & Wagner, 2001: 526 (Afrocrania) Holotype O° - in: National Museums of Kenya (NMK), Nairobi COL 2007/231 -234 5PT 0°22°N/34°50°E, 1600m, - 7- 11.11.99, Th. Wagner leg. / Paratypus - Afrocrania - Kenya, Kakamega F., - kakamegaensis - Middelhauve & Wagner 7.-11. Februay 1999 Kakamega Forest, Western, Kenya, Africa O° ITN, 34°53°E (exact gathering site information; impre- cise data on original label!) laboissierei Wagner, 2001a: 201 (Monolepta) COL 2007/205-207 3 PT Coll. Mus. Congo - Lulua: Kapanga - X.1933 - G.F. Over- laet / Paratypus / Monolepta - laboissierei Wagner / coll. — Th. Wagner October 1933 Kapanga, Lulua, Kasai Occidental, Democratic Republic of the Congo, Africa 8°13°S, 22722 E (gathering site: estimated value) COL 2006/208-209 2 PT 20. October 1951 COL 2006/210 1 PT 12. April 1952 COL 2006/211 1 PT 18. July 1952 Garamba, Parc National de la, Haut-Congo, Democratic Re- public of the Congo, Africa 4°10 N, 29°30°E (gathering site: estimated value) marcoi Freund & Wagner, 2003: 1957 (Bonesioides) COL 2007/230 ı PT Uganda, Semliki Forest - 0°48°N, 30°8°E; 5.-12.11.97 - Th. Wagner leg. / C.a. S7 1 / Paratypus / Bonesioides - marcoi - Freund & Wagner 5.-12. Februay 1997 Semliki Forest, Bundibugyo, Western, Uganda, Africa 0%48'"N, 30%08 E (site information) mpangae Wagner, 2000a: 38 (Monolepta) COL 2007/212-213 20' PT Uganda Prot. - Mpanga Forest, - Toro. 4,800 ft. - 13-23 Nov. 1911. - S. A. Neave. / 1912-193 / Paratypus / Monolepta mpangae - Wagner - Th. Wagner det. 1999 13.-23. November 1911 Mpanga Forest (forest), Central, Uganda, Africa 0°12 N, 32918 E (gathering site: estimated value) naumanni Wagner, 2005: 279 (Monolepta) COL 2007/214 Co HT (Fig. 10) Uganda, District Masindi - Budongo Forest n. Sonso - 1°45°N, 31°35°E; 1.-10.VIL9OS - Th. Wagner leg. / T.r.15 2 / Holotypus / Monolepta - naumanni sp. nov. - Th. Wagner / AfrıGa - specimen ID - 1835 - specimen data - documented - 23.111.2006 168 Karl-Heinz LAMPE, Dirk ROHWEDDER & Carola Schmipr: African Coleoptera Types collected by Th. Wagner Figs 10-18. Monolepta naumanni Wagner, 2005 (10) 9 HT, Monolepta peleae Wagner, 2003 (11) 9 HT, Monolepta sonsoensis Wagner, 2001 (12) & HT, Nephus brevipilosus Fürsch, 1997 (13) @ HT, Pseudoscymnus brunneus Fúrsch, 1997 (14) HT, Epi- lachna carapacola Fürsch, 1997 (15) 0 HT, Epilachna conspergata Fürsch, 1997 (16) 9 HT, Harmonia gilvella Fürsch, 2001 (17) a HT, Scotoscvmnus glabripilosus Fürsch, 1997 (18) 9 HT: Habitus in dorsal view. Bonner zoologische Beiträge 55 (2006) 169 1.-10. July 1995 Sonso River (Budongo Forest), Masindi, Uganda, Africa 1°45°N, 31°35°E (site information) COL 2007/244-245 2 PT 1.-10. July 1995 COL 2007/246-247 2 PT 11.-20. July 1995 Sonso River (Budongo Forest), Masindi, Uganda, Africa 1°45°N, 31°35°E (site information) COL 2007/248 1 PT 5.-12. Februay 1997 Semliki Forest, Bundibugyo, Western, Uganda, Africa 0°48°N, 30°08 E (site information) COL 2006/251 1 PT January 2003 Busumbuli River, Kakamega Forest, Western, Kenya, Africa 0°19'N, 34°52°E imprecise data on original label!) COL 2006/253 1 PT October 2002 Colobus Trail, Kakamega Forest, Western, Kenya, Africa 0°21'N, 3451 E imprecise data on original label!) COL 2006/252 1PT January 2003 (exact gathering site information; (exact gathering site information; Kisere Forest, Kakamega Forest, Western, Kenya, Africa 0°24 N, 34954 E (site information COL 2007/249 1 PT November 2001 COL 2007/250 1 PT January 2003 Yala River, Kakamega Forest, Western, Kenya, Africa 0912N, 34°53°E imprecise data on original label!) (exact gathering site information; peleae Wagner, 2003: 80 (Monolepta) COL 2007/215 CHT (Fig. 11) Rwanda, Rusumo - Ibanda Makera, X.93 - Th. Wagner leg. / T.n.1 1 / Holotypus / Monolepta - peleae - sp. nov. Th. Wagner / AfriGa - specimen ID - 1708 - specimen data - documented - 23.111.2006 October 1993 Ibanda Makera, Rusumo, Kibungo, Rwanda, Africa 2°15° S, 3050'E (gathering site: estimated value) COL 2006/216 1PT 1893 Pretoria, Transvaal, Northern Province, South Africa, Africa 25°45°S, 28°10 E (gathering site: estimated value) COL 2006/217 ı PT 30. July — 2. August 2000 Peramiho, Ruvuma, Tanzanıa, Afrıca 10°39°S, 35227 E (gathering site: estimated value) sonsoensis Wagner, 2001b: 54 (Monolepta) COL 2007/218 CHT (Fig. 12) Uganda, District Masindi - Budongo Forest n. Sonso - 1°45°N, 31°35°E; 1.-10.V11.95 - Th. Wagner leg. / Holoty- pus / Monolepta - sonsoensis Wagner / AfriGa - specimen ID — 1276 - specimen data - documented - 23.111.2006 1.-10. July 1995 Sonso River (Budongo Forest), Masindi, Uganda, Africa 1°45 N, 31°35°E (site information) COL 2006/219 ı PT 1.-10. July 1995 Sonso River (Budongo Forest), Masindi, Uganda, Africa 1°45°N, 31°35°E (site information) tutseki Wagner & Scherz, 2002: 359 (Afrocandezea) Holotype 9 - in: National Museums of Kenya (NMK), Nairobi COL 2007/235 1 PT Uganda, District Masindi - Budongo Forest n. Sonso - 1°45°N, 31°35°W; - 19.-30.V1.95 - Th. Wagner leg. / Paraty- pus / Afrocandezea tutseki - sp. nov. - Scherz & Wagner 2002 19.-30. June 1995 COL 2007/236 1 PT 11.-20. July 1995 COL 2007/237 1 PT 5.-15. January 1997 COL 2007/238-239 2PT September 1997 Sonso River (Budongo Forest), Masindi, Uganda, Africa 1245'N, 31°3S°E (exact gathering site information; error on original label!) COL 2007/240-243 4 PT 26. July — 22. August 1998 Kanyawara (Kibale National Park), Kabarole, Western, Uganda, Africa 034 'N, 30°20°E (gathering site: estimated value) upembaensis Wagner, 2000b: 234 (Monolepta) COL 2007/220 1 PT Congo belge: P.N.U. - Mbuye-Bala (1750 m.) - 1-7-1V-1948 - Mis. G.F. de Witte. 1500a / Paratypus / Monolepta - upem- baensis - Wagner / coll. - Th. Wagner 1.-7. April 1948 Mbuye-Bala (stream)(= Buye-Bala (stream)), Upemba, Parc National de la, Katanga, Democratic Republic of the Con- go, Afrıca 8°54'S, 26°59 E (gathering site: estimated value) ursulae Wagner, 2003: 67 (Monolepta) COL 2007/221, 257-258 3PT S.W. - AFRIKA - Okahandja 1.1956. - W. Hoesch leg., ded. - Eing.Nr.6, 1956. / Paratypus / Monolepta - ursulae - Th. Wagner January 1956 170 Karl-Heinz LAMPE, Dirk ROHWEDDER & Carola SCHMIDT: African Coleoptera Types collected by Th. Wagner Okahandja, Okahandja, Otjozondjupa, Namibia, Africa 21°59'S, 16°5’E (gathering site: estimated value) COL 2006/224, 265-267 3 PT 12. January 1985 COL 2006/268-270 3 PT 27. March 1988 Dóbra, Windhoek, Khomas, Namibia, Africa 22°20°S, 17°06E (gathering site: estimated value) COL 2006/271 1 PT 14.-22. January 1982 Kombat, Grootfontein, Otjozondjupa, Namibia, Äfrica 19°43°S, 17°43°E (gathering site: estimated value) COL 2006/272 1 PT 20. February - 6. March 1979 Gobabeb, Swakopmund, Erongo, Namibia, Africa 23°33°S, 15°02°E (gathering site: estimated value) COL 2006/223 , 264 2 PT 9.-15. March 1985 Naos, Rehoboth, Hardap, Namibia, Africa 23°12°S, 16%46'E (gathering site: estimated value) COL 2006/261 1 PT 15. December 1974 COL 2006/262 1PT 8.-12. March 1979 COL 2006/222, 259-260 3 PT 8.-13. January 1982 COL 2006/263 1 PT 17.-20. March 1982 Okahandja, Okahandja, Otjozondjupa, Namibia, Africa 21959'S, 165'E (gathering site: estimated value) COL 2006/273 1 PT 18.-20. March 1991 Osona, Okahandja, Otjozondjupa, Namibia, Africa 22°04 S, 16°53°E (gathering site: estimated value) COL 2006/274 ı PT 7. February 1975 COL 2006/275-276 2PT 8. February 1975 COL 2006/273 ı PT 2. March 1975 Windhoek, Windhoek, Khomas, Namibia, Africa 22°35°S, 17°05°E (gathering site: estimated value) COCCINELLIDAE brevipilosus Fúrsch, 1997: 18 (Nephus) COL 2006/13 Co HT (Fig. 13) Rwanda, Nyakabuye - Cyamudongo X.93 - Th. Wagner leg. / C.g. 12 / Holotypus - Nephus O” - brevipilosus - Fürsch, 1995 October 1993 Cyamudongo, Nyakabuye, Cyangugu, Rwanda, Africa 2°33°S, 28°59°E (gathering site: estimated value) COL 2006/14 109 PT October 1993 Karengera, Cyangugu, Rwanda, Africa 2°31°S, 29°02°E (gathering site: estimated value) COL 2006/15-16 29 PT October 1993 Cyamudongo, Nyakabuye, Cyangugu, Rwanda, Africa 2°33°S, 2859'E (gathering site: estimated value) brunneus Fúrsch, 1997: 20 (Pseudoscymnus) COL 2006/53 CO HT (Fig. 14) Rwanda, Nyakabuye - Cyamudongo X.93 - Th. Wagner leg. / Cg 11 / Holotypus - Pseudoscymnus - brunneus O - Fürsch, 1995 October 1993 Cyamudongo, Nyakabuye, Cyangugu, Rwanda, Africa 2°33°S, 28759 E (gathering site: estimated value) COL 2006/54-55, 57-60 6 PT COL 2006/56, 61 2.0 "Pr October 1993 Cyamudongo, Nyakabuye, Cyangugu, Rwanda, Africa 2°33°S, 28°59'E (gathering site: estimated value) carapacola Fürsch, 1997: 26 (Epilachna) COL 2006/41 Co HT (Fig. 15) Rwanda, 1700 m - Cyamudongo, X.93 - Th. Wagner leg. / C.g. 17 1 / Holotypus - Epilachna 0° - carapacola - Fürsch, 95 October 1993 Cyamudongo. Nyakabuye, Cyangugu, Rwanda, Africa 2°33°S, 28759 E (gathering site: estimated value) COL 2006/42 19 PT October 1993 Cyamudongo, Nyakabuye, Cyangugu, Rwanda, Africa 2°33'S, 2859 E (gathering site: estimated value) conspergata Fúrsch, 1997: 24 (Epilachna) COL 2006/43 oO HT (Fig. 16) Rwanda, Nyungwe - Kamiranzovu, 2000 m - Th. Wagner leg. X.93/C.g. 2 4 / Holotypus - Epilachna O” - conspergata - Fúrsch, 95 October 1993 Kamiranzovu, Nyungwe Forest, Cyangugu, Rwanda, Africa 2°29°S, 2909 E (gathering site: estimated value) COL 2006/44 1?PT COL 2006/45 1PT October 1993 Kamiranzovu, Nyungwe Forest, Cyangugu, Rwanda, Africa 2929'S, 2909 E (gathering site: estimated value) COL 2006/46 19 PT October 1993 Karengera, Cyangugu, Rwanda, Africa 2°31°S, 29°02°E (gathering site: estimated value) Bonner zoologische Beiträge 55 (2006) 171 fissilobis Fürsch, 2001b: 103-104 (Pharoscymnus) Holotype © - in: National Museums of Kenya (NMK), Nairobi COL 2006/107-115 9 PT Kenya, Nairobi, Langata, - 1°20°S/36°46°E, 1650 m, 12.- 14.1.1999, - Th. Wagner leg. / T.s.4 4 / Paratypus - Pharoscym- nus - fissilobis - Fürsch, 1999 12.-14. January 1999 Langata (Nairobi), Nairobi, Kenya, Africa 1°20°S, 36°46°E (site information) gilvella Fürsch, 2001b: 98-99 (Harmonia) COL 2006/1185 CO HT (Fig. 17) Uganda, Semliki Forest - 0°48’ N, 3098” E; 5.-12.11.97 - Th. Wagner leg. / C.a. S3 7 / Holotypus - Micraspis O' - gilvel- la - Fürsch, 1999 5.-12. Februay 1997 Semliki Forest, Bundibugyo, Western, Uganda, Africa 0°48°N, 30°08’E (site information) COL 2006/6-8, 1186-1190 8 PT 5.-12. Februay 1997 Semliki Forest, Bundibugyo, Western, Uganda, Africa 0°48°N, 30°08’E (site information) glabripilosus Fürsch, 1997: 22 (Scotoscymnus) COL 2006/62 oO HT (Fig. 18) Rwanda, 1700 m - Karengera, X.93 - Th. Wagner leg. /C.g.1 14 / Holotypus - Scotoscymnus - glabripilosus O - Fürsch, 1995 October 1993 Karengera, Cyangugu, Rwanda, Africa 2°31°S, 2902 E (gathering site: estimated value) COL 2006/63-64, 70, 73, 75-80 10 PT October 1993 Cyamudongo, Nyakabuye, Cyangugu, Rwanda, Africa 2°33°S, 2859 E (gathering site: estimated value) COL 2006/65, 67-69, 81 5iPT October 1993 Karengera, Cyangugu, Rwanda, Africa 2°31°S, 29°02°E (gathering site: estimated value) COL 2006/66, 72, 74 3 PT October 1993 Kamiranzovu, Nyungwe Forest, Cyangugu, Rwanda, Africa 2°29'S, 29°09’E (gathering site: estimated value) COL 2006/71 1 PT October 1993 Ibanda Makera, Rusumo, Kibungo, Rwanda, Africa 2°15°S, 3050 E (gathering site: estimated value) gracilis Fürsch, 1997: 20 (Ortalia) COL 2006/5 CO HT (Fig. 19) Rwanda, Rusumo - Ibanda Makera, X.93 - Th. Wagner leg. / Holotypus - Ortalia - gracilis © - Fürsch, 95 / Ortalia ba- sinigra Ma. - det. H. Fürsch 03 October 1993 Ibanda Makera, Rusumo, Kibungo, Rwanda, Africa 2°15°S, 30°50°E (gathering site: estimated value) goellnerae Fürsch, 2001b: 100 (Platynaspis) COL 2007/58-59 29 PT Uganda, Semliki Forest - 0°48°N, 30°8°E; 5.-12.11.97 - Th. Wagner leg. / Paratype - Platynaspis - goellnerae Y - Fürsch, 1999 5.-12. Februay 1997 Semliki Forest, Bundibugyo, Western, Uganda, Africa 0°48'N, 30°08’E (site information) kenyensis Fürsch, 2001b: 101-102 (Boschalis) Holotype G - in: National Museums of Kenya (NMK), Nairobi COL 2006/116 ı PT Mt. Kenya, NP-HQ, - 2550 m, 0°10.5°S/37°8.8°E; - 11.99 Th. Wagner leg. / P.1.17 2 / Paratypus - Boschalis kenyensis - Fürsch, 99 Februay 1999 Mount Kenya (mts.), Central, Kenya, Africa 0°10°S, 37°11’E imprecise data on original label!) COL 2006/117 1 PT January 1999 (exact gathering site information; Gatamayu, Kikuyu, Central, Kenya, Africa 0°58'S, 36°42°E (site information) COL 2006/118 1PT 18. Februay 1999 SE-Aberdare, Central, Kenya, Africa 0°23'S, 36°46'E (site information) magnifica Fúrsch, 2006: 114-115 (Cheilomenes) COL 2007/56 CO HT (Fig. 20) Kenya, Busumbuli, 1600 m - Kakamega Forest - 0,27%N / 34,88°E - W. Freund leg. X.01 / T.n.138 1 - IX.-X.2001 / Holotypus - Cheilomenes - magnifica O' - Fürsch, 2005 October 2001 Busumbuli River, Kakamega Forest, Western, Kenya, Africa 0°19'N, 34°52°E (exact gathering site information; imprecise data on original label!) COL 2007/57 January 2003 Kisere Forest, Kakamega Forest, Western, Kenya, Africa 0°24'N, 3454 E (site information) 19 PT maximus Fürsch, 1997: 22-24 (Scotoscymnus) COL 2006/87 CHT (Fig. 21) Rwanda, Nyakabuye - Cyamudongo X.93 - Th. Wagner leg. / Cg 9 / Holotypus I - Scotoscymnus - maximus - Fúrsch, 95 October 1993 Cyamudongo, Nyakabuye, Cyangugu, Rwanda, Africa 2°33°S, 2859'E (gathering site: estimated value) COL 2006/88, 90-106 18 PT October 1993 Cyamudongo, Nyakabuye, Cyangugu, Rwanda, Africa 2°33°S, 28°59°E (gathering site: estimated value) 172 Karl-Heinz LAMPE, Dirk ROHWEDDER & Carola SCHMIDT: African Coleoptera Types collected by Th. Wagner ENTE) Figs 19-27. Ortalia gracilis Fürsch, 1997 (19) Y HT, Cheilomenes magnifica Fürsch, 2006 (20) @ HT, Scotoscymnus maximus Fürsch, 1997 (21) 9 HT, Ortalia minima Fürsch, 2001 (22) & HT, Nephus rugulipennis Fürsch, 1997 (23) & HT, Scymnus rwan- densis Fúrsch, 1997 (24) & HT, Telsimia striata Fürsch, 1997 (25) & HT, Epilachna struwei Fürsch, 2001 (26) & HT, Boschalis wagneri Fürsch, 1997 (27) & HT: Habitus in dorsal view. Bonner zoologische Beitráge 55 (2006) 173 COL 2006/89 1 PT October 1993 Karengera, Cyangugu, Rwanda, Africa 2°31°S, 29°02°E (gathering site: estimated value) minima Fürsch, 2001b: 99 (Ortalia) COL 2006/4 CO HT (Fig. 22) Uganda, District Masindi - Budongo Forest n. Sonso - 1°45°N, 31°35°E; 15.-25.1.97 - Th. Wagner leg. / C.a.35 1 / Holotypus - Ortalia O” - minima - Fiirsch, 1999 15.-25. January 1997 Sonso River (Budongo Forest), Masindi, Uganda, Africa 1°45°N, 31°35°E (site information) montivagans Fürsch, 2001a: 91-92 (Nephus) Holotype G - in: National Museums of Kenya (NMK), Nairobi COL 2006/9, 11 2 PT Mt. Kenya, NP-HQ, - 2550 m, 0°10.5°S, 37°8.8°E; - 11.99 Th. Wagner leg. / Av 16 2 / Paratypus - Nephus - montivagans - Fürsch, 1999 Februay 1999 Mount Kenya (mts.), Central, Kenya, Africa OASIS MINE imprecise data on original label!) COL 2006/10 1 PT 18. Februay 1999 SE-Aberdare, Central, Kenya, Africa 0°23°S, 36°46°E (site information) (exact gathering site information; rubida Fúrsch, 2001b: 104 (Epilachna) Holotype @ - in: National Museums of Kenya (NMK), Nairobi COL 2006/31-40 10 PT Mt. Kenya, Chogoria Rte., - 0°09°S/37°26°E, 2800 m, - 25.11.1999, Th. Wagner leg. / Paratype - Epilachna - rubida - Fúrsch, 2000 25. Februay 1999 Chogoria Rte., Mount Kenya (mts.), Eastern, Kenya, Africa 0°09°S, 37726 'E (site information) rubridorsis Fürsch, 2001a: 92-93 (Nephus) Holotype ©’ - in: National Museums of Kenya (NMK), Nairobi COL 2006/17 19 PT Kenya Kikuyu, Gatamayu - 2330 m 0%58'S/36%42'E, - 2.99 leg. Th. Wagner / Paratypus - Nephus (Sidis) - rubridorsis Q - Fürsch 2000 Februay 1999 Gatamayu, Kikuyu, Central, Kenya, Africa 0°58'S, 36°42°E (site information) rugulipennis Fürsch, 1997: 18 (Nephus) COL 2006/12 CO HT (Fig. 23) Rwanda, Rusumo - Ibanda Makera, X.93 - Th. Wagner leg. / Holotypus - Nephus 0° - rugulipennis - Fürsch, 1995 October 1993 Ibanda Makera, Rusumo, Kibungo, Rwanda, África 2°15°S, 30°50°E (gathering site: estimated value) rwandensis Fúrsch, 1997: 16 (Scymnus) COL 2006/47 CHT (Fig. 24) Rwanda, Rusumo - [banda Makera, X.93 - Th. Wagner leg. /T.n.5 3 / Holotypus - Scymnus (Pullus) - rwandensis O” - Fursch, 1995 October 1993 Ibanda Makera, Rusumo, Kibungo, Rwanda, Africa 2°15°S, 30°50'E (gathering site: estimated value) COL 2006/48 - 52 5 PT October 1993 Ibanda Makera, Rusumo, Kibungo, Rwanda, Africa 2°15°S, 30°50°E (gathering site: estimated value) striata Fúrsch, 1997: 20-22 (Telsimia) COL 2006/131 CHT (Fig. 25) Rwanda, 1700 m - Cyamudongo, X.93 - Th. Wagner leg. / C.g.4 2 / Holotypus - Telsimia - striata O” - Fürsch, 1995 October 1993 Cyamudongo, Nyakabuye, Cyangugu, Rwanda, Africa 2°33°S, 28°59'E (gathering site: estimated value) COL 2006/132 19 PT COL 2006/133-134 2 PT October 1993 Cyamudongo, Nyakabuye, Cyangugu, Rwanda, Africa 2°33°S, 28°59°E (gathering site: estimated value) struwei Fürsch, 2001b: 105-106 (Epilachna) COL 2006/125 CHT (Fig. 26) Rwanda, Mont - Bisoke, X.92 / Holotypus - Epilachna - struwei O” - Fürsch, 1999 October 1992 Volcano Visoke (Bisoko), Ruhengeri, Rwanda, Africa Volcano National Park, 1°28°S, 29°29°E (gathering site: estimated value) COL 2006/126-130 5 PT October 1992 Volcano Visoke (Bisoko), Volcano National Park, Ruhengeri, Rwanda, Africa 1°28°S, 29°29°E (gathering site: estimated value) vulpecula Fúrsch, 2001a: 93 (Nephus) Holotype 9 - in: National Museums of Kenya (NMK), Nairobi COL 2006/18, 29 2 PT Kenya, Kikuyu, Gatamayu, - 2330 m 0°58°S/36°42’E - 11.99 Th. Wagner leg. / P.14 2 / Paratypus - Nephus - vulpecula - Fúrsch, 1999 Februay 1999 Gatamayu, Kikuyu, Central, Kenya, Africa 0°58°S, 36°42°E (site information) COL 2006/19-28, 30 11 PT 18. Februay 1999 SE-Aberdare, Central, Kenya, Africa 0°23'S, 36°46'E (site information) 174 Karl-Heinz LAMPE, Dirk ROHWEDDER & Carola SCHMIDT: African Coleoptera Types collected by Th. Wagner wagneri Fúrsch, 1995: 30 (Boschalis) COL 2006/119 CHT (Fig. 27) Rwanda, Nyakabuye - Cyamudongo X.93 - Th. Wagner leg. / C.g.9 3 / Holotype - Boschalis - wagneri O” - Fürsch, 1995 October 1993 Cyamudongo, Nyakabuye, Cyangugu, Rwanda, Africa 2°33°S, 28°59 E (gathering site: estimated value) COL 2006/120-124 5 PT October 1993 Cyamudongo, Nyakabuye, Cyangugu, Rwanda, Africa 2°33°S, 2859'E (gathering site: estimated value) wagneri Fürsch, 2001b: 103 (Scotoscymnus) Holotype O” - in: National Museums of Kenya (NMK), Nairobi, 3 paratypes dto. COL 2006/82-86 5PT Kenya, SE-Aberdare, - 0°23 S/36°46 E, 2750 m, - 18.11.99 Th. Wagner leg. / P.l. 263 / Paratypus — Scotoscymnus - wagneri - Fürsch, 1999 18. Februay 1999 SE-Aberdare, Central, Kenya, Africa 0°23°S, 36°46°E (site information) wagneri Fürsch, 2006a: 116; 2006b: 266 (Stethorus) Holotype O” - in: National Museums of Kenya (NMK), Nairobi, 32 paratypes dto. COL 2007/60-65 6PT Kenya, Kakamega F., - 0222 N/34950'E, 1600 m, - 7-11.11.99, Th. Wagner leg. / Paratypus - Stethorus - wagner! - Fürsch, 2005 7.-11. Februay 1999 Kakamega Forest, Western, Kenya, Afrıca 0°17 N, 34°53 E (exact gathering site information; imprecise data on original label!) COL 2007/66 1 PT October 2002 Busumbuli River, Kakamega Forest, Western, Kenya, Africa 0%19'N, 3452 E (exact gathering site information; imprecise data on original label!) COL 2007/67-68 2 PT October 2001 COL 2007/69-74 6 PT January 2002 COL 2007/75 1PT October 2002 COL 2007/76-78 3PT January 2003 Colobus Trail, Kakamega Forest, Western, Kenya, Africa 0°21 N, 3451 "E (exact gathering site information; imprecise data on original label!) COL 2607/79 1 PT January 2002 COL 2007/80-81 2 PT January 2003 Isiukhu River, Kakamega Forest, Western, Kenya, Africa 0°20°N, 34°53°E (exact gathering site information: imprecise data on original label!) COL 2007/82-84 3 PT January 2002 COL 2007/85-86 2 EX October 2002 COL 2007/87-88 2.PT, January 2003 Kisere Forest, Kakamega Forest, Western, Kenya, Africa 0°24°N, 34754 E (site information) COL 2007/89-94 6 PT January 2002 COL 2007/95-97 3PT January 2003 Yala River, Kakamega Forest, Western, Kenya, Africa O°12°N, 3453 E (exact gathering site information; imprecise data on original label!) DERMESTIDAE rusumoensis Háva & Herrmann, 2002: 254 (Orphinus) COL 2007/254 Co HT (Fig. 28) Rwanda, Rusumo - Ibanda Makera, X.93 - Th. Wagner leg. / Holotypus - Orphinus (Orphinus) - rusumoensis n. sp. O - det. J. Hava & A. Herrmann 2002 October 1993 Ibanda Makera, Rusumo, Kibungo, Rwanda, Africa 2°15°S, 30°50°E (gathering site: estimated value) wagneri Hava & Herrmann, 2002: 253 (Orphinus) COL 2007/255 Co HT (Fig. 29) Rwanda, Rusumo - Ibanda Makera, X.93 - Th. Wagner leg. / Holotypus - Orphinus (Orphinus) - wagneri n. sp. O” - det. J. Hava & A. Herrmann 2002 October 1993 Ibanda Makera, Rusumo, Kibungo, Rwanda, Africa 2°15°S, 30°50°E (gathering site: estimated value) HYDROPHILIDAE magnus Hebauer, 2002b: 6 (Allocotocerus) COL 2006/135-136 2 PT Kenya, Kakamega F., - 0°22°N/34°50°E, 1600 m, - 7-11.11.99, Th. Wagner leg. / Paratypus - Allocotocerus - magnus sp. n. - des. F. Hebauer 7.-11. Februay 1999 Kakamega Forest, Western, Kenya, Africa 0°17'N, 34253 E (exact gathering site information; imprecise data on original label!) COL 2006/137-138 ZPT 05.-15. Jan. 1997 Sonso River (Budongo Forest), Masindi, Uganda, Africa 1°45°N, 31°35°E (site information) Bonner zoologische Beitráge 55 (2006) 175 melinus Hebauer, 2001: 384 (Enochrus) COL 2006/141 1 PT Uganda District Masindi - Budongo Forest n. Sonso - 1°45°N, 31°35°W 19.-30.V1.95 - Th. Wagner leg. / Paraty- pus / Enochrus (Meth.) - melinus sp. n. - det. F. HEBAUER 19.-30. June 1995 Sonso River (Budongo Forest), Masindi, Uganda, Africa 1°45°N, 31°35°E (exact gathering site information; error on original label!) perssoni Hebauer, 2002a: 267 (Enochrus) COL 2006/140 1 PT Kenya, SE-Aberdare, - 0°23’N/36°46 E, 2750 m, - 16.- 21.11.1999 Th. Wagner leg. / Paratypus / Enochrus (Meth.) - perssoni sp. n. - det. F. HEBAUER 16.-21. Februay 1999 SE-Aberdare, Central, Kenya, Africa 0°23°N, 36°46°E (site information) wagneri Hebauer, 2002b: 14 (Helochares) COL 2006/139 1 PT Kenya, Kakamega F. - 0°22°N/34°50°E, 1600 m, - 7-11.11.99, Th. Wagner leg. / Paratypus - Helochares - wagneri sp. n. - des. F. Hebauer 7.-11. Februay 1999 Kakamega Forest, Western, Kenya, Africa 017 N, 34°53°E (exact gathering site information; imprecise data on original label!) LAEMOPHLOEIDAE wagneri Karner, 1997: 95-98 (Notolaemus) COL 2006/142 oO HT (Fig. 30) Rwanda, Rusumo - Ibanda Makera, X.93 - Th. Wagner leg. / T.n. 5 / Holotypus - Notolaemus - wagneri - n.sp. O' - M. Karner 1996 October 1993 Ibanda Makera, Rusumo, Kibungo, Rwanda, Africa 2°15°S, 30°50°E (gathering site: estimated value) COL 2006/143 109 PT COL 2006/144-148 59 PT October 1993 Ibanda Makera, Rusumo, Kibungo, Rwanda, Africa 2°15°S, 30°50°E (gathering site: estimated value) MELYRIDAE adrastus Evers, 1998: 63-64 (Sphinginopalpus) COL 2006/164 CHT (Fig. 31) Rwanda, Rusumo - Ibanda Makera, X.93 - Th. Wagner leg. / T.n.8 / Von Teclea nobilis gefoggt / Holotypus O” / Sphin- ginopalpus - adrastus Evers - A. Evers det. 1997 October 1993 Ibanda Makera, Rusumo, Kibungo, Rwanda, Africa 2°15°S, 3050 E (gathering site: estimated value) carapae Evers, 1998: 60-62 (Attalus) COL 2006/149,151,153 392 PT Rwanda, Nyungwe - Kamiranzovu, 2000 m - Th. Wagner leg. X.93 / C.g.2 / Von Carapa grandiflora gefoggt / Paratypus Q / Attalus - carapae Evers - A. Evers det. 1997 October 1993 Kamiranzovu, Nyungwe Forest, Cyangugu, Rwanda, Africa 2°29'S, 2909 E (gathering site: estimated value) COL 2006/150 19 PT October 1993 Cyamudongo, Nyakabuye, Cyangugu, Rwanda, Africa 2°33°S, 2859'E (gathering site: estimated value) COL 2006/152 109 PT October 1993 Karengera, Cyangugu, Rwanda, Africa 2°31°S, 2902 E (gathering site: estimated value) karengerae Evers, 1998: 62 (Attalus) COL 2006/154 19 AT Rwanda, 1700 m - Karengera, X.93 - Th. Wagner leg. /C.g.1 1 / Von Carapa grandiflora gefoggt / Allotypus Y / Attalus - karengerae Evers - A. Evers det. 1997 October 1993 Karengera, Cyangugu, Rwanda, Africa 2931'S, 2902 E (gathering site: estimated value) lateridentatus Evers, 1998: 64 (Colotes) COL 2006/165 CHT (Fig. 32) Rwanda, Rusumo - Ibanda Makera, X.93 - Th. Wagner leg. / T.n.6 / Von Teclea nobilis gefoggt / Holotypus O / O / Colotes lateridentatus Evers - A. Evers det. 1997 October 1993 Ibanda Makera, Rusumo, Kibungo, Rwanda, Africa 2°15°S, 30°50°E (gathering site: estimated value) COL 2006/166-170 592 PT October 1993 Ibanda Makera, Rusumo, Cyangugu, Rwanda, Africa 2°15°S, 30°50°E (gathering site: estimated value) stilifer Evers, 1998: 60 (Afrocarphurus) COL 2006/163 19 PT Rwanda, Karengera - Cyamudongo X.93 - Th. Wagner leg. /C.g.3 / Von Carapa grandiflora gefoggt / Paratypus @ / Afro- carphurus - stilifer Evers - A. Evers det.1997 / 1 Paratypus —Q behalten October 1993 Cyamudongo, Nyakabuye, Cyangugu, Rwanda, Africa 2°33°S, 2859 'E (gathering site: estimated value) tecleae Evers, 1998: 64-65 (Morphotroglops) COL 2006/162 12 AT Rwanda, Rusumo - Ibanda Makera, X.93 - Th. Wagner leg. / T.n.8 / Von Teclea nobilis gefoggt / Allotypus Q / Mor- photroglops - tecleae Evers - A. Evers det.1997 176 Karl-Heinz LAMPE, Dirk ROHWEDDER & Carola SCHMIDT: African Coleoptera Types collected by Th. Wagner Figs 28-36. Orphinus rusumoensis Hava & Herrmann, 2002 (28) 9 HT, Orphinus wagneri Hava & Herrmann, 2002 (29) O HT, Notolaemus wagneri Karner, 1997 (30) $ HT, Sphinginopalpus adrastus Evers, 1998 (31) & HT, Colotes lateridentatus Evers, 1998 (32) @ HT, Afroebaeus wagneri Evers, 1998 (33) & HT, Attalus wagneri Evers, 1998 (34) © HT, Stenus arborarius Puthz, 1999 (35) @ HT, Stenus wagnerianus Puthz, 1999 (36) © HT: Habitus in dorsal view. Bonner zoologische Beitráge 55 (2006) 177 October 1993 Ibanda Makera, Rusumo, Kibungo, Rwanda, Africa 2°15°S, 30°50°E (gathering site: estimated value) wagneri Evers, 1998: 65 (Afroebaeus) COL 2006/155 Oo HT (Fig. 33) Rwanda, Karengera - Cyamudongo X.93 - Th. Wagner leg. / C.g.4 / Von Carapa grandiflora gefoggt / Holotypus 9° / Afroebaeus - wagneri Evers - A. Evers det. 1997 October 1993 Karengera, Cyangugu, Rwanda, Africa 2°31'S, 29°02°E (gathering site: estimated value) COL 2006/158,160 29 PT COL 2006/159 10 PT October 1993 Karengera, Cyangugu, Rwanda, Africa 2°31°S, 29°02°E (gathering site: estimated value) COL 2006/161 19 PT October 1993 Cyamudongo, Nyakabuye, Cyangugu, Rwanda, Africa 2°33°S, 28°59'E (gathering site: estimated value) wagneri Evers, 1998: 62-63 (Attalus) COL 2006/156 Q HT (Fig. 34) Rwanda, Karengera - Cyamudongo X.93 - Th. Wagner leg. /C.g.1 / Von Carapa grandiflora gefoggt / Holotypus Q / Attalus - wagneri Evers - A. Evers det. 1997 October 1993 Karengera, Cyangugu, Rwanda, Africa 2°31°S, 29°02°E (gathering site: estimated value) COL 2006/157 19 PT October 1993 Kamiranzovu, Nyungwe Forest, Cyangugu, Rwanda, Africa 2°29°S, 2909 E (gathering site: estimated value) STAPHYLINIDAe arborarius Puthz, 1999a: 15-17 (Stenus) COL 2006/171 CHT (Fig. 35) Uganda, District Masindi - Budongo Forest n. Sonso - 1°45°N, 31°35°W 11.-20.VILOS - Th. Wagner leg. / T.n.17 4 / Puthz, 1998 / Stenus - arborarius - spec. nov. - det. V. Puthz 1998 / 1 pt behalten 11.-20. July 1995 Sonso River (Budongo Forest), Masindi, Uganda, Africa 1°45°N, 31°35°E (exact gathering site information; error on original label!) COL 2006/172 15.-25. January 1997 Sonso River (Budongo Forest), Masindi, Uganda, Africa 1°45°N, 31935'E (gathering site) 10 PT wagneri Puthz, 1999b: 21 (Megalopinus) COL 2007/279 Q HT (Fig. 36) Uganda District Masindi - Budongo Forest n. Sonso - 1°45°N, 31°35°W 19.-30.V1.95 - Th. Wagner leg. / T.r.3 1 / Holotypus / Megalopinus - wagneri - spec. nov. - det. V, Puthz 1998 19.-30. June 1995 Sonso River (Budongo Forest), Masindi, Uganda, Africa 1°45°N, 31°35°E (exact gathering site information; error on original label!) wagnerianus Puthz, 1999a: 18-19 (Stenus) COL 2006/173 CO HT (Fig. 37) Rwanda, Nyungwe - Kamiranzovu, 2000 m - Th. Wagner leg. X.93 / ©’ - Holotypus / Stenus - wagnerianus - spec. nov. - det. V. Puthz 1998 / 1 pt behalten October 1993 Kamiranzovu, Nyungwe Forest, Cyangugu, Rwanda, Africa 2°29°S, 2909 E (gathering site: estimated value) COL 2006/174 109 PT October 1993 Kamiranzovu, Nyungwe Forest, Cyangugu, Rwanda, Africa 2°29'S, 29%09'E (gathering site: estimated value) Acknowledgements. We thank Dr. Bradley J. Sinclair (ZFMK, Bonn) for checking the English text and Dr. Thomas Wagner (University Koblenz) for valuable information. REFERENCES ADLBAUER, K. 2006. Weitere neue Bockkäfer aus der Äthiopi- schen Region (Coleoptera, Disteniidae und Cerambycidae). Les Cahiers Magellanes 55: 1-21. Evers, M. J. A. 1998. Die kronenbewohnenden Malachiidae (Coleoptera) dreier Baumarten in Rwanda und Kivu. Bonner Zoologische Beitráge 48 (1): 59-66. FREUND, W. & WAGNER, TH. 2003. Revision of Bonesioides Laboissiere, 1925 (Coleoptera; Chrysomelidae; Galerucinae) from continental Africa. Journal of Natural History 37: 1915-1976. FRIESER, R. 1997. Teilergebnisse der von Thomas Wagner in Zen- tralafrika im Rahmen der Baumkronenbenebelung erbeuteten Anthribiden (Coleoptera: Anthribidae). Acta Coleopterologi- ca XI (1): 1120. FÜRSCH, H. 1995. Revision der Gattung Boschalis Weise, 1897, mit Beschreibung neuer Arten. (Coleoptera, Coccinellidae). Mitteilungen der Münchner Entomologischen Gesellschaft 85: 21-31 FÜRSCH, H. 1997. Coccinellidae (Coleoptera) aus Rwanda. Bon- ner Zoologische Beiträge 47 (1/2):13-39. FÜRSCH, H. 2001a. Zur Coceinelliden-Fauna der Wälder in Zen- tral- und Ostafrika, mit Beschreibungen neuer Arten (Cole- optera, Coccinellidae). Entomologische Zeitschrift 111 (3): 90-94. 178 Karl-Heinz LAMPE, Dirk ROHWEDDER & Carola SCHMIDT: African Coleoptera Types collected by Th. Wagner Furscu, H. 2001b. Zur Coccinelliden-Fauna der Wälder in Zen- tral- und Ostafrika, mit Beschreibungen neuer Arten (Cole- optera, Coccinellidae). Entomologische Zeitschrift 111 (4): 98-107. FÜRSCH, H. 2006a. Zur Marienkáferfauna des Kakamega Forest in Kenya mit biogeographischen Anmerkungen (Coleoptera, Coccinellidae). Entomologische Zeitschrift 116 (3): 113-117. FÜRSCH, H. 2006b. Die afrotropischen Arten von Stethorus Wei- se, 1885 (Coleoptera: Coccinellidae, Seymninae). Entomolo- gische Zeitschrift 116 (6): 261-268. HAvA, J. & HERRMANN, A. 2003. Orphinus wagneri sp. n. and Orphinus rusomoensis sp. nov. (Coleoptera: Dermestidae) from Rwanda. Entomologische Zeitschrift 113 (8): 253-254 HEBAUER, F. 2001. Taxonomische Studien zur Hydrophiliden- Gattung Enochrus Thomson, 1859. 2. Teil : Die afrikanischen Arten der Untergattung Methydrus Rey, 1885. A: Die Enoch- rus meracus-Gruppe (Coleoptera, Hydrophilidae). Beiträge zur Entomologie 51 (2): 375-391. HEBAUER, F. 2002a. Taxonomische Studien zur Hydrophiliden- Gattung Enochrus Thomson, 1859. — 3. Teil: Die afrikanis- chen Arten der Untergattung Methydrus Rey, 1885. B.: Die Enochrus natalensis-Gruppe. Beiträge zur Entomologie 52 (1): 255-269. HEBAUER, F. 2002b. New Hydrophilidae of the Old World (Coleoptera, Hydrophilidae). Acta Coleopterologica XVIII (3): 3-24. KARNER, M. 1997. A new species of Notolaemus Lefkovitch from Rwanda (Coleoptera: Laemophloeidae). Bonner Zoologische Beiträge 47 (1-2): 95-98. MIDDELHAUVE, J. & WAGNER, TH. 2001. Revision of Afrocra- nia (Coleoptera: Chrysomelidae, Galerucinae). Part I: Species in which the males have head cavities or extended elytral ex- trusions. European Journal of Entomology 98: 511-532. Putuz, V. 1999a. Zwei neue afrikanische Arten der Gattung Ste- nus Latreille, 1796 und eine taxonomische Bemerkung (Col., Staphylinidae). Zeitschrift der Arbeitsgemeinschaft Osterrei- chischer Entomologen 51 (1): 15-20. Putuz, V. 1999b. Megalopinus wagneri sp.n. aus Uganda (Co- leoptera, Staphylinidae). Zeitschrift der Arbeitsgemeinschaft Österreichischer Entomologen 51 (1): 21-24. WAGNER, TH. 2000a. New Monolepta species (Coleoptera: Chrysomelidae, Galerucinae) from Eastern Africa. Entomol- ogische Zeitschrift 110 (2): 34-40. WAGNER, TH. 2000b. Revision of Afrotropical Monolepta Chevrolat, 1837 (Coleoptera, Chrysomelidae, Galerucinae). Part I: Species with red and black coloured elytra, pronotum and head, with description of new species. Entomologische Zeitschrift 110 (8): 226-237. WAGNER, TH. 200la. New Monolepta species (Coleoptera, Chrysomelidae, Galerucinae) from Central and Southern Africa. Entomologische Blátter 96: 199-209. WAGNER, TH. 2001b. Revision of Afrotropical Monolepta Chevrolat, 1837 (Coleoptera: Chrysomelidae, Galerucinae). Part II: Species with red elytra, pronotum and head, with de- scriptions of new species. Bonner Zoologische Beiträge 50 (1/2): 49-65. WAGNER, TH. 2002. Revision of Afrotropical Monolepta species (Coleoptera, Chrysomelidae, Galerucinae). Part III: Species with red elytra and yellow prothorax, including descriptions of new species. Deutsche Entomologische Zeitschrift 49 (1): 27-45. WAGNER, TH. & SCHERZ, X. 2002. Afrocandezea gen. nov. from tropical Africa (Coleoptera: Chrysomelidae, Galerucinae). En- tomologische Zeitschrift 112 (12): 357-362. WAGNER, TH. 2003. Revision of afrotropical Monolepta Chevro- lat, 1837 (Coleoptera, Chrysomelidae, Galerucinae). — Part IV: Species wıth red head and thorax and black elytra or black elytra with red apex, with description of new species. Anna- les Sciences Zoologiques, Miscellanea 49: 37-89. WAGNER, TH. 2005. Revision of the vincta Species-group of Monolepta Chevrolat, 1837 from Africa, Arabia and the Near East (Coleoptera: Chrysomelidae, Galerucinae). Bonner Zo- ologische Beitráge 53 (1/2): 255-282. Authors’ addresses: Karl-Heinz LAMPE (corresponding author), Dirk ROHWEDDER & Carola SCHMIDT, Zoologi- sches Forschungsmuseum Alexander Koenig (ZFMK), Adenauerallee 160, 53113 Bonn, Germany. E-Mail: k.lampe.zfmk(@uni-bonn.de Received: 22.12.2006 Accepted: 08.01.2007 Corresponding editor: M. Schmitt Bonner zoologische Beiträge werden publiziert im Eigenverlag Zoologisches Forschungsmuseum Alexander Koenig, Adenauer- allee 160, D-53113 Bonn (Germany). Die Zeitschrift erscheint mit vier Heften im Jahr, zum Preis von 11,50 € je Heft bzw. 46,- € je Band incl. Versand. Korrespon- denz betreffend Abonement, Kauf oder Tausch bitten wir zu richten an die Bibliothek, Zoologisches Forschungsmuseum Alex- ander Koenig, Adenauerallee 160, D-53113 Bonn (Germany); E-Mail: d.steinebach.zfmk@uni-bonn.de. © Zoologisches Forschungsmuseum Alexander Koenig (ZFMK), Bonn, Germany. ISSN 0006-7172 Satz: Eva-Maria Levermann, Kaiserstr. 151, D-53113 Bonn (Germany); E-Mail: emlevermann@netcologne.de. Druck: JF@CARTHAUS, Bonn, Germany. Bonner zoologische Beiträge Editorial Board (in brackets editorial competence) Prof.Dr. Michael SCHMITT (editor-in-chief, Coleoptera, theory) Tel.: +49 228-9122 286, Fax: +49 228-9122 332; E-Mail: m.schmitt.zfmk@uni-bonn.de. Y = . Renate VAN DEN ELZEN (Vertebrata, except Mammalia), Tel.: +49 228-9122 231, Fax: +49 228-9122 212; E-Mail: r.elzen.zfmk@uni-bonn.de. Y Ss . Bernhard A. HUBER (Invertebrata, except Insecta), Tel.: +49 228-9122 294, Fax: +49 228-9122 212; E-Mail: b.huber.zfmk@uni-bonn.de. Dr. Gustav PETERS, (Mammalia, bioacustics), Tel.: +49 228-9122 262, Fax: +49 228-9122 212; E-Mail: g.peters.zfmk@uni-bonn.de. Dr. la Bradley S. SINCLAIR (Diptera, language advisor), Entomology Laboratory - CFIA, K.W. Neatby Bldg., C.E.F. 960 Carling Ave., Ottawa, ON, Canada KIA 0C6; E-Mail: sinclairb(@inspection.ge.ca. Dr. Dieter STUNING (Insecta, except Coleoptera and Diptera), Tel.: +49 228-9122 220, Fax: +49 228-9122 212; E-Mail: d.stuening.zfmk(@uni-bonn.de. Editorial office: Adenauerallee 160, D-53113 Bonn, Germany. Advisory Board Pro. Dr. Miguel Angel ALONZO-ZARAZAGA, Museo nacional, Ciencias Naturales, E-28006 Madrid; E-Mail: zarazaga@mncn.csis.es. Prof. Dr. Ulrike Aspöck, Naturhistorisches Museum, 2. Zoologische Abteilung (Insekten), Postfach 417, A-1014 Wien; E-Mail: ulrike.aspoeck@nhm-wien.ac.at. Prof. Dr. Paolo Aupisio, Universita di Roma “La Sapienza”, Dipl. Biol. Anim e dell’Uomo (Zool.), Viale dell’Universitä 32, 1-00185 Roma, Te.: +39 6-49914744; E-Mail: paolo.audisio@uniromal.it. Prof. Dr. Aaron M. BAUER, Villanova University, Department of Biology, 800 Lancaster Avenue, Villanova, PA 19085-1699, USA, Tel: +1-610-519-4857, Fax: +1-610-519-7863; E-Mail: aaron.bauer@villanova.edu. Dr. Júrgen HAFFER, Tommesweg 60, D-45149 Essen, Tel.: +49 201-710426; E-Mail: j.haffer@web.de. Dr. Jeremy D. HoLLOwaY, The Natural History Museum, Department of Entomology, Cromwell Road, London, SW7 SBD, U.K.; E-Mail: j.holloway(@nmh.ac.uk. Dr. Marion KOTRBA, Zoologische Staatssammlung, Diptera, Münchhausenstr. 21, D-81247 München, Tel.: +49 89-8107 147, Fax: +49 89-8107 300; E-Mail: marion.kotrba@zsm.mwn.de. Prof. Dr. Boris KRYSTUFEK, Slovenian Museum of Natural His- tory, P. O. Box 290, SI-1001 Ljubljana; E-Mail: boris.krystufek@zrs.upr.si. Pro. Dr. Sven O. KULLANDER, Swedish Museum of Natural His- tory, Department of Vertrebrate Zoology, P. O. Box 50007, SE-104 05 Stockholm; E-Mail: sven.kullander@nrm.se. Prof. Dr. Steven PERRY, Rheinische Friedrich-Wilhelms-Univer- sität, Institut für Zoologie, Poppelsdorfer Schloss, D-53115 Bonn, Tel: +49 228-73 3807; E-Mail: perry@uni-bonn.de. Dr. Wolfgang SCHAWALLER, Staatliches Museum für Naturkunde, Rosenstein 1, D-70191 Stuttgart, Germany, Tel.: +49 711-8936 221, Fax: +49 711-8936 100; E-Mail: schawaller.smns(@naturkundemuseum-bw.de. Dr. W. David Sissom, Dept. of Life, Earth and Environmental Sciences, W. Texas A. & M. University, WTAMU Box 60808, Canyon, Texas 79016, USA; E-Mail: dsissom@wtamu.edu. = ". Miguel VENCES, Technische Universitat Carolo-Wilhelmi- na, Zool. Inst., Abt. Evolutionsbiol., Mendelssohnstr. 4, D-38106 Braunschweig, Tel.: +49 531-391 3231, Fax: +49 531-391 3222; E-Mail: m.vences(@tu-braunschweig.de. PD Dr. Heike WAGELE, Rheinische Friedrich-Wilhelms-Univer- sität, Institut für Evolutionsbiologie und Ökologie, D-53121 Bonn, Tel.: +49 228 73 5159, Fax: +49 234-322 4114; E-Mail: hwaegele@evolution.uni-bonn.de. Dr. Erich WEBER, Eberhard-Karls-Universität, Zoologische Schausammlung, Sigwartstr. 3, D-72076 Tübingen, Germany; E-Mail: erich.weber@uni-tuebingen.de. Inhalt A a DI 3 9088 I WAGNER, Philipp & BOHME, Wolfgang: A new species of the genus Trapelns Cuvier, 1816 (Squamata: Agamidae) from arid central Africa ZACHOS, Frank E:; OTTO, Marthe; HMWE, San San & HARTL, Günther B.: A Populationsgenetische Untersuchungen zur Differenzierung des schleswig-holsteinischen Rehwildes (Capreolus capreolus Linnaeus, 1758; Artiodactyla: Cervidae) HAHN, Ingo: Bioacoustic Characteristics and Population Numbers of Endemic Cinclodes oustaleti baeckstroemii (Aves: Furnariidae) Lónnberg, 1921 of Alejandro Selkirk Island, Chile HAHN, Ingo: Biogeographical Isolation and Bioacoustics: the Juan Fernandez Firecrown, Sephanoides fernandensis (Aves: Trochilidae) (King, 1831), of Robinson Crusoe Island, Chile VAN ROOSMALEN, Marc G. M.; FRENZ, Lothar; VAN HOOFT, Pim; DE IONGH, Hans H. & LEirs, Herwig: A New Species of Living Peccary (Mammalia: Tayassuidae) from the Brazilian Amazon DUBATOLOV, Vladimir & HOLLOWAY, Jeremy D.: A new species of the Creatonotos transiens-group (Lepidoptera: Arctiidae) from Sulawesi, Indonesia WAGNER, Philipp & BOHME, Wolfgang: Herpetophauna Kakamegensis: The amphibians and reptiles of Kakamega Forest, western Kenya AUDISIO, Paolo & DE BIASE, Alessio: A new Meligethes of the ML pubescens species-group from South Africa (Coleoptera, Nitidulidae, Meligethinae) FORSCHLER, Marc Imanuel & SIEBENROCK, Karl Heinz: Morphological differentiation of mainland Citril Finches, Carduelis [citrinella) citrinella and insular Corsican (Citril) Finches, Cardnelis [citrinella) corsicanus ROHWEDDER, Dirk; LAMPE, Karl-Heinz & SCHMIDT, Carola: African Coleoptera Type Specimens collected by Thomas Wagner in the collection of the ZFMIX Buchbesprechungen / Book Reviews JUNKER, Thomas (2004): Die zweite Darwinsche Revolution — Geschichte des Synthetischen Darwinismus in Deutschland 1924 bis 1950 (M. SCHMITT, Bonn) Merz, B. (ed.) 2006. Phylogeny, Taxonomy, and Biology of Tephritoid flies (Diptera, Tephritoidea) (B. J. SINCLAIR, Ottawa) BRAUN, Monika & DIETERLEN, Fritz (Hrsg,): Die Säugetiere Baden-Württembergs (G. PETERS, Bonn) Titelbild/ Cover illustration: Wild giant peccary, Pecari maximus sp. nov. (see contribution of Marc G. M.VAN ROOSMALEN, pp. 105-112) || 01431 81 89 101 105 159 88 104 158 | Jill { i Bonner zoologische Beiträge Special Issue: Proceedings of the 2nd International Workshop on Opisthobranchia, ZEMK, Bonn, Germany, September 20th to 22nd, 2006 ; p 0 ze A y Y \| / A des OW) RT YH BD air a E A Leere ck 24 Guest Editor: Heike Wägele Herausgegeben vom Zoologischen Forschungsmuseum Alexander Koenig, Bonn Band >> Heft 3/4 200 / (2006) ZU Leibniz Gemeinschaft Bonner zoologische Beiträge werden publiziert im Eigenverlag Zoologisches Forschungsmuseum Alexander Koenig, Adenauer- allee 160, D-53113 Bonn (Germany). Die Zeitschrift erscheint mit vier Heften im Jahr, zum Preis von 11,50 € je Heft bzw. 46,- € je Band incl. Versand. Korrespon- denz betreffend Abonement, Kauf oder Tausch bitten wir zu richten an die Bibliothek, Zoologisches Forschungsmuseum Alex- ander Koenig, Adenauerallee 160, D-53113 Bonn (Germany); E-Mail: d.steinebach.zfmk@uni-bonn.de. © Zoologisches Forschungsmuseum Alexander Koenig (ZFMK), Bonn, Germany. ISSN 0006-7172 Satz: Eva-Maria Levermann, Kaiserstr. 151, D-53113 Bonn (Germany); E-Mail: emlevermann@netcologne.de. Druck: JFoCARTHAUS, Bonn, Germany. Bonner zoologische Beitrage Editorial Board (in brackets editorial competence) Prof. Dr. Michael SCHMITT (editor-in-chief, Coleoptera, theory) Tel.: +49 228-9122 286, Fax: +49 228-9122 332; E-Mail: m.schmitt.zfmk(wWuni-bonn.de. Q ". Renate VAN DEN ELZEN (Vertebrata, except Mammalia), Tel.: +49 228-9122 231, Fax: +49 228-9122 212; E-Mail: r.elzen.zfmk(@uni-bonn.de. = . Bernhard A. HUBER (Invertebrata, except Insecta), Tel.: +49 228-9122 294, Fax: +49 228-9122 212; E-Mail: b.huber.zfmk(@uni-bonn.de. = . Gustav PETERS, (Mammalia, bioacustics), Tel.: +49 228-9122 262, Fax: +49 228-9122 212; E-Mail: g.peters.zfmk(@uni-bonn.de. 2 . Bradley S. SINCLAIR (Diptera, language advisor), Entomology Laboratory - CFIA, K.W. Neatby Bldg., C.E.F. 960 Carling Ave., Ottawa, ON, Canada KIA 0C6; E-Mail: sinclairb(@inspection.ge.ca. D . Dieter STUNING (Insecta, except Coleoptera and Diptera), Tel.: +49 228-9122 220, Fax: +49 228-9122 212; E-Mail: d.stuening.zfmk(@uni-bonn.de. Editorial office: Adenauerallee 160, D-53113 Bonn, Germany. Advisory Board Prof. Dr. Miguel Angel ALONZO-ZARAZAGA, Museo nacional, Ciencias Naturales, E-28006 Madrid; E-Mail: zarazaga(@@mnen.csis.es. Prof. Dr. Ulrike Aspöck, Naturhistorisches Museum, 2. Zoologische Abteilung (Insekten), Postfach 417, A-1014 Wien; E-Mail: ulrike.aspoeck(@nhm-wien.ac.at. Prof. Dr. Paolo Aupisıo, Universita di Roma “La Sapienza”, Dipl. Biol. Anım e dell’Uomo (Zool.), Viale dell’Universitä 32, 1-00185 Roma, Te.: +39 6-49914744; E-Mail: paolo.audisio@uniroma it. Prof. Dr. Aaron M. BAUER, Villanova University, Department of Biology, 800 Lancaster Avenue, Villanova, PA 19085-1699, USA, Tel: +1-610-519-4857, Fax: +1-610-519-7863; E-Mail: aaron.bauer@villanova.edu. Dr. Jürgen HAFFER, Tommesweg 60, D-45149 Essen, Tel.: +49 201-710426; E-Mail: j.haffer@web.de. Dr. Jeremy D. HoLLowAY, The Natural History Museum, Department of Entomology, Cromwell Road, London, SW7 5BD, U.K.; E-Mail: j.holloway@nmh.ac.uk. Dr. Marion KoTRBA, Zoologische Staatssammlung, Diptera, Münchhausenstr. 21, D-81247 München, Tel.: +49 89-8107 147, Fax: +49 89-8107 300; E-Mail: marion.kotrba@zsm.mwn.de. Prof. Dr. Boris KRYSTUFEK, Slovenian Museum of Natural His- tory, P. O. Box 290, SI-1001 Ljubljana; E-Mail: boris.krystufek@zrs.upr.si. Pro. Dr. Sven O. KULLANDER, Swedish Museum of Natural His- tory, Department of Vertrebrate Zoology, P. O. Box 50007, SE-104 05 Stockholm; E-Mail: sven.kullander@nrm.se. Prof. Dr. Steven PERRY, Rheinische Friedrich-Wilhelms-Univer- sität, Institut für Zoologie, Poppelsdorfer Schloss, D-53115 Bonn, Tel: +49 228-73 3807; E-Mail: perry(@uni-bonn.de. Dr. Wolfgang SCHAWALLER, Staatliches Museum für Naturkunde, Rosenstein 1, D-70191 Stuttgart, Germany, Tel.: +49 711-8936 221, Fax: +49 711-8936 100; E-Mail: schawaller.smns@naturkundemuseum-bw.de. Dr. W. David Sissom, Dept. of Life, Earth and Environmental Sciences, W. Texas A. & M. University, WTAMU Box 60808, Canyon, Texas 79016, USA; E-Mail: dsissom@wtamu.edu. Dr. Miguel VeNces, Technische Universität Carolo-Wilhelmi- na, Zool. Inst., Abt. Evolutionsbiol., Mendelssohnstr. 4, D-38106 Braunschweig, Tel.: +49 531-391 3231, Fax: +49 531-391 3222: E-Mail: m.vences@tu-braunschweig.de. Prof. Dr. Heike WAGELE, Rheinische Friedrich-Wilhelms-Uni- versität, Institut für Evolutionsbiologie und Ökologie, D-53121 Bonn, Tel.: +49 228 73 5159, Fax: +49 234-322 4114; E-Mail: hwaegele@evolution.uni-bonn.de. Dr. Erich WeBer, Eberhard-Karls-Universitat, Zoologische Schausammlung, Sigwartstr. 3, D-72076 Tubingen, Germany; E-Mail: erich.weber@uni-tuebingen.de. Bonner zoologische Beiträge Band 55 (2006) Heft 3/4 Seiten 181-190 Bonn, November 2007 Distribution of homarine in some Opisthobranchia (Gastropoda: Mollusca)* Sven AFFELD!)2), Heike WAGELE2)3), Conxita AVILA, Stefan KEHRAUS!) & Gabriele M. KóNIG!) Institut für Pharmazeutische Biologie, Universität Bonn, Bonn, Germany “Institut für Evolutionsbiologie und Okologie, Universität Bonn, Bonn, Germany 3Zoologisches Forschungsmuseum Alexander Koenig, Bonn, Bonn, Germany 4Dept. of Animal Biology (Invertebrates), Facultat de Biologia, Universitat de Barcelona, Barcelona, Spain *Paper presented to the 2nd International Workshop on Opisthobranchia, ZFMK, Bonn, Germany, September 20th to 22nd, 2006 Abstract. Homarine, a zwitterionic natural product, ıs only known from some marine invertebrates, including the opi- sthobranch Phestilla lugubris (Klein 1986 in KARUSO 1987). This natural product is described as having antipredatory and antifouling effects. In the current study we present new results on the distribution of homarine in Opisthobranchia and in a few of their food organisms. Our results indicate that homarine is mainly found in members of the Cladobran- chia (with one exception) and is probably derived from enidarian food sources. Other investigated opisthobranch taxa, which feed on a variety of other marine organisms (algae, Porifera, Bryozoa etc.), lack homarine. An exception is the ce- phalaspid Aglaja tricolorata in which homarine was detected, and foraging of this predatory slug on homarine contai- ning cladobranchs seems likely. It is shown in Marionia blainvillea that the characteristic glandular cells in the epider- mis of the Dendronotoidea are not adaptations to store homarine. A Keywords. Natural products, Cnidaria, food organisms, Cladobranchia, Marionia blainvillea, histoldgy. 1. INTRODUCTION A wide variety of natural products is known from Opistho- branchia (KARUSO 1987; AvILA 1995; CIMINO et al. 1999; CIMINO et al. 2001; CIMINO & GAVAGNIN 2006; WAGELE et al. 2006). Some of the opisthobranchs are of particular interest due to their extreme toxicity and the high struc- tural diversity of their secondary metabolite content. One of these groups is the algae- or cyanobacteria-feed- ing taxon Anaspidea, which store a high diversity of sec- ondary metabolites. Apart from terpenes, which are the major structural class in Anaspidea, bioactive polyketides and peptides were also found (CAREFOOT 1987; YAMADA & KIGOSHI 1997; CIMINO et al. 2001). One of these pep- tides, i.e. dolastatin 10, showed potent cytotoxic activity. In clinical studies (phase II), however it had no effect nei- ther against advanced colorectal cancer (SAAD et al. 2002), advanced pancreaticobiliary cancers (KINDLER et al. 2005), nor advanced breast cancer (PEREZ et al. 2005). The Sacoglossa, which mainly feed on algae of the ta- xon Chlorophyta, obtain secondary metabolites from their food. One of these compounds, the depsipeptide kahalalide F that Elysia rufescens takes up from its food alga Bry- opsis spec. and algal associated bacteria is currently in a Dana PO, Lie f (ll a clinical trial (phase II) against hepatocellular carcinoma, non-small cell lung cancer and melanoma (HAMANN 2004). In some cases these sequestered chemicals are bio- transformed by the slugs. Thus Mediterranean Oxynoe oli- vacea, Lobiger serradifalci and Ascobulla fragilis, all of which feeding on Caulerpa prolifera, take up the algal sesquiterpenoid caulerpenyne and modify it into oxytox- in-1 and -2, which have an enhanced deterrent effect against fish compared to caulerpenyne (CIMINO et al. 1990; GAVAGNIN et al. 1994a). According to CIMINO & GHISELIN (1998), CIMINO et al. (1999) and MARIN & Ros (2004) highly derived sacoglossans are able to form their defen- sive compounds de novo. E.g. GAVAGNIN et al. (1994b) showed that the polypropionate elysione is produced by Elysia viridis itself. Within the Nudibranchia, the sponge-feeding Doridoidea are probably the group among the Opisthobranchia best examined for the presence of secondary metabolites, most of which are dietary derived terpenes (see e.g. the reviews Karuso et al. 1987; AvILA 1995; CIMINO et al. 1999; Gav- AGNIN & FONTANA 2000; GARSON 2006; WAHIDULLAH et al. 2006; MIYAMOTO 2006). De novo biosynthesis, how- ever, has also been shown, e.g., for the sesquiterpenes 182 Sven AFFELD et al.: Homarine in Opisthobranchia polygodial in Dendrodoris limbata (CIMINO et al. 1983) and ent-pallescensin A in Doriopsilla areolata (GAVAGNIN et al. 2001). In contrast, the nudibranch taxon Cladobranchia has been poorly investigated chemically. Traditionially, Clado- branchia are divided into three taxa, the monophyletic Ae- olidoidea and Dendronotoidea, and the paraphyletic Arminoidea (see WAGELE & WILLAN 2000). Members of the Cladobranchia mainly feed on octocorals and hydro- zoans (MCDONALD & NYBAKKEN 1997). Within the Armi- noidea few terpenes have been detected. The briarane diterpenoids verecynarmin A to G and the cembranoid pre- verecynarmin are known from Armina maculata and its prey organism, the octocoral Veretillum cynomorium (GUERRIERO et al. 1987, 1988, 1990). Leminda millecra exhibits four different sesquiterpenoids, millecrone A and B, as well as millecrol A and B (CIMINO et al. 2001). They all are sequestered while feeding on alcyonariids. Further investigation revealed more natural compounds: isofura- nodiene, (+)-8-hydroxycalamenene, algoafuran, cubebenone, and a series of seven triprenylquinones and hydroquinones. Alcvonium fauri and the gorgonian Lep- togorgia palma seem to be the source of at least some of these compounds (McPHAIL et al. 2001). Origin of jano- lusimide, extracted from Janolus cristatus (SODANO & SPINELLA 1986) seems to be unknown, a de novo biosyn- thesis can not be excluded (CIMINO et al. 2001). Terpenes are also present in Dendronotoidea. Tritonia hamnerorum e.g. sequesters the feeding deterrent sesquiterpene julieannafuran from its food, which is the sea fan Gorgonia ventalina (CRONIN et al. 1995). Sever- al terpenes (rubifolide, pukalide, cuparane sesquiter- penoides) and other compounds (tochuinyl acetate and its dihydro-derivative, ptilosarcenone) are collected from Tochuina tetraquetra. All these products seem to be food derived (CIMINO et al. 2001). Further compounds in oth- er members of Dendronotoidea are summarized by CIMI- no et al. (2001): An unknown Tritonia sp. revealed a prostanoid, also found in the octocoral Telesto, and Tri- toniella belli sequestered a chimyl alcohol from the oc- tocoral Clavularia frankliniana. De novo biosynthesis of terpenes is recorded from one member of the Dendrono- toidea: BARSBY et al. (2002) showed through feeding ex- periments with [1,2-!3C5] acetate that Melibe leonina pro- duces at least one of its defensive metabolites (2,6-di- methyl-5-heptenal) itself. In the Aeolidoidea terpenes are only known from the genus Phyllodesmium. P. longicirrum accumulates three cem- branoid diterpenes, trocheliophorol, thunbergol and epoxythunbergol, from the soft coral Sarcophyton troche- liophorum (COLL et al. 1985). P. guamensis specifically sequesters the cembranoid | 1-beta-acetoxypukalide from its preferred prey, the soft coral Sinularia maxima (SLAT- TERY et al. 1998). Only very few records for other com- pounds are available. CIAVATTA et al. (1996) discovered two prenylchromanols (1 and 2) and a prenyl-p-hydroxy acid in the skin of Cratena peregrina. None of these com- pounds was detected in the preferred food of C. peregri- na, namely the hydrozoan Eudendrium racemosum. On the other hand, other metabolites, usually not considered as deterrent to predators have been isolated from Euden- drium and its aeolid predators: carotenoids from Flabel- lina iodinea (MCBETH 1972), sterols from Cratena pere- grina, Flabellina affinis and F. lineolata (CIMINO et al. 1980), carotenoids and alkaloids from Phestilla melanobrachia (OKUDA et al. 1982), sterols and alkaloids from P. lugubris (TARGETT 1983; KARUSO 1987). KARU- so (1987, after Klein 1986) reported also the presence of homarine in the latter species. Members of the Aeolidoidea are known for storing enido- cysts from their prey organisms in special structures lo- cated at the cerata tips (see Fig. 1A). Here, a cnidosac is formed as part of the digestive gland, and in the cells of this sac, intact cnidocysts are incorporated and maintained functional. It is assumed that the slugs use these cnidosysts as a major means of defense (e.g. EDMUNDS 1966; GREEN- WOOD & MARISCAL 1984a, b). This assumption is support- ed by the studies of FRICK (2003), showing that Flabelli- na verrucosa responds to the presence of predators by the variation of nematocyst incorporation. The incorporation of enidocysts is regarded as the main key character in the evolution of the Aeolidoidea (WÄGELE 2004). Nevertheless, in some cases secondary metabolites seem to be of importance for aeolids as well, e.g. in the genus Phyllodesmium where no incorporation of cnido- cysts takes place (SLATTERY et al. 1998). The same holds true for Phestilla lugubris, which shows no cnidocysts in their enidosacs, but homarine was found (KARUSO 1987, after Klein 1986). Homarine was first isolated in 1933 by HOPPE-SEYLER. Its biological function has been controversially discussed ever since. This product seems to be restricted to marine or- ganisms, and within the Metazoa it is only found in in- vertebrates (CARR et al. 1996). Therefore it was suspect- ed that its purpose lies in osmoregulation. BEERS (1967) studied the distribution of homarine in Crustacea, Echi- nodermata and Tunicata and from these results infered that the presence of homarine is “in basic accord with a role in cellular osmotic phenomena”. DALL (1971) though could not find any contribution of homarine in osmotic processes in decapod crustaceans. Nevertheless, it is wide- ly accepted that homarine serves as an osmolyte in ma- rine algae. Initial studies have been carried out by DICK- SON & KirST (1986) with the green alga Platvmonas sub- Bonner zoologische Beiträge 55 (2006) 183 cordiformis. SLATTERY et al. (1997) showed that the soft coral Gersemia antarctica releases a potent antimicrobial mixture of organic compounds into the surrounding sea- water, and homarine is the compound responsible for most of this biological activity. In the tissues of the anthozoans Actinia equina and Calliactis parasitica, homarine was de- tected at concentrations of 1.5 and 2.1 mg/g dry weight (MATHIAS et al. 1960). BERKING (1986, 1987) detected homarine in several hydrozoans, where 1t regulates the colony morphology and prevents the metamorphosis in larvae. Detection of homarine in gastropods is rare. MC- CLINTOCK (1994) found the substance in the Antarctic Marseniopsis mollis (Caenogastropoda, Marseniidae). It is probably derived from epizoic bryozoans and hydro- zoans growing on the ascidian Cnemidocarpa verrucosa, the main food of M. mollis and serves as a feeding deter- rent against the seastar Odontaster validus. Since homarine was found in aeolids without cnidocysts, the question arouse, whether the occurence of homarine with a possible defensive function is more widespread in opisthobranchs.The current study focuses on the distribu- tion of homarine in several opisthobranch taxa, with a main focus on Cladobranchia, since this group 1s hardly investigated concerning its chemical defensive systems. Furthermore the localisation of this compound within the A 12% and, 0 SA el Á Saad Wek body of the dendronotoid Marionia blainvillea (Fig. 1B) was investigated in detail. The epidermal tissue of the mantle of M. blainvillea is densely packed with glands of unknown function (WAGELE et al. 2006) (Fig. 1C). There- fore a role of these glands in chemical defense was con- sidered likely. 2. MATERIAL AND METHODS Specimens of the opisthobranchs and of their food were collected worldwide during several field trips. Table 1 lists the investigated specimens, the collection details, and the means of extraction. The animals were fixated either in alcohol (ethanol or methanol) or by deep freezing. The supernatant alcohol was taken off and used for the preparation of a crude extract, or in the case of the deep frozen samples extraction with methanol was carried out. The solvent was evaporated under reduced pressure with a rotary evaporator in order to obtain a crude extract. The detection of homarine was carried out via 'H NMR spectroscopy on a Bruker Avance 300 DPX spectrometer, using D-O as solvent. Fig. 1. A. typical cnidosac (here Aeolidia papillosa). B. Marionia blainvillea C. Histological section of dorsal mantle epitheli- um of Marionia blainvillea. Note the single vacuoles in the cells staining homogenously dark violet. 184 Sven AFFELD et al.: Homarine in Opisthobranchia Table 1. Species investigated for homarine, with information on location, date of collection, extraction type and result. Food or- ganism or substrate 1s indicated in the last column. These information are taken from MCDONALD & NYBAKKEN (1997) if not stated otherwise. + homarine present; — no signal of homarine present in IH NMR and total amount of crude extract >100mg; ? insuffi- cient amount of crude extract (<10mg); Column 5 (Extraction): fluid in brackets indicate that only the alcohol for fixation was used for the preparation of a crude extract Higher ranking Taxon/Species Aeolidoidea Cratena pilata (Gould, 1870) Cuthona caerulea (Montagu, 1840) Cuthona gymnota (Couthouy, 1838) Flabellina affinis (Gmelin, 1791) Hermissenda crassicornis (Eschscholtz, 1831) Phestilla lugubris (Bergh, 1870) Phyllodesmium spec. Dendronotoidea Marionia blainvillea (Risso, 1818) Marionia blainvillea (Risso, 1818) Tritonia striata Haefelfinger, 1963 Tritonia manicata Deshayes, 1853 Doridoidea Dendrodoris grandiflora Rapp, 1827 Platydoris argo (Linné, 1767) Adalaria proxima (Alder & Hancock, 1854) Archidoris pseudoargus (Rapp, 1827) Cephalaspidea Scaphander lignarius (Linne, 1758) Aglaja tricolorata Renier, 1807 Anaspidea Aplysia spec. Phyllaplysia spec. Locatıon Number of Indiv. Fixation Date of collection Opisthobranchia Blanes, Spain 04.11.1993 Banyuls-sur-Mer, France 05.2006 Blanes, Spain 04.11.1993 Banyuls-sur-Mer, France 05.2006 5 Indiv. 24.03.1994 Blanes, Spain Deep frozen EtOH Deep frozen EtOH Deep frozen Data taken from KARUSO 1987 (after Klein 1986) Lizard Island, Australia 14.07.2006 5 Indiv. 09.2004 Blanes, Spain Blanes, Spain 10 Indiv. 05.2006 Blanes, Spain 15 Indiv. 04.-07.2004 11 Indiv. 03.-08.2004 Blanes, Spain Tierra de Malgrat, Spain 05.11.2004 Giglio, Italy 05.2005 07.1996 Bretagne, France Tierra de Malgrat, Spain 05.11.2004 Blanes, Spain 5 Indiv. 11.11.2004 Giglio, Italy 05.2005 La Coruna, Spain 08.03.2006 Banyuls-sur-Mer, France 09.1994 MeOH Deep frozen EtOH Deep frozen Deep frozen EtOH 96° o EtOH 70% EtOH 93% EtOH 96% EtOH 70% EtOH 70% EtOH 96% EtOH Extraction MeOH (EtOH) MeOH (EtOH) MeOH (MeOH) MeOH (EtOH) MeOH MeOH (EtOH) (EtOH) (EtOH) (EtOH) (EtOH) (EtOH) (EtOH) (EtOH) Homarine Food Tubularia spp., and other hydrozoans diverse hydrozoan species Tubularia spp., and other hydrozoans Eudendrium spp. Examined specimens found on Tubularia crocea, also on diverse hydrozoans Porites spp. Specialised on Xeni- idae. (pers. obs. HW) e.g. Eunicella singu- laris, also on other gorgonians Paralcyonium ele- gans (see Schmekel & Portmann 1982) Clavularia sp., Cornularia spp. (Alcynonacea) Sponges e.g. /rcinia fasciculata, Clathria toxystila, Spongia officinalis Demospongia Bryozoa Demospongia Predatory e.g. Polychaeta Predatory, other Opisthobranchia Higher ranking Taxon/Species Pleurobranchoidea Bathyberthella antarctica Willan & Bertsch, 1987 Tylodinoidea Umbraculum umbraculum (Lightfoot, 1786) Location Antarctica 61°43,30°S 59%12,40"W Bonner zoologische Beiträge 55 (2006) 185 Number of Indiv. Fixation Extraction Homarine Food Date of collection 27.11.1996 EtOH 70% (EtOH) unknown 08.09.1998 EtOH 80% (EtOH) - Demospongia Meteor Bank ST 486 (N-Atlantic) 29°45,7° N 28°22.9° W Other gastropods „Prosobranchia“ Littorina littorea Nordstrand, Germany 10 Indiv. EtOH 98% (EtOH) — Linne, 1758 22.08.2006 ve j Food of the examined Opisthobranchia Octocorallia Eunicella singularis Banyuls-sur-Mer, France 05.2006 EtOH 98% (EtOH) + (Esper, 1791) Xenniidae Lizard Island, Australia 14.07.2006 MeOH (MeOH) = Hydrozoa Tubularia crocea Blanes, Spain 04.11.1993 Deep frozen MeOH + Agassiz, 1862 From | g of the crude extract of one colony of the gor- gonian Eunicella singularis 4 mg of homarine were 1so- lated by RP-18 VLC and HPLC. The VLC was carried out using a 2.5 cm glass column filled 10 cm with Silica gel Polygoprep 60-50 C18. Elution was performed using a wa- ter/methanol gradient starting with pure water, yielding 13 fractions. The HPLC was performed on a Shimadzu system equipped with a Merk Hitachi Pump L-7100 and a Shi- madzu SPD-M6A Photodiode Array UV-VIS detector, us- ing a Knauer C¡g Eurospher-100 (5 um, 250x80 mm) column. A constant flow of 1 mL/sec was applied using Water/Methanol (90:10 V/V) as eluent. The five deep frozen specimens of Marionia blainvillea were dissected before extraction into mantle, foot, diges- tive gland and the rest of the viscera, in order to examine the distribution of homarine within the body of the ani- mal. For histological investigation, one specimen each of the dendronotoid Mariona blainvillea and the aeolid Cuthona caerulea have been preserved in formaldehyde/seawater. After dehydration in alcohol, they were embedded in hy- droxyethylmethacrylate for serial sectioning and sections (2.5 um) were stained with toluidine blue. Investigation was performed under a light microscope. 3. RESULTS The survey of several members of the Opisthobranchia re- vealed that the natural compound homarine 1s present in all examined samples of Cladobranchia, apart from a new species of Phyllodesmium from Lizard Island. This com- pound was also not found in other members of the Opisthobranchia, except for Aglaja tricolorata. Results of this survey are listed in Table 1. The low field shifted 'H NMR signals of homarine 6 7.8 — 8.9 ppm (see Fig. 2) are very characteristic for this com- pound, and can easily be detected in 'H NMR spectra of crude extracts. In all cases they did not overlap with any other resonance signals. The detection of these resonance signals is a clear indication for the presence of homarine, however only if the concentration of the compound is suf- ficient, i.e. traces will not be detected. For specific localisation of the compound different body parts of M. blainvillea were extracted separately. 'H NMR spectra of the extracts show that homarine is present throughout the body (see Fig. 3 A to D). Histological investigation revealed that the epidermis of M. blainvillea contains many epithelial glandular cells with one large vacuole filling the whole cell. The vacuole contains a substance that is stained homogeneously dark violett with toluidine blue (Fig. 1C). These glandular cells are evenly distributed over the dorsal and lateral notum. 186 Sven AFFELD et al.: Homarine in Opisthobranchia | | | O | j | | N Ä | |! CH, O IN I | AA y \ \ | \ Aa ar) ran Mal po j x 4.000 88 86 84 8.2 8.0 7.8 | | | N hh ie | N | | If Mo . A SS Ñ PPM 86 84 82 80 78 76 74 72 70 68 66 64 62 60 58 56 54 52 50 48 46 44 Fig. 2. 'H NMR spectrum of purified homarine in methanol-d,. 300 MHz, and chemical structure of homarine. The epithelium of Cuthona caerulea showed very few glandular cells, these were usually filled with dark stain- ing granular substances. 4. DISCUSSION Secondary metabolites clearly seem to play a role in the biotic interactions with potential predators throughout all systematic groups of Opisthobranchia. These chemicals are often taken up from the food while de novo synthesis is rare (review see CIMINO et al. 2004). Only a few field studies on the deterrent effect of the ob- served compounds have been carried out (THOMPSON 1960; AvILA & PAUL 1997; SLATTERY et al. 1998; JOHN- SON & WILLOWS 1999; MARIN et al. 1999; AvILA et al. 2000; BECERRO et al. 2001; IKEN et al. 2002; ROGERS et al. 2002; PENNEY 2004). While even a few studies focus on parasites on opisthobranchs (ARNAUD 1978; Ho 1981: CAREFOOT 1987; JENSEN 1987; Huys 2001; SCHRÓDL 2002, 2003), the effect of secondary metabolites on pathogens is hardly investigated. TEEYAPANT et al. (1993a, b) studied the uptake of secondary metabolites in 7y/o- dina perversa (Tylodinoidea) from its food sponge Aplvsina (Verongia) aerophoba and the antimicrobial ac- tivity of the sponge metabolites. However they only found antimicrobial activity in compounds that were not pres- ent in Tylodina perversa. Incorporation of homarine is shown here for slugs feed- ing on Hydrozoa as well as on the gorgonian octocoral £. singularis. The examined Hydrozoa contained homarine, which is in accord with its role as a morphogen, suppos- edly in all hydrozoans (BERKING 1986, 1987). The exam- ined gorgonian £. singularis also contained homarine as described for other gorgonians where it serves as an an- tifouling agent (TARGETT et al. 1983). Uptake of homa- rine can also occur by predation on opisthobranchs con- taining homarine. According to RUDMAN (1972), members of the genus Aglaja are active predators, feeding on vagile prey. Whereas some aglajid species, like A. cylindrica, spe- cialize on polychaetes and nemertines, others (e.g. A. au- reopunctata) mainly focus on opisthobranchs. It seems likely that Aglaja tricolorata obtained the homarine by feeding on other opisthobranchs. Several of the investigated species belong to the Aeoli- doidea with functional cnidocysts incorporated, which are a good means of defense against predatory fishes (FRICK 2003) but certainly useless against crustaceans and prob- ably without effect against seastars. Both latter groups are potential predators on opisthobranchs, though only few thorough investigations on predation have been carried out (review see WAGELE et al. 2006). The efficiency of homa- rine as a feeding deterrent is shown e.g. in Marseniopsis mollis (MCCLINTOCK 1994). But similar studies have nev- er been performed for opisthobranchs and further inves- tigations are needed to clarify the role of this compound in cladobranchs. The presence of a further defensive sys- tem, like homarine, would provide additional protection, e.g. for the cnidocyst containing Cuthona caerulea. In species without cnidocysts, e.g. Marionia blainvillea, homarine might even represent the sole defensive strate- gy. Bonner zoologische Beitráge 55 (2006) 187 x 4.000 == Sy Ty 8.8 as 8.4 8.2 80 T T i r 1 T y i - — mi zu PPM 90 8.0 70 60 50 40 3.0 20 vo Fig. 3. 'H NMR spectra of crude extracts of dissected Marionia blainvillea in methanol-dy. 300 MHz. A. Mantle, B. Foot, C. Digestive gland, D. Viscera without digestive gland. 188 Sven AFFELD et al.: Homarine in Opisthobranchia Homarine could also serve as an antibacterial agent in the mucus layer of the slugs, independent of its supposed func- tion as a feeding deterrent, since any kind of mucus is a perfect environment for bacteria or fungi. Therefore, or- ganisms with a mucus layer can adopt two ways of keep- ing their body surface clean. They can either produce big amounts of mucus that wash off continuously or add a sub- stance that inhibits growth of microorganisms. The for- mer strategy is known, e.g. from certain soft corals (DUCK- LOW & MITCHELL 1979; DUERDEN 1986), the latter could be the case in the Aeolidoidea and Dendronotoidea. WAGELE et al. (2006) noted the presence of homogenous- ly staining violet glands as a characteristic of Dendrono- toidea. In their survey on glandular structures in Opistho- branchia, they observed these glands for nine (out of 17 investigated) members of the Dendronotoidea. These glands are absent in any members of the Aeolidoidea and “Arminoidea”. We confirmed the presence of these par- ticular glands in Marionia blainvillea and their absence in Cuthona caerulea. The uniform distribution of homa- rine in the different body parts of M. blainvillea suggests that no bioaccumulation in the glandular epithelium takes place in this case. Homarine is a very small and very po- lar molecule, therefore it is likely that it is taken up pas- sively in the digestive gland or even in the whole diges- tive system. Quantitative analyses still should be carried out to support this supposition. 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Authors’ adresses: Sven AFFELD, Institut für Phar- mazeutische Biologie, Universität Bonn, Nussallee 6, 53121 Bonn, Germany, E-mail: sven.affeld@uni-bonn.de; Heike WÄGELE (corresponding author) Institut für Evo- lutionsbiologie und Ökologie, Universität Bonn, An der Immenburg 1, 53121 Bonn, Germany, and Zoologisches Forschungsmuseum Alexander Koenig, Bonn, Germany, E-mail: hwaegele@evolution.uni-bonn.de; Conxita AvI- LA, Dept. of Animal Biology (Invertebrates), Facultat de Biologia, Universitat de Barcelona, Av. Diagonal, 645 08028 Barcelona, Spain, E-mail: conxita.avila@ub.edu; Stefan KEHRAUS, Institut für Pharmazeutische Biologie, Universität Bonn, Nussallee 6, 53121 Bonn, Germany; Gabriele König, Institut für Pharmazeutische Biologie, Universität Bonn, Nussallee 6, 53121 Bonn, Germany, E-mail: g.koenig@uni-bonn.de. Bonner zoologische Beitráge Band 55 (2006) Heft 3/4 Seiten 191-202 Bonn, November 2007 Utility of H3-Genesequences for phylogenetic reconstruction — a case study of heterobranch Gastropoda —* Angela DNAPOLI, Ceyhun TAMER)), Susanne FRANSSEN)), Lisha NADUVILEZHATH!) & Annette KLUSSMANN-KOLB!) Department of Ecology, Evolution and Diversity — Phylogeny and Systematics, J. W. Goethe-University, Frankfurt am Main, Germany *Paper presented to the 2nd International Workshop on Opisthobranchia, ZFMK, Bonn, Germany, September 20th to 22nd, 2006 Abstract. In the present study we assessed the utility of H3-Genesequences for phylogenetic reconstruction of the He- terobranchia (Mollusca, Gastropoda). Therefore histone H3 data were collected for 49 species including most of the ma- jor groups. The sequence alignment provided a total of 246 sites of which 105 were variable and 96 parsimony informa- tive. Twenty-four (of 82) first base positions were variable as were 78 of the third base positions but only 3 of the se- cond base positions. H3 analyses showed a high codon usage bias. The consistency index was low (0,210) and a substitution saturation was observed in the 34 codon position. The alignment with the translation of the H3 DNA sequences to amino-acid sequences had no sites that were parsimony-informative within the Heterobranchia. Phylogenetic trees were reconstructed using maximum parsimony, maximum likelihood and Bayesian methodologies. Nodilittorina unifasciata was used as outgroup. The resolution of the deeper nodes was limited in this molecular study. The data themselves were not sufficient to clar- ify phylogenetic relationships within Heterobranchia. Neither the monophyly of the Euthyneura nor a step-by-step evo- lution by the “basal” groups was supported. A conclusion about the monophyly of Opisthobranchia and Pulmonata could not be extracted from our data because we did not have any resolution at this point. We believe histone H3 alone provides no new marker for studying deep molecular evolution of the Heterobranchia due to the high grade of conservation and the low phylogenetic signal. Surprisingly there was a good resolution on the genera level. Analyses conducted with maximum parsimony and Bayesian inference (using all data) recovered all (or nearly all) genera mostly with statistically significantly supported nodes. Fur- ther studies focusing on the possible utility of histone H3 for the resolution of recent splits will be necessary. Keywords. Heterobranchia, Opisthobranchia, histone H3, molecular phylogeny. 1. INTRODUCTION Many questions regarding gastropod phylogeny have not yet been answered such as the molecular confirmation of the Heterobranchia concept based on morphological stud- ies from HASZPRUNAR (1985, 1988). This taxon contains the Pentaganglionata HASZPRUNAR, 1985 also known as Euthyneura SPENGEL, 1881 (with the Opisthobranchia and Pulmonata) and several mostly little known “basal” groups (e.g. Valvatoidea, Omalogyroidea, Architectonicoidea, Rissoelloidea and Pyramidelloidea) which present a step- by-step evolution towards the euthyneuran level of organ- isation (HASZPRUNAR 1988). The hyperstrophy of the pro- toconch is the most important autapomorphous character of the Heterobranchia. The Euthyneura are characterised by the presence of two additional (so-called parietal) gan- glia. However, the monophyly of the Euthyneura has not been clarified by molecular studies, yet. In some studies they are recovered monophyletic (COLGAN et al. 2000, 2003; KNUDSEN et al. 2006) in others not (THOLLESSON 1999). The Pulmonata and Opisthobranchia can be sepa- rated by characters respective of the nervous system (pres- ence of a procerebrum and cerebral bodies in pulmonates and presence of a rhinophoral nerve in Opisthobranchia and Pyramidelloidea). However the molecular confirma- tion regarding the monophyly of the Opisthobranchia (VONNEMANN et al. 2005; GRANDE et al. 2004a) and the Pulmonata (TILLIER et al. 1996, DAYRAT et al. 2001) is still a matter of debate. There is no comprehensive investiga- tion concerning the “basal” groups. Only a few represen- tative taxa (e.g. Valvatoidea — Cornirostra pellucida, Ar- chitectonicoidea — Philippea lutea, Pyramidelloidea — Pyramidella dolabrata) have been included in current mo- lecular studies (COLGAN et al. 2000; GRANDE et al. 2004a, 2004b). 192 Angela DINAPOLI et al.: Utility of H3-Gene in Heterobranchia In recent years molecular systematic analyses in gas- tropods have utilised a variety of genetic markers, e.g. nu- clear 28S ribosomal RNA and/or 18S ribosomal RNA or mitochondrial 16S ribosomal RNA and/or cytochrome ox- idase subunit I (TILLIER et al. 1994, 1996; DAYRAT et al. 2001; VONNEMANN et al. 2005; THOLLESSON et al. 1999; REMIGIO & HEBERT 2003). Nevertheless, new genetic markers are needed for the resolution of certain phyloge- netic relationships (especially regarding deeper nodes). Partial fragments of the gene coding for the extremely con- servative H3 protein (MAXSON et al. 1983) were first used to clarify arthropod molecular evolution (COLGAN et al. 1998) and later polychaete (BROWN et al. 1999), gastro- pod (COLGAN et al. 2000, 2003), polyplacophoran (OKUSU et al. 2003), cephalopod (LINDGREN et al. 2004) and hexa- pod (KJER et al. 2006) phylogeny. All studies used a com- bined dataset in their approaches. In their study of gas- tropod phylogeny, Colgan et al. (2000) did not find a monophyletic Heterobranchia while within the Euthyneu- ra, the Opisthobranchia are paraphyletic with respect to the pulmonates. Very similar phylogenetic relationships were shown in COLGAN et al. (2003). The Heterobranchia as well as the Opisthobranchia and Pulmonata are rarely recovered as monophyletic in these studies. In the present study we wanted to test the utility of H3 gene sequences for phylogenetic reconstruction within the Heterobranchia (focusing primarily on the Opistho- branchia). We were especially interested in testing whether H3 is suitable to resolve deeper nodes within het- erobranch phylogeny. Therefore, partial histone H3 data were collected for 49 species including most of the ma- jor groups (Euthyneura with Opisthobranchia and Pul- monata and “basal” groups with Valvatoidea, Architecton- icoidea, Omalogyroidea, Rissoelloidea and Pyramidel- loidea). 2. MATERIALS AND METHODS 2.1. Specimens and DNA extraction The studied taxa and the accession numbers are listed in Table 1. Twenty of the 49 sequences are taken from Gen- Bank. Opisthobranchia are represented by 26 species (in- cluding 11 suborders). Nodilittorina unifasciata (Caenogastropoda Cox, 1960) was used as an outgroup. DNA was extracted from ethanol-preserved individuals us- ing the DNeasy Tissue Kit from Qiagen (Hilden, Ger- many). Table 1 . Taxonomic positions and collecting locations of the sampled taxa. Accession numbers of sequences included in the ana- lyses (ZSM = Zoologische Staatssammlung); published sequences taken from GenBank are marked with an asterisk. Major Taxon Species Caenogastropoda Littorinoidea Littorinidae Conidae Campanilidae Opisthobranchia Nudibranchia Tethydidae Discodorididae Arminidae Pleurobranchoidea Pleurobranchidae Tylodinoidea Umbraculidae Cephalaspidea Scaphandridae Philinidae Gastropteridae Anaspidea Akeridae Aplystidae Aplysiidae Aplysiidae Thecosomata Cavoliniidae Creseidae Gymnosomata Pneumodermatidae Nodilittorina unifasciata (Gray, 1826) Conus miles Linnaeus, 1758 Campanile symbolicum Iredale, 1917 Tethys fimbria Linne, 1767 Discodoris atromaculata (Bergh, 1880) Armina neapolitana (Delle Chiaje, 1824) Pleurobranchaea meckeli Leue, 1813 Umbraculum umbraculum (Lightfoot, 1786) Scaphander lignarius (Linne, 1758) Philine aperta (Linnaeus, 1767) Gastropteron meckeli Kosse, 1813 Akera bullata Müller, 1776 Aplvsia californica Cooper, 1863 Aplysia cf. juliana Quoy & Gaimard, 1832 Bursatella leachii de Blainville, 1817 Clio pyramidata Linné, 1767 Creseis sp. Pneumoderma cf. atlantica (Oken, 1815) Locality GenBank Accession Number Genbank AF033705* Genbank AF033684* Genbank AF033683* Blanes, Spain EF133468 Genbank DQ280013* Banyuls-sur-Mer, France EF133469 Blanes, Spain EF133470 Atlantic Ocean, Meteor Bank EF133471 Blanes, Spain EF 133472 Genbank DQ093508* Blanes, Spain EF 133473 Kattegat, Denmark EF133474 Miami, USA EF133475 Genbank AF033675* Dingo Beach, Australia EF133476 Canary Islands; Spain EF 133477 Genbank DQ280012* USA, Atlantic EF133478 Bonner zoologische Beiträge 55 (2006) 193 Major Taxon Species Sacoglossa Placobranchidae Elysia tímida (Risso, 1818) Placobranchidae Elysia pusilla (Bergh, 1872) Placobranchidae Elysia crispata Morch, 1863 Placobranchidae Elysia viridis (Montagu, 1804) Cylindrobullidae Cylindrobulla beauii Fischer, 1857 Acochlidia Hedylopsidae Hedylopsis spiculifera (Kowalewsky, 1901) Microhedylidae Unela glandulifera (Kowalewsky, 1901) Architectibranchia Hydatinidae Micromelo undatus (Bruguiere, 1792) Acteonoidea Acteonidae Pupa solidula (Linné, 1758) Acteonidae Rictaxis punctocaelatus (Carpenter, 1864) Bullinidae Bullina lineata (Gray, 1825) Pulmonata Systellommatophora Onchidiidae Onchidium sp. Onchidiidae Onchidella floridana (Dall, 1885) Onchidiidae Onchidella sp. Stylommatophora Charopidae Hedleyoconcha delta (Pfeiffer, 1857) Siphonariidae Siphonaria serrata (Fischer, 1807) Siphonariidae Siphonaria concinna Sowerby,1824 Siphonariidae Siphonaria zelandica (Quoy & Gaimard, 1832) Amphibolidae Salinator solida (Schacko, 1878) Eupulmonata Ellobiidae Ophicardelus ornatus (Ferussac, 1821) “basal” Heterobranchia/Triganglionata Pyramidelloidea Pyramidellidae Turbonilla lactea (Linné, 1758) Pyramidellidae Turbonilla sp. Architectonicoidea Architectonicidae Heliacus variegatus (Gmelin, 1791) Architectonicidae Philippea lutea (Lamarck, 1822) Valvatoidea Cornirostridae Cornirostra pellucida (Laseron, 1954) Orbitestellidae Orbitestella vera Powell, 1940 Orbitestellidae Orbitestella sp. Omalogyroidea Omalogyridae Omalogyra burdwoodiana Strebel, 1908 Rissoelloidea Rissoelidae Rissoella elongatospira Ponder, 1966 Rissoelidae Rissoella micra Finlay, 1924 Rissoelidae Rissoella cystophora Finlay, 1924 2.2. DNA amplification and sequencing The following degenerated primers were used: H3-F: 5”- ATG GCT CGT ACC AAG CAG AC(ACG) GC-3” and H3-R: 5”-ATA TCC TT(AG) GGC AT(AG) AT(AG) GTG AC-3” (Colgan et al. 1998) and produced a 246 bp prod- uct. The PCR profile was as follows: 95 °C for 5 min, fol- lowed by 35 cycles of 30 s at 95 °C, 25 sat 52 °C, 45 s at 72 °C and a final extension at 72 °C for 5 min and Taq Polymerase, recombinant from Invitrogen (Karlsruhe, Germany) was used. All products were purified using the QIA quick Gel Extraction Kit from Qiagen (Hilden, Ger- many) and sequenced in both directions with a CEQ 2000 Beckmann Coulter using the CEQ DTCS Quick Start Kit Locality GenBank Accession Number Blanes, Spain EF133479 Genbank DQ534792* Genbank DQ534790* Genbank DQ534790* Florida, USA EF133480 Rovinj, Croatia EF133481 Rovinj, Croatia EF133482 Genbank DQ093513* Dingo Beach, Australia EF133483 Cayucos, Californica, USA EF133484 Genbank AF033680* Genbank AF033706* Florida, USA EF 133485 Genbank DQ093511* Genbank AF033693* South Africa EF133486 South Africa EF133487 Genbank AF033713* Genbank AF033712* Genbank AF033707* Roscoff, France EF133488 Wellington, New Zealand EF 133489 Tropical aquarium (ZSM 20012193) EF133490 Genbank AF033708* Genbank AF033685* Wellington, New Zealand EF561623 Leigh, New Zealand EF561624 Antarctic (ZSM Mol-20021228) EF133491 Wellington, New Zealand EF561622 Wellington, New Zealand EF561620 Wellington, New Zealand EFS61621 (Krefeld, Germany). First we sequenced only one frag- ment per specimen. Moreover, to avoid mistakes accord- ing to the conservative character of the H3 gene, we se- quenced a random sample of 5 species a second time whereby no varieties could be detected. 2.3. Sequence alignment Sequences were aligned manually using the software pack- age BioEdit version 7.0.5 (HALL 1999). The H3 DNA se- quences were translated into the amino acid sequences in GeneDoc version 2.6.002 (NICHOLAS & NICHOLAS 1997). The alignment is available from the authors upon request. 194 Angela DINAPOLI et al.: Utility of H3-Gene in Heterobranchia 2.4. Statistical tests Codon usage statistics were calculated using GCUA ver- sion 1.2 (MCINERNEY 1997). The purpose of this function is to calculate the Number (N) of times a particular codon is observed in an alignment and also to calculate the Rel- ative Synonymous Codon Usage (RSCU) values for the dataset. RSCU values define the number of times a par- ticular codon is observed relative to the number of times that the codon would be observed in the absence of any codon usage bias. Without any codon usage bias, the RSCU value would be 1.00. A codon that is used less fre- quently than expected will have a value of less than 1.00 and a codon that is used more frequently than expected will have a volume of more than 1.00 (MCINERNEY 1997). The degree of bias (Yy2/n) which is the sum of the 72 val- ues for the individual amino acids divided by the total number of inferred residues (n) for the combination of da- ta from all species (SHIELDS et al. 1988) was determined. The substitution saturation was calculated for all 3 codon positions using the method developed by XıA et al. (2003) implemented in the software package DAMBE version 4.2.13 (XIa & XIE 2001). 2.5. Phylogenetic reconstruction Appropriate models for the analyses were selected after running Modeltest version 3.4 (POSADA & CRANDALL 1998) and using the Akaike information criterion (AIC) (see tab. 2). The following analyses were conducted using PAUP* ver- sion 4.0 b10 (SWOFFORD, 2002) (settings: heuristic search strategy; tbr; gaps were treated as fifth bases): a) Maxi- mum parsimony for all data and b) Maximum likelihood for all data. Table 2. Information on used models. Bootstrapping (FELSENSTEIN 1985) was performed for maximum parsimony with 1000 replicates and for maxi- mum likelihood with 100 replicates. The following analyses were conducted using MrBayes version 3.1.2 (RONQUIST & HUELSENBECK 2003): Bayesian inference: a) all data (with one model for all three codon positions), b) all data (with codon specific models) and e) only codon position one and two (third codon position excluded; with one model for codon position one and two). For Bayesian inference a Metropolis Chain Monte Carlo analysis with four chaims and 1 000 000 generations was performed with the first 1000 trees ignored as burn-in. 3. RESULTS 3.1. Statistical tests The sequences provided a total of 246 sites of which 105 were variable and 96 parsimony informative. Twenty-four (of 82) first base positions were variable as were 78 of the third base positions but only 3 of the second base posi- tions. Insertion/deletion events (indels) were not observed in any of the groups. The amino acid alignment had no sites that were parsimony-informative within the Hetero- branchia. H3 analyses showed a high codon usage bias (Tab. 3). The bias was principally against the use of A and U in the third codon position. ¥ tests were performed for all amino acids and revealed that the null hypothesis which is the expect- ed equal usage of the codons can be rejected for all amino acids with a significance level of 0,001 excepting histi- dine (p<0,05). For aspartic acid the null hypothesis can not be rejected (p=0,21). The degree of bias (Sy?/n) showed a high value of 0,617. Proportion of invariable sites Codon-Position Model Gamma distribution shape parameter [st Position GTR+I a=equal 2nd Position TVMef+l a=equal 3d Position TVM+G a=0.8835 Ist and 214 Position GTR+H+G a=0.9167 Ist 29d and 3'd Position GTR+H+G a=1.0265 Pinvar=0.5692 Pinvar=0.8647 Pinvar=equal Pinvar=0.6909 Pinvar=0.5408 Ñ | Bonner zoologische Beitráge 55 (2006) 195 Table 3. Codon Usage Bias. N = number of times a particular codon is observed in a dataset (alignment). RSCU values = num- ber of times a particular codon is observed, relative to the number of times that the codon would be observed in the absence of any codon usage bias. Amino acids (AA) are indicated by the three letter abbreviations. AA Codon N RSCU Phe UUU 6 (0.08) Phe UUC 142 (1.92) Leu UUA (0.05) Leu UUG 43 (0.66) Tyr UAU 2 (0.04) Tyr UAC 96 (1.96) Ter UAA 0 (0.00) Ter UAG 0 (0.00) Ter UGA 0 (0.00) Leu CUU 48 (0.74) Leu EUC 81 (1.24) Leu CUA 2 (0.03) Leu CUG 214 (3.28) His CAU 31 (1.27) His CAC 18 (0.73) Gln CAA 41 (0.28) Gln CAG 252 (1.72) Ile AUU 21 (0.43) Ile AUC 126 (2.57) Ile AUA 0 (0.00) Asn AAU 0 (0.00) Asn AAC 0 (0.00) Lys AAA 135 (0.61) Lys AAG 306 (1.39) Val GUU 9 (0.18) Val GUC 89 (1.82) Val GUA 4 (0.08) Val GUG 94 (1.92) Asp GAU 38 (0.78) Asp GAC 60 (1.22) Glu GAA 69 (0.70) (1.30) Glu GAG 128 The consistency index in maximum parsimony analyses was low (0,210) as well as the retention index (0.444). Us- ing the method developed by XIA et al. (2003) a substi- tution saturation was observed in the third codon position (1,, 0,543 > I... 0,298). To support this observation the en- tropy for each position in the alignment was calculated us- ing BioEdit Version 7.0.5.2. Codon position one showed an average entropy value of 0,09 whereas the value for the second codon position was 0,009 and 0,69 for the third codon position. If the nucleotides occur more or less equal- ly at a certain position within the alignment the entropy is highest with the value of 1,36. AA Codon N RSCU Ser UCU 45 (1.38) Ser UCC 34 (1.04) Ser UCA 13 (0.40) Ser UCG 8 (0.24) Cys UGU 0 (0.00) Cys UGC 0 (0.00) Trp UGG 0 (1.00) Pro CCU 83 (1.36) Pro ECC 107 (1.75) Pro CCA 42 (0.69) Pro CCG 13 (0.21) Arg CGU 233 (2.59) Arg CGC 127 (1.41) Arg CGA 18 (0.20) Arg CGG 9 (0.10) Thr ACU 3 (0.87) Thr ACC 146 (2.38) Thr ACA 45 (0.73) Thr ACG 1 (0.02) Ser AGU 8 (0.24) Ser AGC 88 (2.69) Arg AGA 74 (0.82) Arg AGG 78 (0.87) Ala GCU 201 (1.49) Ala GCC 279 (2.07) Ala GCA 47 (0.35) Ala GCG 12 (0.09) Gly GGU 32 (0.87) Gly GGC 38 (1.03) Gly GGA 72 (1.96) Gly GGG 5 (0.14) Met AUG 49 (1.00) 3.2. Phylogenetic analyses The maximum parsimony 50 % majority-rule consensus tree (Fig. 1) showed all genera (Aplysia, Elysia, Onchidel- la, Siphonaria, Orbitestella, Turbonilla and Rissoella) re- covered as monophyletic. However, some of the bootstrap supports were low and there was no bootstrap support for a monophyletic Rissoella. Beyond the genera level, only Architectonicoidea were detected as monophyletic. All other nodes lacked support. The genera Orbitestella, Turbonilla, Onchidella and Aplvsia and again, the Architectonoicoidea were found to be monophyletic in the maximum likelihood analyses. The 196 Angela DINAPOLI et al.: Utility of H3-Gene in Heterobranchia Fig. 1. 50 % majority-rule consensus tree of maximum parsimony of 14 most parsimonious trees based on histone H3 data set (nucleotides), number of parsimony informative characters = 96, consistency index (CI) = 0,210. Nodilittorina unifasciata Campanile symbolicum Conus miles Clio pyramidata Omalogyra burdwoodiana Akera bullata 100 Bursatella leachii Unela glandulifera Pneumoderma cf. atlantica Cylindrobulla beauii Aplysia californica 78 Aplysia cf. juliana Philine aperta Creseis sp. Cornirostra pellucida Elysia timida Elysia pusilla Elysia crispata 836 Flysia viridis Gastropteron meckeli Orbitestella vera Orbitestella sp. Tethys fimbria Heliacus variegatus Philippea lutea Salinator solida Pleurobranchaea meckeli Scaphander lignarius Umbraculum umbraculum Hedleyoconcha delta Onchidium sp. Hedylopsis spiculifera Pupa solidula Rictaxis punctocaelatus Micromelo undatus Bullina lineata Turbonilla lactea %E= Turbonilla sp. Ophicardelus ornatus Armina neapolitana Discodoris atromaculata Rissoella elongatospira Rissoella micra 100 Rissoella cystophora Onchidella floridana 92h Onchidella sp. Siphonaria concinna 52 Siphonaria serrata 83k= Siphonaria zelandica Bonner zoologische Beiträge 55 (2006) 197 Fig. 2. 50 % majority rule consensus Bayesian inference cladogram for the histone H3 dataset (based on nucleotides); Bayesian posterior probabilities provided at the branches. 0.97 0.88 0.77 0.99 0.56 0.97 0.73 0.67 0.97 0.84 0.88 0.64 1.00 0.98 0.96 0.70 0.99 0.90 0.62 1.00 0.99 1.00 0.62 0.58 0.58 Nodilittorina unifasciata Conus miles Omalogyra burdwoodiana Campanile symbolicum Creseis sp. Unela glandulifera Philine aperta Bursatella leachii Aplysia californica Aplysia cf. Juliana Clio pyramidata Heliacus variegatus Philippea lutea Akera bullata Pneumoderma cf. atlantica Cylindrobulla beauil Cornirostra pellucida Pupa solidula Rictaxis punctocaelatus Micromelo undatus Bullina lineata Turbonilla lactea Turbonilla sp. Gastropteron meckeli Orbitestella vera Orbitestella sp. Elysia pusilla Elysia timida Elysia crispata Elysia viridis Tethys fimbria Salinator solida Ophicardelus ornatus Hedleyoconcha delta Onchidium sp. Hedylopsis spiculifera Scaphander lignarius Umbraculum umbraculum Pleurobranchaea meckeli Discodoris atromaculata Armina neapolitana Onchidella floridana Onchidella sp. Rissoella elongatospira Rissoella micra Rissoella cystophora Siphonaria concinna Siphonaria serrata Siphonaria zelandica 198 Angela DINAPOLI et al.: Utility of H3-Gene in Heterobranchia Fig. 3. 50 % majority rule consensus Bayesian inference phylogram for the histone H3 dataset (based on nucleotides, 3d codon position excluded); Bayesian posterior probabilities provided at the branches. Nodilittorina unifasciata Campanile symbolicum Omalogyra burdwoodiana 0.57 Conus miles Discodoris atromaculata Cornirostra pellucida Micromelo undatus Salinator solida Ophicardelus ornatus Siphonaria zelandica Siphonaria concinna Siphonaria serrata Hedleyoconcha delta Onchidella sp. Onchidella floridana Onchidium sp. Bullina lineata Rictaxis punctocaelatus Pupa solidula Unela glandulifera Hedylopsis spiculifera Cylindrobulla beauii Elysia viridis Elysia pusilla Creseis Sp. Clio pyramidata 0.9 Aplysia cf. juliana Gastropteron meckeli Scaphander lignarius Umbraculum umbraculum Pleurobranchaea meckeli Elysia timida 0.62 Elysia crispata Tethys fimbria 0.70 Armina neapolitana Philine aperta 0.72 Pneumoderma cf. atlantica Heliacus variegatus 0.78 Philippea lutea Orbitestella vera 0.81 Orbitestella sp. Turbonilla lactea 1.00 Turbonilla sp. Akera bullata Bursatella leachii Aplysia californica 0.53 Rissoella elongatospira 0.99 Rissoella micra 0.99 Rissoella cystophora 0.1 Bonner zoologische Beitráge 55 (2006) 199 basis of the 50 % majorıty-rule bootstrap tree (tree not shown) resembled a comb. There was no resolution of the deep nodes. The 50 % majority-rule consensus Bayesian inference cladogram (with one model for all three codon positions) (Fig. 2) also recovered all genera as monophyletic. Only the Bayesian posterior probability for Siphonaria was low (0.58) because only values above 0.95 are statistically sig- nificant. Beside the genera level Architectonicoidea were monophyletic and the Caenogastropoda formed a clade to- gether with Omalogyra. All other nodes had no statisti- cally significant support. The 50 % majority-rule consensus Bayesian inference cladogram (with codon specific models) (tree not shown) was quite similar to Figure. 2. All genera except Rissoel- la and Architectonicoidea were found to be monophyle- tic. The remaining nodes were supported by Bayesian pos- terior probabilities below 0.95. The 50 % majority rule consensus Bayesian inference phy- logram (with 314 codon position excluded and with one model for codon position one and two) (Fig. 3) recovered only the genera Rissoella and Turbonilla as monophylet- ic while the Caenogastropoda together with Omalogvra were grouped separately from the rest of the taxa. All oth- er nodes had no statistically significant support. 4. DISCUSSION Molecular investigations of deep-level relationships with- in the Gastropoda have been made difficult due to a lack of slowly evolving genes. Hence, a number of different markers have been utilized to solve this problem. Analy- ses of nuclear genes like the 28S ribosomal RNA and/or the 18S ribosomal RNA have provided a number of im- portant insights into gastropod relationships at several lev- els (TILLIER et al. 1994, 1996; DAYRAT et al. 2001; Von- NEMANN et al. 2005). The same applies to mitochondrial genes like the 16s ribosomal RNA (THOLLESSON et al. 1999) or the cytochrome oxidase subunit I (REMIGIO & HEBERT 2003). COLGAN et al. (2000, 2003) used the his- tone H3 protein in combination with other genes to clar- ify gastropod molecular evolution. However, many aspects of gastropod phylogeny remain unclear such as the mo- lecular confirmation of the Heterobranchia concept based on morphological studies from HASZPRUNAR (1985, 1988). At the moment there is no comprehensive molec- ular study of heterobranch phylogeny especially one in- cluding the “basal” taxa (e. g. Pyramidelloidea, Architec- tonicoidea, Valvatoidea, Omalogyroidea and Rissoel- loidea). In this study we wanted to present a primary molecular insight into heterobranch phylogeny while simultaneous- ly testing the utility of the gene coding for the highly con- served protein histone H3 for resolving the deeper nodes within this taxon. Unfortunately, the present study did not provide a robust phylogenetic hypothesis for the relationships among dif- ferent lineages of Heterobranchia based on H3-Genese- quences. Neither the monophyly of the Euthyneura nor a step-by-step evolution by the “basal” groups was support- ed. A conclusion about the monophyly of Opisthobranchia and Pulmonata could not be extracted from our data be- cause we did not have any resolution at this point. The first to investigate the value of histone H3 were CoL- GAN et al. (1998). They wanted to combine small nuclear ribonucleic acid U2 data and histone H3 to investigate arthropod molecular evolution. However, partitioned da- ta for H3 and U2 were incongruent according to Incon- gruence Length Difference tests. Using H3 data only, anomalous nodes appeared in their phylogenies while some possessed decay indices of 1. Therefore, their data were not sufficient to clarify relationships within major arthropod groups. BROWN et al. (1999) investigated the DNA sequence da- ta of 34 Polychaeta species for partial histone H3, U2 snR- NA and two segments of 28S rDNA (Dl and D9-10 ex- pansion regions). When using H3 only, BROWN et al. (1999) found a lack of concordance with morphological results and argued that the inclusion of all H3 data is im- appropriate for the phylogenetic levels under investiga- tions. COLGAN et al. (2000) and later COLGAN et al. (2003) used partial histone H3 (327bp) for the investigation of gastro- pod phylogeny. In COLGAN et al. (2000), where the authors used 36 sequences of histone H3 only, using the chiton Ischnochiton australis as an outgroup, no clades were re- tained in the bootstrap analyses. H3 with the third codon position excluded, retained only the higher Vetigastropo- da (bootstrap support = 68 %). In COLGAN et al. (2003) in which H3 alone was used to recover phylogenetic relationships within Gastropoda, on- ly the clade of the Patellogastropoda with a support of 52 % was recovered. When the third codon position was ex- cluded, none of the expected groups were recovered. Oxuso et al. (2003) were the first to apply DNA sequence data to reconstruct the phylogeny of the molluscan class Polyplacophora. Their use of 59 sequences of histone H3 resolved deeper nodes than the mitochondrial genes did while the strict consensus tree nested the two Gastropo- 200 Angela DINAPOLI et al.: Utility of H3-Gene in Heterobranchia da Viviparus georgianus and Siphonaria pectinata with- in Polyplacophora. A combined approach to the phylogeny of Cephalopoda (Mollusca) using 18S rRNA, 28S rRNA, histone H3 and COI underscored the aim of the study presented by LIND- GREN et al. (2004). The strict consensus tree for the over- all optimal parameter set for 66 sequences of histone H3 alone did not show monophyly for any classes investigat- ea: KJER et al. (2006) investigated the molecular phylogeny of Hexapoda (supermatrix approach with 137 taxa; 375bp). They only recovered one ordinal level node when using only H3. According to our phylogenetic results and the results in the papers listed above histone H3 alone provides no new marker for studying deep molecular evolution of the Het- erobranchia due to the high grade of conservation and the low phylogenetic signal for deeper nodes. There were several indices defined by the results of our statistical tests supporting this assumption. We observed a high codon usage bias (see tab. 2) in our alignment which was also indicated by an increasing frequency of C- and G-ending codons and fewer A- and U-ending codons (SHIELDS et al. 1988). High C+G content at silent sites re- flects the effect of selection (SHIELDS et al. 1988) while selective constraints against certain codons might reduce the amount of phylogenetic noise caused by synonymous substitution at either first or third codon positions (BROWN et al. 1999). However, our data suggest that a bias in codon usage will not necessarily be indicative of the phylogenetic utility of a sequence. Despite a high codon usage bias our computed phylogenetic trees showed a poor resolution of the deeper nodes. An explanation for this could be that the pressure to obtain the favoured codon had partially obscured the phylogenetic signal. COLGAN et al. (1998, 2000) made similar observations and conclud- ed that apparent, high codon-usage bias as found for the H3 data does not necessarily result in high phylogenetic consistency for DNA sequences. In the studies presented by Brown et al. (1999) a lack of agreement of the H3 analyses with morphology occurs despite very high codon usage bias. They concluded that whilst selective con- straints may have reduced the absolute rate of synonymous substitutions, the pressure in favor of (homoplastic) resti- tution of the favoured codon has at least a partially ob- scured phylogenetic signal. Hence, codon usage bias does not necessarily mean that a gene sequence will be phylo- genetically useful. The degree of bias (Yx2n) showed a high value of 0,617. It was similar to the values observed in gastropods (0.60) (COLGAN et al. 2000) and polychaetes (0.665) (Brown et al. 1999) and higher than the values of Drosophila melanogaster (FITCH & STRAUSBAUGH 1993) and arthro- pods (0.37) (COLGAN et al. 1998). Another indication suggesting the problems of H3 as a marker for studying deep molecular evolution was the high grade of conservation indicated by the lack of parsimo- ny-informative sites in the amino acid alignment. Additional important evidence was the observed substi- tution saturation at the third codon position. A saturation is caused by multiple-hits which render homoplasious changes. Homoplasy on the basis of saturation in substi- tution is one of the major problems in molecular phylo- genetics (TILLIER et al. 1996). This problem generally be- comes more relevant at progressively higher taxonomic levels (BOORE & BROWN 1998). Ifnumerous substitutions occur at the same position, a hiding or completely eras- ing of the ancient phylogenetic signal could be the result (Lopez et al. 1999). In order to avoid a decrease of the phylogenetic information contained in the sequences, we excluded the third codon position in further analyses. However, this was the position with the most variable sites (78 of 82 positions) in our data set. With the exclusion there was no phylogenetic information left for a resolu- tion of the deeper nodes. Trees which resemble combs at the base resulted (see fig. 3). The entropy was calculated for each position in the alignment to further assess the in- fluence of the 3 codon positions. A high entropy value im- plies that the nucleotides occur almost equally at this po- sition within the alignment. An almost equal distribution of the nucleotides at one position indicates a-less selec- tive constraint allowing a higher frequency of substitu- tions. The average values for each of the three codon po- sitions indicate a less selective constraint for the third codon position hence supporting the previous result of this codon position being saturated. It is questionable if given a larger data set, the noise will eventually succumb to the signal. There are few examples where the expansion of an ambiguous data set has result- ed in a convincing phylogeny (BOORE & BROWN 1998). Surprisingly, there was a good resolution on genera lev- el. Analyses conducted with maximum parsimony and Bayesian inference (with all data) recovered all (or near- ly all) genera mostly with statistically significant support- ed nodes. However, our findings should be consider pre- liminary and further studies including more genera are necessary to test the possible utility of histone H3 for the resolution of recent splits. In other studies (COLGAN et al. 1998; OKuso et al. 2003; LINDGREN et al. 2004) some (but not all) genera were found to be monophyletic but due to the question the authors intended to answer, their studies Bonner zoologische Beiträge 55 (2006) 201 in regards to taxon sampling lacked genera represented by more than one species. In conclusion, still slowly evolving genes for the resolu- tion of the deeper nodes in gastropod phylogeny are miss- ing. To test the Heterobranchia concept as outlined by HASZPRUNAR (1988) other markers have to be found with sufficient variability but no substitution saturation. Acknowledgements. We are grateful to Hermann Dreyer for providing the H3 primers and to the following colleagues and friends for providing samples: Gilianne and Jon Brodie (Townsville), Alan Hodgson (Grahamstown), Michael Schródl (Munich), Enrico Schwabe (Munich), Verena Vonnemann (Bochum), Heike Wágele (Bonn), Tim Wollesen (Frankfurt), as well as Bruce Marshall (Te Papa Tongarewa, Wellington) for his support during a collecting trip to New Zealand. This project was funded by the German Science Foundation (KL 1303/4-1). REFERENCES BOORE, J. L. & Brown, W. M. 1998. Big trees from little genomes: mitochondrial gene order as a phylogenetic tool. Current Opinion in Genetics and Development 8(6): 668-674. BROWN, S., ROUSE, G., HUTCHINGS, P. & COLGAN, D. 1999. As- sessing the usefulness of histone H3, U2 snRNA and 28S rD- NA in analyses of polychaete relationships. Australian Jour- nal of Zoology 47: 499-516. COLGAN, D. J., MCLAUCHLAN, A., WILSON, G. D. F., LIVINGSTON, S. P., EDGECOMBE, G. 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Reconstruction of the Phylogeny of the Opisthobranchia (Mollusca: Gastropoda) by means of 18S and 202 Angela DINAPOLI et al.: Utility of H3-Gene in Heterobranchia 28S rRNA Gene Sequences. Journal of Molluscan Studies 71: Authors’ addresses: Angela DINAPOLI (corresponding au- 113-125. o thor) and Annette KLUSSMANN-KOLB, Institut for Ecolo- XIA, X. & XIE, Z. 2001. DAMBE: Data analysis in molecular ey, Evolution and Diversity — Phylogeny and Systemat- biology and evolution. Journal of Heredity 92: 371-373. : ; : : Xrá. XX, 7: Sar, Ms Caen, L. de Wang, Y 2003, aw. 195 J. W. Goethe-University, Frankfurt am Main, Ger- index of substitution saturation and its application. Molecu- Many, a.dinapoli@wbio.uni-frankfurt.de. lar Phylogenetics and Evolution 26: 1-7. Bonner zoologische Beiträge Band 55 (2006) Heft 3/4 Seiten 203-222 Bonn, November 2007 Biogeographic considerations of the Opisthobranchia (Mollusca: Gastropoda) fauna from the Brazilian littoral and nearby areas“ Francisco J. GARCÍA?), Marta DOMINGUEZ?) & Jesús S. TRONCOSO?) Departamento de Sistemas Físicos, Químicos y Naturales; Facultad Ciencias Experimentales; Universidad Pablo de Olavide, Sevilla, Spain 2)Departamento de Ecología y Biología Animal, Facultad Ciencias del Mar, Universidad de Vigo, Vigo, Spain *Paper presented to the 2nd International Workshop on Opisthobranchia, ZFMK, Bonn, Germany, September 20th to 22nd, 2006 Abstract. The aim of this paper is to examine the biogeographical distribution of the littoral and sub-littoral opistho- branch gastropods from Brazil and nearby areas. On the basis of published literature and personal data, a zoogeographic study was undertaken and the Brazilian region was compared with the Caribbean and Argentinean regions. The Brazili- an littoral has been divided into six zones, based in the oceanographic features established by CASTRO € MIRANDA (1998). 466 species belonging to the orders Cephalaspidea, Anaspidea, Sacoglossa, Notaspidea and Nudibranchia from the Ca- ribbean to the Argentinean region were considered in this study. The number of opisthobranch species is highest in Ca- ribbean areas. Along the Brazilian littoral, South Brazilian Bight is the region with highest richness, while Ormoco-Ama- pá and Amazon shelf are the areas with lower species numbers. The similarity analysis shows that some possible geo- graphic barriers act to the distribution of the opisthobranchs. For each locality considered in the study, the percentage of species that extends northward is higher than southward. Keywords. Brazil, Caribbean region, Argentinean region, biogeography, species list. 1. INTRODUCTION Faunistic inventories give fundamental information for many basic and applied scientific disciplines, such as ecol- ogy and biogeography (STORK & SAMWAYS 1995). An es- sential tool to establish inventories is taxonomy, which supplies a reference system for the biodiversity (BISBY 1995). The level of faunistic and taxonomical knowledge from different regions varies considerably, which raises a problem for the accomplishment of biogeographic stud- ies. Nevertheless, the elaboration of this type of studies offers insights into possible models of distribution of a tax- onomic group throughout a more or less extensive area. Opisthobranch gastropods are well represented in most marine habitats from equatorial to polar regions. Ernst and Eveline Marcus studied this group for more than thirty years in West Atlantic temperate and warmer waters. Re- cently, other authors have provided new contributions to the knowledge of the Brazilian opisthobranchs (TRONCOSO et al. 1998; García et al. 2002; GARCÍA & TRONCOSO 2003, 2004; PADULA & ABSALAO 2005; PoLa et al. 2005; DOMÍNGUEZ et al. 2006 a, b; VALDÉS et al. 2006). The ac- tual level of knowledge of the opisthobranchs from Brazilian and Caribbean regions (1.e. MARCUS 1977; MAR- cus & MARCUS 1967a; THOMPSON 1977, 1980; ESPINOSA & ORTEA 2001; ROSENBERG 2005; VALDÉS et al. 2006) per- mits some biogeographical considerations; however, fu- ture studies along some areas of Brazilian coast are nec- essary to obtain a more complete knowledge of this fau- na. The work of Marcus and Marcus was focused around Sáo Paulo and Rio de Janeiro areas. The necessity of more faunistic studies in Brazil is obvious when comparing the number of species cited along Brazilian coasts (280 species, after MARCUS 1977) with, for example, the Iber- 1an Peninsula (523 species, after CERVERA et al. 2004). Here, we present a study on the diversity of the Opistho- branchs along the Brazilian coasts, using as biogeograph- ic areas the six zones defined by CASTRO & MIRANDA (1998). These Brazilian zones are also compared with Caribbean and Argentinean regions (1.e. MUNIAIN 1997; SCHRÓDL 1999; ROSENBERG 2005). 204 Francisco J. GARCIA et al.: Biogeography of Brazilian Opisthobranchia 2. MATERIAL AND METHODS 2.1. Biogeographical areas This research was conducted by comparing littoral and sub-littoral Opisthobranch fauna from the Brazilian shores with those from the Caribbean and Argentinean bio- geographic regions. Species checklists were compiled by combining data from bibliographical sources and person- al observations (included in the references). The total num- ber of species considered for this study is 466, belonging to the orders Cephalaspidea, Anaspidea, Sacoglossa, No- taspidea and Nudibranchia (Table 1). In this paper, we adopted the six oceanographic zones de- fined in CASTRO & MIRANDA (1998) for Brazil (Fig. 1). The geographical limits and features of these areas are shown in Table 2. BER (72) = CR-Cro (79) MY CC-CR (173) =, Wl (250) CRo-O ABC (55) CRa-O (178) 0-4 (12) S (74) EBS (27) ACR (61) SBB (121) SBS (18) Fig. 1. Limits of the areas considered along Brazilian coasts, based on CASTRO & MIRANDA (1998) and nearby zones based on BrIGGS (1974). The number of species considered for each region is indicated. Within the Caribbean region we have considered the provinces defined in BRIGGS (1974) (Fig. 1): CC-CR, extends from Cape Canaveral to Cabo Romano CR-CRo, extends from Cape Romano to Cape Rojo CRo-O, extends from Cape Rojo to the mouth of the Orinoco River O-A, extends from Orinoco River to Amapa ABC, Includes Aruba, Bonaire and Curacao WI West Indies (WI) Bermudas Is (BER). In addition, we consider the Argentinean region (ARG) between the borders of Brazil to Uruguay up to 43-44°S (Chubut). 2.2. Community analysis and measurement of biodi- versity In order to examine diversity within the opisthobranch communities, data by region were subjected to a multi- variate analysis using the Bray Curtis similarity measure and non-metric Multidimensional Scaling Ordination (MDS). The Bray-Curtis index (BRAY & CURTIS 1957) was cho- sen because it does not consider double absences in its cal- culations. The results were then graphically described using dendrograms with the UPGMA (unweighted pair- group methods using arithmetic averages) aggregation al- gorithm (SNEATH & SOKAL 1973). The ordination analy- ses were carried out by means of an MDS (non metric mul- tidimensional scaling program) based on the similarity ma- trix between stations. For two different station groups a requirement is to iden- tify which species account for the observed assemblage difference (CLARKE & GORLEY, 2001). The SIMPER rou- tine was used to identify taxa that greatly contributed to differentiate station groups. The software used was P.R.I.M.E.R. (Plymouth Routines in Multivariate Ecolo- gical Research) version 5.2.8. for Windows. 3. RESULTS 466 species of opisthobranchs were found to occur in the Western Atlantic Ocean from Cape Canaveral to the Ar- gentinean province (the Atlantic Magellanic region is not included). Table 3 shows the number of species by order or suborder for each zoogeographic area. The number of Bonner zoologische Beiträge 55 (2006) 205 opisthobranch species for each area varıes remarkably. WI (West Indies) has the highest number of species (250 species), although, in general, the number is also high in the Caribbean areas Cro-O and CC-CR. Along the Brazilian zones, the number of species 1s lower and there is also a notable difference between the different zones. SBB is the zone with the highest number of species while AS is the zone with the lowest number (122 and 12 species, respectively). The number of species from the Ar- gentinean region (ARG) is moderately low. 3.1. Faunal affinities The cluster analysis using the six Brazilian zones shows a first division in which AS separate from the remaining zones (Fig. 2A). A second division separates SBS, and fi- nally, the rest of the areas splits into two groups; a group including the north-eastern Brazilian zones EBS and NBS, and the other including ACR and SBB. A two-dimension- al representation of the analysis MDS shows this same grouping pattern (Fig. 2B). Similarity A Fig. 2. Cluster classification (A) and MDS ordination (B) of the biogeographic areas based on the presence-absence of spe- cies in the areas from Brazil. 4 SBS AO-A ® Arg AAS Awı A CC-CR A CRo-O A CR-CRo A ABC A Ber 4 SBB AACR 4 EBS 4 NBS 0 20 40 60 80 100 A Similarity A Stress: 0,09 CC-CR A SBS E] B Fig. 2. Cluster classification (A) and MDS ordination (B) of the biogeographic areas based on the presence-absence of spe- cies in all areas included in this study. Black triangles, Brazili- an regions; white triangles. Caribbean provinces; black circle, Argentinean region. Including all geographical areas from Cape Canaveral and Bermudas to Chubut (Argentinean region), the cluster analysis and MDS show a first division in which Orinoco-Amapa (O-A), the Brazilian areas SBS and AS and the Argentinean region (ARG) separate from the re- maining regions (Fig. 3). Those separate in two groups. One includes the north-eastern Brazilian zones EBS and NBS, while the other includes the remaining zones. The latter divides itself into two subgroups, one including the Brazilian regions SBB and ACR, and the other subgroup including all Caribbean provinces. The SIMPER analysis identified four distinct groups. In group | (SBS, O-A, ARG, AS), the following taxa con- tribute to the similarity (up to 85%): Cephalaspidea (Acteocina bidentata, Volvulella persimilis, Acteocina can- dei, Acteon pelecais) and Notaspidea (Pleurobranchaea inconspicua). Group 2 (WI, CC-CR, CRo-O, CR-CRo, ABC, Ber) is mainly characterised by species of the or- 206 Francisco J. GARCÍA et al.: Biogeography of Brazilian Opisthobranchia ders Cephalaspidea (Volvulella, Bulla, Haminoea, Hydati- na, Micromelo), Notaspidea (Umbraculum, Pleuro- branchus), Anaspidea (Stylocheilus, Bursatella, Aplysia, Bosellia), Sacoglossa (Cylindrobulla, Oxynoe, Tridachia, Elvsia) Dendronotacea (Scyllaea) and Nudibranchia (Spurilla, Chromodoris). Group 3 (SBB, ACR) is charac- terised by the presence of species of the order Nudi- branchia, mainly Doridina (Dendrodoris krebsii, Cadlina rumia, Tyrinna evelinae, Diaulula greelevi, Discodoris evelinae, Chromodoris clenchi), Arminina (Armina) and Aeolidina (Spurilla neapolitana, Phidiana lynceus, Fla- bellina engeli, Glaucus atlanticus). Group 4 (EBS, NBS) is determined (up to a cumulative 90%) by the presence of the orders Nudibranchia (Doto divae, Diaulula gree- leyi), Notaspidea (Pleurobranchaea inconspicua, Umbrac- ulum umbraculum, Berthellina quadridens), Anaspidea (Phyllaplysia engeli, Stvlocheilus striatus, Aplysia dacty- lomela, Aplysia parvula, Bursatella leachii), Saccoglos- sa (Cylindrobulla beauii, Oxynoe antillarum, Elysia tuca), and Cephalaspidea (Philine sagra, Hydatina vesicaria, Mi- cromelo undatus, Haminoea elegans, Chelidonura petra) In terms of dissimilarity, the species Spurilla neapolitana, Scyllaea pelagica, Pleurobranchus areolatus, Tridachia crispata, Aplysia dactylomela, Aplysia fasciata, Oxynoe antillarum contribute greatly to differentiate groups | and 2. Group | differed from group 3 due to Phidiana lynceus, Spurilla neapolitana, Flabellina engeli, Facelina coenda, Anteaeolidiella indica, Berghia benteva, Taringa telopia, Tyrinna evelinae, Siraius ilo, Jorunna spazzola, Okenia zoobotryon, Hallaxa apefae, Doris verrucosa, Discodoris evelinae, Chromodoris neona, Dendrodoris krebsii, Diaul- 80 70 60 50 40 30 20 10 CC- | CR- | Cro- CR | CRo O 0 Fig. 4. | i Y E OSO aaa ax [aafesfwifm ||| o po pos 0 [a] me] ss [one camas [%fels,efolol;s}ols is: |m ula greeleyi, Cadlina rumia, Chromodoris clenchi, Berthella agassizii, Berthella stellata, Aplysia dacty- lomela, Aplysia fasciata, Oxynoe antillarum, Ascobulla ul- la, Navanax aenigmaticus. Group 1 differed from group 4 due to Doto divae, Diaulula greeleyi, Phyllaplysia en- geli, Stylocheilus striatus, Berthellina quadridens, Aplysia dactylomela, Aplysia parvula, Oxynoe antillarum, Elysia tuca, Micromelo undatus, Chelidonura petra, Atys ri- iseanus. Differences between group 2 and group 4 were mainly due to Tridachia crispata, Aplysia fasciata, Che- lidonura petra. The group 2 differed from group 3 due to Facelina coenda, Berghia benteva, Scyllaea pelagica, Sir- aius ilo, Hallaxa apefae, Chromodoris neona, Tridachia crispata, Philine mera, Cylindrobulla beauii, Acteon pele- cais. And finally the species Phidiana Iynceus, Flabelli- na engeli, Glaucus atlanticus, Facelina coenda, Armina muelleri, Anteaeolidiella indica, Berghia benteva, Doto di- vae, Taringa telopia, Tyrinna evelinae, Siraius ilo, Oke- nia zoobotryon, Hallaxa apefae, Chromodoris neona, Dendrodoris krebsii, Cadlina rumia explained most of the dissimilarity between groups 3 and 4. Figure 4 shows the number and percentage of endemic species for each biogeographic area. Those species con- sidered as endemic have been cited only at one zoogeo- graphic region from the area of study. Geographic distri- bution along other biogeographic regions was not consid- ered for this study. The level of endemism varies notably along the different zoogeographic areas. The highest val- ue was found in the Argentinean region (68.6%). This high value is due to a southward distribution to the Magellan region of the fauna from this area. The Brazilian zones Number and percentage of endemic species for each faunistic area. Bonner zoologische Beiträge 55 (2006) 207 have percentages between 0 % in Amazon Shelf (AS) and East Brazilian Shelf (EBS), and 28.7 % in South Brazil- ian Bight (SBB). Among the Caribbean provinces, the per- centage of endemism varies between 7.1 %, at Aruba, Bonaire and Curacao (ABC), and 22 % in the West Indi- an region. Table 4 shows the percentage of species from each region present in other localities. The percentage is increasing, when extended northward, and decreasing, when extend- ed southward. 4. DISCUSSION An aspect to be considered in this study is that the data on species distribution and the level of knowledge of the communities vary across the geographical regions. Thus, the results presented here must be considered as tentative. The number of opisthobranch species between biogeo- graphic areas varies remarkably being higher in the Caribbean: West Indies (WI: 250 species), Cape Rojo- Orinoco (CRo-O: 178) and Cape Canaveral-Cape Romano (CC-CR: 173). The south Brazilian Bight (SBB) is the zone in Brazil with the highest richness (122 species), fol- lowed by the north Brazilian Shelf (NBS: 75) and the Abrolhos-Campos Region (ACR: 62). The areas with the lowest species richness are Oricono-Amapa (O-A) and Amazon Shelf (AS), both with 12 species, followed by South Brazilian Shelf (SBS) with 18 species. This could be related to oceanographic conditions, such as the effect of the Amazon River in the AS zone, and with a differ- ence in sampling effort; the south Brazilian Bight is the zone where Ernst and Eveline Marcus did many of their collections. The first division observed in the similarity analysis in- cludes four geographical regions with features that seem to act as barriers to the distribution of opisthobranchs. The coasts from Orinoco-Amapa and Amazon Shelf are influ- enced by the Amazon and Orinoco rivers whose plumes spread north-westward for more than 1,000 km into the North Atlantic (CASTRO & MIRANDA 1998). These areas are characterised by soft bottom, turbid waters and fresh- water runoff and they have been recognised as barriers to the dispersal of corals (Cox & Moore 2000), rocky shore gastropods (VERMEN 1978) and shallow water reef fish- es (GILBERT 1972). The scarce opisthobranch fauna from Orinoco-Amapa and Amazon Shelf and the composition of species present in these areas are likely related to en- vironmental characteristics of the region. Species present in these areas are generally cephalaspideans and armi- nacean, which frequently live in sand or mud. A similar situation is found on the South Brazilian Shelf, where the low species numbers may be related to the ef- fect of the Patos Lagoon River plume with an annual mean discharge of about 2000 m3s-! (MARQUES et al. 2006). This area is influenced during the winter by Subantarctic wa- ter. The coastal area of the Argentinean biogeographic province is a transition zone characterised by processes of mixing and instability of the water masses. This province includes geographical features like the Rio de la Plata. The estuaries influence the primary and secondary production in the area and consequently, the distribution of species. Moreover, there is an interaction off the coast between the Malvinas current flowing on the slope from the south with cold Subantarctic waters rich in nutrients and the Brazilian current, with temperatures higher than 20° C and salinity over 36.0 ppt. This determines the pres- ence of eurythermal and euryhaline species (BOSCH! 2000). The fauna of opisthobranchs is formed mainly by nudi- branchs, which have their northern distribution limit at the border between the Argentinean region and the South Brazilian Shelf, extending southwards to Subantarctic re- gions. Cluster analyses show two Brazilian groups, composed of NBS-EBS and ACR-SBB. In general, these groups coin- cide geographically with those indicated by FLOETER et al. (2001) to the reef-fish fauna of the Brazilian coast. These authors considered several regions like the South and South-eastern coastal reefs, from the Guarapari islands to Santa Catarina (areas included in our analysis as the group ACR-SBB), and the North-eastern coast, extending from the Manuel Luis reefs to Abrolhos Archipelago (areas in- cluded in our analysis as the group NBS-EBS). The southern and south-eastern coastal reefs, cited by FLOETER et al. (2001) show lower mean annual water tem- perature, relatively higher primary production and a large shelf width. The reef-fish fauna living in this area appears to be the richest of Brazil, due to the mix of tropical and subtropical elements. The area is subjected to a relative- ly intense seasonal upwelling promoted by the South At- lantic Central Water, bringing low-temperature (<1 8°C) and nutrient-rich waters close to the coastline (EKAU & KnoPPERS 1999). FLOETER et al. (2001) stated that a con- siderable number of Caribbean reef fishes found in this region are absent from the north-eastern sites. We found a similar pattern in opisthobranchs. The cluster analysis groups ACR-SBB closer to the Caribbean provinces than to other Brazilian zones. On the other hand, ACR and SBB are the Brazilian areas with the highest richness in opistho- branchs. This could be related with environmental features (as is observed for reef-fishes; FLOETER et al. 2001). Nev- ertheless, in addition to environmental factors, differences 208 Francisco J. GARCIA et al.: Biogeography of Brazilian Opisthobranchia in the richness of opisthobranchs for each area may de- pend upon other factors, such as discrepancies in sampling effort. The fauna from Brazil is better known in the South Brazilian Bight and Abrolhos-Campos zone, where Eve- line and Ernst Marcus conducted research for over 30 years. The north-eastern region as is described by FLOETER et al. (2001) for fishes, which nearly overlap with the group NBS-EBS, is characterised by its relatively warm waters, a weak seasonal signal and a small vertical temperature gradient; the circulation is influenced northward by the North Brazilian Current, and southward by the Brazilian Current (CASTRO & MIRANDA 1998). The reef formation consists of coralline algal crusts over a rocky substrate, hermatypic and fire corals, as well as sponges (FLOETER et al. 2001). The narrow and open shelf is an oligotroph- ic system almost entirely covered by carbonate sediments due to little freshwater input and the coast is influenced by the South Equatorial Current (KNopPERS et al. 1999). In these areas herbivorous Sacoglossa and Anaspidea are more abundant. This trend of an increase in abundance in herbivores towards the tropical zone was previously ob- served in the Brazilian reef fishes (FERREIRA et al. 2004). 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Malacolog 4.1: A Database of Western At- lantic Marine Mollusca. [WWW database (version 4.1.0)] URL http: //www.malacolog.org/. ScHróDL, M. 1999. Zoogeographic relationships of the Magel- lan Nudibranchia (Mollusca: Opisthobranchia) with particu- lar reference to species from adjacent regions. Scientia Ma- rina 63(Supplement 1): 409-416. SNEATH, P. H. A. & SOKAL, R. R. 1973. Numerical Taxonomy. The principles and practice of numerical classification. WH Freeman, San Francisco. Stork, N. E. & SAmMways, M. J. 1995. Inventorying and Mon- itoring. Pp. 453-544 in: Heywoob, V.H. (ed.) Global Biodi- versity Assessment. Cambridge University Press, Cambridge, UK. TEMPLADO, J.; LUQUE, A. A. & ORTEA, J. 1987. A new species of Aegires Loven, 1844 (Opisthobranchia: Doridacea: Aegit- etidae) from the Caribbean Sea: Aegires ortizi spec. nov., with comparative descriptions of the North Atlantic species of this genus. The Veliger 29 (3): 303-307. THompson, T. 1977. Jamaican Opisthobranchs Molluscs I. 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GARCIA (corresponding author), Departamento de Sistemas Fisicos, Quimicos y Naturales; Facultad Ciencias Experimentales; Universidad Pablo de Olavide, Sevilla, Spain, E-mail: fjgarcia@us.es; Marta DOMINGUEZ, Departamento de Ecologia y Biologia Animal, Facultad Ciencias del Mar, Universidad de Vi- go, Campus de Vigo, E-36200, Vigo, Spain; Jesus S. TRONCOSO, Departamento de Ecologia y Biologia Animal, Facultad Ciencias del Mar, Universidad de Vigo, Campus de Vigo, E-36200, Vigo, Spain. E-mail: malvarez(@ uvigo.es; troncoso(Vuvigo.es. 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SUO[ 1U9$UISI.] DULDODIY 8561 ‘AUDI Y UOSS[O PROIÍSUOIUL DUIIODIY (0061 ‘USN 9 [[LLISA) sua.uımsop DULIOAJIF (1981 ‘AUSIQIO P) 19puba DUIIOAJIP (92781 ABS) DID/NIDUDI DULDODIY (FEST “ISUIA) PI] PulDoalayF (Ir8I Ausigıo,p) PIDJUIPIG DULIOIAJOP (1061 eneurA) sısuopnuntog DULDOD)D 7 PRG “UIS[IAAA Y USW PID.90 VULIOIAJIP somods Francisco J. GARCIA et al.: Biogeography of Brazilian Opisthobranchia N a Species Bulla striata Bruguiere, 1792 Bullina exquisita MeGinty, 1955 Bullina torrei (Aguayo & Rehder, 1936) Chelidonura berolina Marcus & Marcus, 1970 Chelidonura cubana Ortea & Martinez, 1997 Chelidonura hirundinina (Quoy & Gaimard, 1833) Chelidonura hummelincki Marcus & Marcus, 1970 Chelidonura juancarlosi Ortea & Espinosa, 1998 Chelidonura mariagordae Ortea, Espinosa & Moro, 2004 Chelidonura petra Marcus, 1976 Chelidonura sabina Marcus & Marcus, 1970 Crenilabium exile (Jeffreys, 1870) Cylichna alba (Brown, 1827) Cylichna auberii (d’Orbigny, 1841) Cylichna discus Watson, 1883 Cylichna eburnea A. E. Verrill, 1885 Cylichna georgiana (Strebel, 1908) Cylichna krebsii Morch, 1875 Cylichna verrillii Dall, 1889 Cylichna vortex (Dall, 1881) Cylindrobulla beauii P. Fischer, 1857 Cylindrobulla gigas Mikkelsen, 1998 Diaphana caribaea Espinosa, Ortea & Fernandez-Garces, 2001 Diaphana seguenzae (Watson, 1886) Gastropteron chacmol Gosliner, 1989 Gastropteron hamanni Gosliner, 1989 Gastropteron rubrum (Rafinesque, 1814) Gastropteron vespertilium Gosliner & Armes, 1984 Haminoea antillarum (d’Orbigny, 1841) Haminoea elegans (Gray, 1825) Haminoea glabra (A. Adams, 1850) Haminoea petitii (d’Orbigny, 1841) Haminoea solitaria (Say, 1822) Haminoea succinea (Conrad, 1846) Hydatina vesicaria (Lightfoot, 1786) Japonacteon pusillus (Forbes, 1844) Micromelo undatus (Bruguiére, 1792) Mysouffa cumingii (A. Adams, 1855) Mysouffa turrita (Watson, 1883) Navanax aenigmaticus (Bergh, 1893) Navanax orbygnianus (Rochebrunne, 1881) Ovulactaeon meekii Dall, 1889 Philine alba Mattox, 1958 Philine argentina Carcelles, 1947 Philine caballeri Ortea, Espinosa & Moro, 2001 Philine candeana (d’Orbigny, 1841) Philine falklandica A. W. B. Powell, 1951 O+O++++++0 [e ocoooo CR-Cro CRo-O O-A + + 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 + 0 0 + 0 0 0 + 0 0 0 0 0 0 0 + 0 0 + 0 - - + + 0 0 0 0 + ES 0 0 0 0 + 0 + + 0 + 0 0 + - + + 0 0 0 + 0 0 0 0 0 + 0 0 0 0 0 + 0 0 0 + 0 0 0 0 oOOooOoooO00 0 0 0000 + Wi BER 0 0 it 0 + 0 + 0 + 0 “+ 0 + 0 + 0 + 0 0 0 0 0 0 0 7 0 + 0 0 0 0 0 + 0 0 0 0 0 + + + 0 0 0 + 0 + 0 0 0 0 0 + + 0 0 + 0 + 0 + + ES Zn 0 0 + + 0 0 E 0 + 0 + 0 + 0 0 0 0 0 + 0 + 0 0 0 Nn oooooooo 0 oooo NBS EBS ACR SBB_ SBS + 0 + + + 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 + + 0 0 0 0 0 0 0 0 0 0 0 0 0 + 0 0 0 0 0 0 0 0 0 + 0 0 0 + 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 + 0 0 0 0 0 ! ! 0 0 0 0 0 0 0 0 0 0 0 0 0 + 0 0 0 0 0 0 0 + 0 0 0 0 0 0 0 0 0 + 0 0 0 0 0 0 > 0 0 0 0 : + + 0 0 0 0 0 + 0 0 0 0 0 0 0 0 0 0 0 0 0 0 + - 0 0 0 0 0 0 0 0 + + 0 0 0 0 - - + 0 0 0 0 0 0 0 + 0 0 0 0 0 0 + 0 0 0 0 0 0 0 0 + 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 + ARG ocoo+00000000000000000o0o Bonner zoologische Beitráge 55 (2006) | | | | | | | | | | | | | TL6I "SNIDABIA] ISNIAADU DIPJASOg EL6| 'SNOABIA] AVAUULAOI DI]ISOZ £96] ‘Spunuupy vaggiins piuljaysiag (0161 “SMIRIJA] 29 sale) jj D]¡NQODS Y ESG] “OHJ-10AN1J psouuiof SiSdOIS A dy (yb8 | “UDAOT) pIsapou DI.DP]Y L961 Aue DUDISPXA] D/]2]NA]04 (SL81 YQ) DI994 D]]9]NAJOA (SL81 UdIOJA) Sıyruns.od D]/9]NA]04 (£88] UoseM) PpNoOLAdnvd pjopna]og (5881 “Ysng) pınunu D]]9NA]O4 (LEST “PMOD) pıpıng PIUOPIJO] 1881 [ed uosjpa dopunydnas (2481 'swepy 2 spoysimw) smiv1usojsund vopuvydoos SS61 AUIDIJA Kugspd sapunydnasy +y881 ILLA “4 “Y sıjıgou sapunydvay 9561 sg 9DS10] 1opunydoas L961 ‘SHOR 2 SODIBIA sn1imp dopupydoas (1881 TIRO) Prrydowsypg dopunydvas (£981 “Y9OJA) Duisvaid pulouny 1061 usa “4 “Y pnardsuooun pumuny (£961 “SMIBIA Y SNOIBIA) 9DA1P DULDUNY TH] ‘AUBIQIO .P VIDLYSIUAS DINIBULY LET ISA v Dpu1u vpnoisury (OrsI ‘suepy 'g :D) smiv1.9soj0und sixpjo1y (1481 ‘AUSIQIO,P) vID9]MS PSMIIY (£881 UOSJeM ) pyıpds PSNIIY 6961 “SNDABJA] 29 SNIIBIN DSOS DSNIIY (6881 TIRO) Pla.od psnjay 181 UIOJA sypydiuo vsnjay (1881 ea) 19/9144 vsnioy (6881 'Teq) prnuop psnjay Z961 ‘SIAM 9 STAM PI0N21/DUDI PSNIIY] (1281 “SÁdIYOf) savao sainounidg (1181 “Suou3nig) sa nosnisoqo snjnouniAg (5881 ‘ysng) 51/0909 snnounidg ESG] “SOOIBIA VINI DINISN] (9961 “SMIBIA IZ SNOIBIA) Psnd sısdounyg 1007 “esourdsq 29 EINIG 1290 sisdou1]1yd L68 | “UOSIPAM DID.ISOAYIDAL DUYY A 6961 SMILE Y SODIBIA 2UUDUANY] OULU (1481 KUSIQIO P) 2.1508 OUI 6881 TIRA Prvunyd au yd 6961 SWR Y SNDIBIA] Po BULLY 6881 Tea unpmqipunfur ou yd 0L81 “SIBS “JA Psonxayf ouyg 6581 “SIS “JN POD aumyd 0 ooo +0o+ +0+ ooo o o FOOoOoOo roooo0o0000 roo+o0 roo0o0o0 rt+tooo ++o++ ooooo+ oco00o00000 o +00000000000o + + Oo++003++ 000 AS OR E =F +00000000o0o+ oooooooo0+ oc0o0o0000o0000000000000o0o00o0o0o0o+00000000000000000o0 ocoo0oooo++00000000000000000000000+O00o00o00o++00 oc00000000000000000000o00oooooo ooo+00++400+000000000000000 ooo+00000000000000000000+000000000000o00o0+ O00O0OoO+ooo co +0+00000000000000000000000%+ 0000000000000 o0ooo COLO Own SS O'S © SO SS RS SS SO + SE SS oS OS OS ES SS So Sc SoSSe. eS +20000000000+0000000000000000000000000o0o+ oo 0000000 SEES MOROS SO ES SEDES ASES SD FF SF SO OOS: CuO SS Fa FF FF O0 Sr Fr Fr SS: SS. SS + SS SS +99 090899 85957 9 599 COSCO SOC 950500995 +5 © SS 959 2.055905 09 oc 5 259 Soo +0 ooo ooo ocoooo SI SS O'S © SO) Ore. Ze +, O c0000000000o DUV SAS HAS MOV SAA SAN SV. aaa DIV V-O 00m MI WII sonads > Francisco J. GARCÍA et al.: Biogeography of Brazilian Opisthobranchia 214 Species Bosellia mimetica Trinchese, 1891 Caliphylla mediterranea A. Costa, 1867 Costasiella lilianae Marcus, 1969 Costasiella nonatoi Marcus & Marcus, 1960 Costasiella ocellifera (Simroth, 1895) Cyerce antillensis Engel, 1927 Cverce cristallina (Trinchese, 1881) Cyerce edmundsi Thompson, 1977 Cyerce habanensis Ortea & Templado, 1989 Elysia canguzua Marcus, 1955 Elysia catulus (Gould, 1870) Elysia cauze Marcus, 1957 Elysia chitwa Marcus, 1955 Elysia chlorotica Gould, 1870 Elysia cornigera Nuttall, 1989 Elysia eugeniae Ortea & Espinosa, 2002 Elysia evelinae Marcus, 1957 Elysia flava A. E. Verrill, 1901 Elysia nisbeti Thompson, 1977 Elysia ornata (Swainson, 1840) Elysia papillosa A. E. Verrill, 1901 Elysia patagonica Muniain & Ortea, 1997 Elysia patina Marcus, 1980 Elysia pratensis Ortea & Espinosa, 1996 Elysia purchoni Thompson, 1977 Elysia serca Marcus, 1955 Elysia subornata A. E. Verrill, 1901 Elysia tuca Marcus & Marcus, 1967 Elysia zuleicae Ortea & Espinosa, 2002 Ercolania cricetus (Marcus & Marcus, 1970) Ercolania funerea (A. Costa, 1867) Ercolania fuscata (Gould, 1870) Ercolania selva Ortea & Espinosa, 2001 Hermaea bifida (Montagu, 1815) Hermaea coirala Marcus, 1955 Hermaea cruciata Gould, 1870 Lobiger souverbii Fischer, 1857 Mourgona germaineae Marcus & Marcus, 1970 Mourgona murca Marcus & Marcus, 1970 Oxynoe aguayoi Jaume, 1945 Oxynoe antillarum Mörch, 1863 Oxynoe azuropunctata Jensen, 1980 Oxynoe panamensis Pilsbry & Olsson, 1943 Phyllobranchillus viridis (Deshayes, 1857) Placida dendritica (Alder & Hancock, 1843) Placida kingstoni Thompson, 1977 Placida verticilata Ortea, 1981 Q 2 A = So++++o++ ++0°00+4 +++ 00404 4 o ++0+0++00+++000+ ++ CR-Cro CRo-O O-A 0 0 0 0 0 0 0 0 0 + 0 0 0 0 0 0 0 + 0 0 0 0 0 0 0 0 te 0 0 0 0 + 0 + 0 oo 0 0 0 + 0 + 0 0 0 + 0 + 0 + + 0 0 0 0 + 0 + 0 + + 0 0 0 0 + 0 + 0 0 0 0 + + 0 + 0 0 0 0 0 + 0 + 0 + 0 0 0 + oo0ooo0o00o00o0o00o00oooo oso 000000000000 0 0 0000 o SO Oo ++-00+ 04 + 00 wi t+ +++ =) +9 +++0+++9000+ + oO +0© oOoooo0o0 +00 0000 oOOOoOooOooO000o0 > Dn o oco00o00o00o00000000o00o00o00on> | | NBS EBS ACR SBB 0 + 0 0 + 0 0 + 0 0 0 + 0 0 0 + 0 0 0 + 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 + 0 0 0 0 0 0 0 = 0 0 0 + 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 + + 0 0 0 0 0 0 0 + 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 = 0 0 0 + + + + 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 + 0 0 0 0 0 0 0 0 0 0 0 + 0 0 0 0 0 0 0 + 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 oo oc00o00o00000000o0o oo ooOooooo0o0 oo 0 0 00000 0000 0 00 7 Bonner zoologische Beiträge 55 (2006) ZOOT ‘ZONSUILIOG 9 OSODUOL| BIDIBL) 10D]DSQD $0.1189 / (98/11 10094817) un nopaiquin un]noD. qui) 0681 RG vUupoiiauy DUIPOJA] (1061 “TILU9A) snadiu SNYIUDAGOAMI] 1061 ‘uosdung X [jeq $19/9D] SNYIUDAQOAMI] +86] mare idsno1 snyoub.1qo.na]d LL6| “UOSdUIOY | apuljaia SAYINDAQOMMI] Y PRG] ‘Snel Sta SNYIUDAGONMMI|] COQ] UIIOJA] 51100210 SNYIUDAQONMI]d 0881 TILDA DP.D] panyouvago.inald (2881 I[LLI9A ) Psago DIDYIUDAGONMNO]J L681 YB19g vnoidsuosul DODYIUDPAGOAMI]J L681 UBIDY 112185P3D PIPYIUDAQOMI] (CORT “Y9OJA) SuSpiuponb purjayLiag (£981 “Y9IOJA) 52.072.119 DUILJAYIAIT PRG "more nun D]]9Y110G (9781 “OSSTY) D1D]]915 P]]941198 (S£81 ‘AUSIQIO,P) porm10801vd pjoyniag (6061 PUB] IB JOBJA) 11218503D D]]941-198 (Z£81 'pleuen wz Aond) $701.18 SNJIIYIOJUS ($781 'pieunen 2 Aon?) ppnvsisuo] sn]19/90/4S (8781 Buey) snul 19 snpayrojAy¢ LL6T ARO vpsvaivus visádo ¡Xy gd SS61 ‘SHIR asua DISAJAD]AYS 6561 egegg vsowp.s Daafımlad (87381 Buey) viafippiad viafiyvlag 9€81 Iddi yg srimzound snyoivjon (8781 Buey) ».19/11G0]0p pafı.ıgp]oq LIST “S[[IAUTe]g 11/9D9] vjjajps.ing 1881 “ULISH 1X09]2 DISáidy £981 “YOY pjnaınd visa dy (1061 “[LL9A) 01.0% pisdjdp TERT “preueo wz Aond) vunynl visa dy (OSQ DA99y Y suULpY) porydv.3093 visa dy 68L1 Y9M04 piwiaspf visA dy QTR] Buey vjawopdjonp visady (1061 ‘Wosdwig 9 |feq) PUI pisa dy L961 MO Y SIOOIBIA 1.19DQ D10yY Z861 APIO IVPNULLDQ P]/9IDA]OA (£981 “YIIOJA) 1UUWD.AYIS DIYIDPIA] (£981 YO) DIOASI.D DIYIDPLLL (1061 "aA “V) Pod vipipLiny | 0007 “esouridsq 9 BO PPzDU pypipiuamy] 0961 “SMOIBIA] A SMILE 15504 LATING (LS61 “SNDIB]JA) 51//29UDA LAB US 9S8] “SODIBIA 51/07 1923US Z96 1 “9DUET SNIDINAOISNS LAS US o oO o oo o 200000000000 oo+0o0+00 ooooococoocoa oOOooooco00o0 oo++00000+0 +O+000003+35+0 Te+rsttstso+++99e ocooo +0 ooooot+o oso Co + + SS ++ So + SO coooooocoococoeooco ooo o+ + oooocoocococooococococoe ++0+ S + SiS + e Ketten POooo JJ so +ed FE FOoox+o F+oo+++ So 55590905 9 90909 599 9 90° 9 999 90 9° 59092 59 9° 90 SO 5900009909 009599 9 90909 + © StOocosOr + +O Oo Oo fF fo +t oF OF oS + Fo +O + 995° Fo 2 5 5 SS Co Oo + 9 SC + Oo oO O'S ocooocoocooocooocoocoocoooocooocoocoor+acacgncocococooocoococortocoococoococoe FOO+O+++A+4++F+++++40+++200+++4+00+00+000++0++000000+0 oco0oo0o0o0o0oo0o0o0o00o0o0o0000000000000000o00o0+o0oo0ooOo SoOooooooooooooooo000 + SE SO SS: Or FO rS Ss rt sS sos HS + +5 #65 SS "SISI SO SO 'O SO -+ Fo se Foo + rss 0 oO Oo SO ocsossooosoooo oco00oooo+o0ooo ooooooooo+ + SIE HO sb SO ++ + DIV V-O OH) CDAD UDI a ans 2 | Z. n =< x | ES = saa x Q « | UV sas aas Francisco J. GARCÍA et al.: Biogeography of Brazilian Opisthobranchia 216 Species Aegires gomezi Ortea, Luque & Templado, 1990 Aegires ortizi Templado, Luque & Ortea, 1987 Aegires sublaevis Odhner, 1932 Ancula espinosai Ortea, 2001 Aphelodoris antillensis Bergh, 1879 Aporodoris millegrana (Alder & Hancock, 1854) Atagema browni Thompson, 1980 Atagema hispida (d’Orbigny, 1837) Atagema prea (Marcus & Marcus, 1967) Cadlina rumia Marcus, 1955 Cadlina scabriuscula (Bergh, 1890) Cadlina sparsa (Odhner, 1921) Ceratophyllidia papilligera (Bergh, 1890) Chromodoris binza Marcus & Marcus, 1963 Chromodoris clenchi (Russell, 1935) Chromodoris dictva Marcus & Marcus, 1970 Chromodoris grahami Thompson, 1980 Chromodoris kempfi Marcus, 1971 Chromodoris neona (Marcus, 1955) Chromodoris perola Marcus, 1976 Chromodoris ponga Marcus & Marcus, 1970 Chromodoris punctilucens Bergh, 1890 Chromodoris roseopicta Verrill, 1900 Corambe evelinae Marcus, 1958 Corambe obscura A. E. Verrill, 1870 Dendrodoris krebsii (Mörch, 1863) Dendrodoris magagnai Ortea & Espinosa, 2001 Dendrodoris senegalensis Bouchet 1977 Dendrodoris warta Marcus & Gallagher, 1976 Diaulula greeleyi (MacFarland, 1909) Diaulula punctuolata (d’Orbigny, 1837) Diaulula worki (Marcus & Marcus, 1967) Discodoris alba White, 1952 Discodoris branneri MacFarland, 1909 Discodoris evelinae Marcus, 1955 Discodoris mortenseni Marcus & Marcus, 1963 Discodoris muta Bergh, 1877 Discodoris notha Bergh, 1877 Discodoris ketos gila (Marcus & Marcus, 1970) Discodoris phoca Marcus & Marcus, 1967 Discodoris purcina Marcus & Marcus, 1967 Discodoris voniheringi MacFarland, 1909 Doridella carambola (Marcus, 1955) Doriopsilla areolata nigrolineata Meyer, 1977 Doriopsilla espinosai Valdés & Ortea, 1998 Doriopsilla pharpa Marcus, 1961 Doris bicolor (Bergh, 1884) CC-CR o+ 2000 + +ooo++o++ +0 000 + CR-Cro CRo-O O-A + OOOO + 44 0° oO Oo +0 00 + + 00 oO ocooo oooooo ++ @OO0+F4+F 04+ 0400+ 4 I $ + + + 0 0 oooo4+40+ 00 ooo eo coo AS NBS 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 + 0 0 0 0 0 + 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 = 0 0 0 + 0 0 0 0 0 0 0 + 0 + 0 0 0 0 0 0 0 0 0 + 0 0 0 + 0 0 0 0 0 0 0 0 0 0 EBS +ooccoocococo o ACR Oo+4S000¿+4+ 00000 ooo OOOOO0 +, 0000, O0OO0O?+000o0 oooo oo S ooo ocoooo+ oo oooooo ooooooo oso oooooooo0ooooo oooo & oo+ 0000000 oO oo ooooocoocoococooo+r+oococoooceooococecocoeoqococo Bonner zoologische Beitráge 55 (2006) (£981 “UY9IOJN) $198.12 $1.10P01]24 0 0 0 0 0 0 0 0 + 0 0 0 0 0 0 0 0 0 0 + 0 0 0 0 0 + 0 0 (161 “SMIP]A]) 19019141 S11OPD.DH 0 0 0 0 0 0 0 + 0 0 0 0 0 0 (0061 “ILLUSA) DIDMIDUMLAPPAD SILOPMIUO) 0 0 0 0 0 0 0 + 0 0 0 0 0 0 (0061 “[[LLI9A ) Paopaı]0 sLIOplyIUCQ 0 0 0 0 0 0 0 + 0 0 0 0 0 0 (1061 “TILLSA) DI01U1U SMOPN>DUO 0 0 0 0 0 0 0 + 0 0 0 0 0 0 (0061 “I[LL9A) D9/9D] SLIOpIyOUE 0 0 0 0 0 0 0 0 0 0 0 0 > 0 (1921 “QUULT) DJD]]9U1D]1Q SILOPIYIUO) 0 0 0 0 0 0 0 + 0 0 0 0 0 0 (1061 “I[LU9A) vioundoainy S1LOPIYIMO 0 0 + + 0 0 0 0 0 0 0 + (0161 “POOM[|[BUIS) 104-1J0G00Z HUIAO 0 0 0 0 0 0 0 0 + 0 0 0 0 0 0007 'esourdsg 29 VIVO 9D.ADUD.IM DUDO 0 0 + 0 0 0 0 0 + 0 0 0 0 0 LS61 “SODIB]A pxodun DUDO 0 0 0 0 0 0 0 0 0 0 0 + 0 0 LOOT “O1OJA 9 19][2Q2) “BIO 51/0391 poumon 0 0 0 0 0 0 0 0 0 0 0 + 0 0 9661 “S9P]LA Y PALO 10/0141 SIMOMYIIXIJA] 0 0 0 + 0 0 0 0 0 0 0 0 0 0 0L61 “SNIABJA] WZ snoiepy D/D9S sisopoydoT 0 0 0 0 0 0 0 0 + 0 0 0 0 0 SOOT “19][eqR) 9 esouIdsq BSG s7IUBOIUI SAYIUDAQI]IYUD 0 0 + + + 0 0 0 + 0 0 0 0 0 (SS61 “SODIB]A) pjozzpds puun.of 0 0 0 0 0 0 0 + 0 0 0 0 0 0 (8881 “undjıoH) D.1q9z s1iopojasdayy 0 0 0 0 0 =f 0 0 + 0 0 + 0 + (0681 YS19Y) 9] o4s s1.0pojasdAy 0 0 0 0 0 0 0 0 + 0 0 + 0 0 +L61 ‘Soysny Y snoieyy apyyın. siopojasdAy 0 0 + 0 0 0 0 0 0 0 0 0 + + (9€81 “Z1MUIS) DI91d $1.10pojas dy 0 0 0 + 0 0 0 0 + 0 0 + 0 0 1/6] “SODIBIA Mm S1.10po]9s dá 0 0 + 0 0 0 0 0 0 0 0 0 0 0 8661 “1LI031/] 29 BIDIBO) “OSODUOL]| $15U9/D] $1.10pOj2S dk y 0 0 0 0 0 0 0 0 0 0 0 + 0 0 9661 “S9PILA Y PALO Ipsoundso sLiopojasdAy] 0 0 0 0 0 0 0 0 + 0 0 + 0 + (1961 “SNDIBIA Y SNOIP]A) Modpg sisopojasdAyy 0 0 0 0 0 0 0 0 + 0 0 + 0 0 1961 “SHOR Y sno] PQ1.19D SL1OPoOJOSdAH 0 0 + 0 0 0 0 0 0 0 0 0 0 0 9007 “OSO2UOL] P PIDIBO "ZONdULWog UNODSOSUDY S1.10PO]dOY + 0 0 0 0 0 0 0 0 0 0 0 0 0 9761 ISUYPO snsoddpd snwmn907dojoff 0 0 0 0 0 0 0 0 + 0 0 + 0 0 7961 Se 2 SNILIAY SNUOS.LOU SNYIUD.AGDKAH 0 0 + + 0 0 0 0 0 0 0 0 0 0 LS6] “SMAIB]A avfady PXD]IPH 0 0 + 0 0 0 0 0 0 0 0 0 0 0 SSH] Mae pjnumu S1.LOPONOL) 0 0 0 0 0 0 0 0 + 0 0 0 0 + (1961 “SNIIBIA] Y Snap) DUPIS SLIOPOSso]H 0 0 0 0 0 0 0 0 + 0 0 0 + 0 (6/81 UBIDA) 1/9/9011 S110POSSO]L) 0 0 0 0 0 0 0 0 0 0 0 0 + 0 +v681 Y3IDG Ppunuu $110poj19L) Te 0 se 0 0 0 + 0 + + 0 0 0 + ( SISNOSOS SS O SS SO r 9 9059090995905 09 099995 O0 90599 59 0 90 9999 9 595 965 coococoocoooocoooooococcocoocoococoocoocoococoocoeoocjooqcoeocoeoocooqocqco oortrooococococoocortrrt+t oooocooocotrtococoocooocoocooocooocoeocoococooo+ocqocoe +O+00++T+00+O+fO0oO+O+S O+S € O+ OO+O+4O+FO+ O+4 O0O+ ++, 00O+0O000O+ MOROS ISSO: + OO SO Oo + CO: SO CO oc 00: + SC CO + OO Oo OC Oo oO + + CO OC So + Oo So OC SO O'S ocoooocooococoocoo+r+ococooooococootrtocoocooocoococeoococoeocertr+r1aqo0qo°o tio Cre OO OOF Oot Oo +O OOO + 007+ rss: Ss + OoCctoeoqocqcortooctrtooctoo + t+ooortortrtortortoococoococoocoorto0o0°o ooooocoocoocococooococococooooqococo otrotrocoocoocoortrtotrtrocoooort+o4+ + SO 5509955059 595959500909 +© oOOOOoOoOoooo9o oo 0 °o°0 0 00000 sonmads DUV SAS JAS MOV Sada SAN SV waa IM OAV V-O OH) 090 UI Biogeography of Brazilian Opisthobranchia Francisco J. GARCÍA et al. a a Species Cuthona genovae (O’ Donoghue, 1929) Cuthona georgiana (Pfeffer, 1886) Cuthona iris Edmunds & Just, 1983 Cuthona perca (Marcus, 1958) Cuthona pumilio Bergh, 1871 Cuthona rubra (Edmunds, 1964) Cuthona tina (Marcus, 1957) Dondice occidentalis (Engel, 1925) Dondice parguerensis Brandon & Cutress, 1985 Eubranchus conicla (Marcus, 1958) Eubranchus convenientis Ortea & Caballer, 2002 Eubranchus leopoldoi Caballer, Ortea & Espinosa, 2001 Eubranchus toledanoi Ortea & Caballer, 2002 Facelina agari Smallwood, 1910 Facelina coenda Marcus, 1958 Facelina goslingii A. E. Verrill, 1901 Facelina karouae (Marcus, 1957) Favorinus auritulus Marcus, 1955 Fiona pinnata (Eschscholtz, 1831) Flabellina dushia (Marcus & Marcus, 1963) Flabellina engeli Marcus & Marcus, 1968 Flabellina hamanni Gosliner, 1994 Flabellina marcusorum Gosliner & Kuzirian, 1990 Flabellina pallida (Verrill, 1900) Flabellina verta (Marcus, 1970) Glaucus atlanticus Forster, 1777 Godiva rubrolineata Edmunds, 1964 Learchis evelinae Edmunds & Just, 1983 Learchis poica Marcus & Marcus, 1960 Limenandra nodosa Haefelfinger & Stamm, 1958 Millereolidia ritmica (Ortea, Caballer & Espinosa, 2003) Nanuca sebastiani Marcus, 1957 Palisa kristenseni (Marcus & Marcus, 1963) Pauleo jubatus Millen & Hamann, 1992 Phidiana lynceus Bergh, 1867 Phidiana patagonica (d’Orbigny, 1836) Phidiana riosi Garcia & Troncoso, 2003 Piseinotecus divae Marcus, 1955 Pseudovermis salamandrops Marcus, 1953 Spurilla alba (Risbec, 1928) Spurilla neapolitana (delle Chiaje, 1844) Tenellia adspersa (Nordmann, 1845) Tenellia fuscata (Gould, 1870) Tenellia pallida (Alder & Hancock, 1855) Tergipes despectus (Johnston, 1835) Tergipes tergipes (Forskál, 1775) CC-CR ooo0oH+ 00 +00 CR-Cro CRo-O O-A 0 + 0 0 0 + 0 > 0 0 0 0 0 0 0 + 0 0 0 + 0 + 0 + 0 0 0 0 0 0 0 0 0 0 0 0 + 0 0 0 0 + 0 0 0 + 0 10) 0 0 + 0 0 0 0 0 0 + 0 0 0 =; 0 + 0 0 0 + 0 + 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 E ak SIE AS | va = o o > 194) oo0oooooo ACR SBB 0 0 0 0 0 0 0 0 0 0 + 0 0 0 0 0 0 0 0 + 0 0 aa 0 0 0 0 0 + 0 0 0 0 0 0 0 0 0 0 0 0 0 ~ + 0 0 0 0 0 0 0 0 + 0 0 0 0 + 0 0 + + 0 0 0 0 0 + 0 0 0 0 0 + 0 0 au 0 0 0 0 0 0 0 0 0 0 0 0 0 0 + 0 0 0 0 0 0 0 + + 0 0 0 0 0 0 0 0 + 0 0 + 0 0 0 0 + + 0 0 + 0 0 0 0 0 T 0 0 + 0 0 = 0 0 0 0 0 0 0 0 0 0 0 0 0 + oO ooooo N N Bonner zoologische Beitrage 55 (2006) C [> :J9JUIM 07< :Pwung I> YINOS VIO 19J2M €C-OC OWI, IT:UION ul Sur pomdp) L7-ST :Pwuung 91 :3upomdyy PITT ou ECASE -ISUUUUINS 9TST -LOJULAA 8T-L7 ‘owns 87-97 HOUMA, 67-L7 :Pwung (Do) Panye.ıadum], rE> AU NM AOIEM DMO.IBJURQqnS VOUT 9E< !PWUNS JUDLIND [zeig 1awumg IBM [EISEOJ USD Javea [Bud 9Uep y ymos IE<-EE> JUdLIND [zeig sjusa9 ZunpomdN OEM [EISEOY Loe [BUDD Suuepy YNOS LE-S’IE JUSLIN O [zeig LE-9€ Juano penojenbg yynos LE-9E JUOLIN IZeIE YON JOARY UOZLUV Juan o1ydo.ysoaH €E> JUDLIN, [IZeIg LION (00, ) Ayunes SIUIAINZ 081 08 [7071 001-09 09-05 tL Or (w) ypdap SALIA -J9US OLIFOTI DET=OS 06 I=S€ SIZO1I S8-0p (UM) JOYS [EJU9UHUO9 JO PLA Sob£7S0S 87 Sol’ STTSoET So€T-SoS I Sos 17808 So8 "Sol Soc Nor SWNT [eIseoJ (SAS) JI9YS uerpzeig wioynog (19S) sig uerpzeig ymos (YOY) u01891 soduiey-soyfoiqy (SIA) JOYS urifizeig UlO]seg (SAN) JOYS ueipizeigg ULOISBIYIION (SV) 31948 uozewv SUOISIY (8661) VANVAIN 9 OULSVD UO paseg ‘s}svoo UBIIZRIH SUOTE POJDPISUO) SEHE dy] JO SONYLIJ PUR SHUT *Z IQL Biogeography of Brazilian Opisthobranchia Francisco J. GARCÍA et al.: 222 Table 3. Number of species by order or suborder for each area considered. CC-CR CR-Cro CRo-O ABC WI BER O-A AS NBS EBS ACR SBB SBS Arg Cephalaspidea 50 39 32 14 88 19 8 9 26 11 14 19 14 10 Sacoglossa = 36 8 29 19 3 14 0 0 7 3 3 18 0 | Anaspidea 14 7 1] 10 14 8 | 0 10 6 6 8 | 0 Notaspidea 6 5 8 3 12 5 | | 10 4 3 5 | 2 Doridina 35 12 39 13 a2 11 0 2 17 2 23 36 0 14 Dendronotina 9 2 16 6 14 3 | 0 3 | 0 8 | 3 Arminina 3 | 4 | 2 0 | 0 0 0 | 2 0 0 Aeolidina 18 5 19 19 31 10 0 0 2 | 12 26 l 9 Total 173 79 178 85 250 72 12 12 75 28 62 122 18 30 Table 4. Percentage of species from each area (vertical column) present in other localities. CC-CR CR-CRo CRo-O ABC WI Ber O-A AS NBS EBS ABR SBB SBS ARG Total of species CC-CR 100 34 33 3 13 23 3 4 24 11 19 34 6 E 1.03 CR-CRo 75 100 66 30 78 39 9 8 34 15 25 of 13 8 19 CRo-O 51 29 100 24 69 21 6 5 28 12 20 23 4 3 178 ABC 69 28 31 100 80 34 2 2 27 18 28 44 5 2 85 WI 50 23 49 27 100 19 5 4 21 10 16 24 4 2 250 BER 61 43 51 40 67 100 6 6 32 17 26 26 8 4 72 O-A 67 58 83 17 100 33 100 17 50 23 42 50 33 42 12 AS 58 50 15 17 73 33 17 100 58 42 42 33 25 25 12 NBS JO 36 65 3 69 31 8 9 100 28 31 39 12 7 To EBS 68 43 75 54 86 43 1] 18 75 100 46 39 18 7 28 ABR 53 32 58 39 65 31 8 8 37 PA 100 61 16 5 62 SBB 48 24 36 30 48 16 5 3 24 9 31 100 10 7 122 SBS 56 56 44 22 61 33 22 17 50 28 56 67 100 28 18 ARG 17 17 IP * 6 17 9 14 9 14 6 9 26 14 100 35 Total of species 173 79 178 85 250 72 12 12 pl 28 62 122 18 33 Bonner zoologische Beiträge Band 55 (2006) Heft 3/4 Seiten 223-229 Bonn, November 2007 Paddle cilia on the cephalic sensory organs (CSOs) of Opisthobranchia (Mollusca: Gastropoda) — genuine structures or artefacts?* Katrin GOBBELER!) & Annette KLUSSMANN-KOLB!) Institute for Ecology, Evolution and Diversity - Phylogeny and Systematics, Johann Wolfgang Goethe-University, Frankfurt am Main, Germany *Paper presented to the 2nd International Workshop on Opisthobranchia, ZFMK, Bonn, Germany, September 20th to 22nd, 2006 Abstract. Paddle cilia are characterised by a curved axoneme at the distal end enclosed by the ciliary membrane. They have been described in numerous different marine invertebrates including one species of Opisthobranchia. There is still controversy about the nature of paddle cilia. Various considerations have been made concerning their function regarding them as genuine structures, whereas other authors claimed paddle cilia to be artefacts. The current study focuses on in- vestigating paddle cilia on the cephalic sensory organs (CSOs) of different opisthobranch species in order to present mo- re data and perhaps clarify the paddle cilia discussion. For this purpose specimens were fixed using two different me- thods. One method comprises an isoosmotic fixation solution which did not induce paddle cilia formation in bivalve lar- vae (SHORT & TAMM 1991). The other method utilises a hypoosmotic fixation solution. Using scanning electron micros- copy paddle cilia can be detected in all investigated species. The abundance of paddle cilia was lower in the majority of specimens fixed with the isoosmotic solution which led us to conclude that paddle cilia are indeed artefacts. Keywords. Scanning Electron Microscopy, isoosmotic fixation. 1. INTRODUCTION Paddle cilia or discocilia were first described by TAMARIN et al. (1974) as cilia whose cylindrical stem curves back upon itself and forms a 360° loop which is completely en- veloped by the ciliary membrane. Since then paddle cil- ia have been discovered in many different marine inver- tebrates (EHLERS € EHLERS 1978; HEIMLER 1978; BONE et al. 1982; MATERA & Davis 1982; CAMPOS & MANN 1988). Various considerations were made concerning the func- tion of this special structure. TAMARIN et al. (1974) pro- posed that paddle cilia function as microscopic spatulas for the application of adhesive plaque material to substrate surfaces. HEIMLER (1978) supposed that they are locomo- tory organs since the disc shaped heads can be interpret- ed as enlargements of the surface to improve the efficien- cy of the ciliary beat. Thus paddle cilia also influenced the nutrition of the larvae investigated in his study. CAM- POS & MANN (1988) detected paddle cilia on the velum of bivalve larvae of two different species. They regarded them as genuine structures and favoured a locomotory and/or chemosensory function although they were not able to prove this assumption, since the occurrence of paddle cilia did not enhance the rate of movement in the exam- ined species (CAMPOS & MANN 1988). So far, only one study deals with the presence of paddle cilia in Opisthobranchia. The investigation of Pleurobran- chaea californica revealed that paddle cilia solely occurred in chemosensitive regions of this species (Davis & MAT- ERA 1982). This fact led Davis & MATERA (1982) to the conclusion that paddle cilia are most likely chemorecep- tors. The dilatations were expected to enlarge the mem- brane surface area, increasing the opportunity for inter- action with chemical substances. Other authors supposed paddle cilia to be artefacts result- ing from osmotic stress, increased temperature, non-phys- iological conditions or fixation (EHLERS & EHLERS 1978; BONE et al. 1982; NIELSEN 1987; SHORT & TAMM 1991; DEINER et al. 1993). SHORT & Tamm (1991) discovered that the occurrence of paddle cilia is associated with the osmolarity of the fixa- tion solution. After the application of an isoosmotic fix- ation solution with 50% seawater no paddle cilia could be detected in their bivalve larvae, whereas other hypoosmot- ic fixation solutions induced the formation of paddle cil- ia. However, the occurrence of paddle cilia was always restricted to certain areas and never affected all cilia of the larvae. NIELSEN (1987) argued that different cilia - even on the same organism - are not equally sensitive to stress 224 Kathrin GÓBBELER & Annette KLUSSMANN-KOLB: Paddle cilia in Opisthobranchia and some cilia are indeed difficult to fix in a normal shape. DEINER et al. (1993) reported that paddle cilia induced by hypotonic solutions usually regain their normal appear- ance if specimens are returned to isotonic solutions. The current study investigates the cilia on the cephalic sen- sory organs (CSOs) of different Opisthobranchia. CSOs are special structures in the head region of sea slugs, which are sensitive to several stimuli. The purpose of this inves- tigation is to detect paddle cilia and clarify the question whether they are genuine structures or artefacts. 2. MATERIAL AND METHODS Specimens of Acteon tornatilis (Linnaeus, 1758) (Acteonoidea), Aeolidiella glauca (Alder and Hancock, 1845) (Nudibranchia), Aplysia punctata Cuvier, 1803 (Anaspidea) and Berthella plumula (Montagu, 1803) (Pleurobranchoidea) were collected in the intertidal at Roscoff and Saint Michel-en-Greve (Brittany, France). Haminoea hydatis (Linnaeus, 1758) (Cephalaspidea) was obtained from a laboratory culture at the J.W.G.-Uni- versity of Frankfurt. We applied the isoosmotic fixation method of SHORT & TAMM (1991) and a hypoosmotic fixation solution in or- der to clarify the nature of paddle cilia on the cephalic sen- sory organs of different Opisthobranchia. The specimens were anaesthetised by an injection of 7% MgCl, in the foot and the CSOs were abscised. Two different methods have been applied: CSOs of investigated species were fixed in 2.5% glutaraldehyde, 1% paraformaldehyde in phosphate buffer, pH 7.2 — 7.4 at room temperature (method 1). The osmolarity of this solution was approximately 250 mOsmols less than the osmolarity of the isoosmotic fix- ation solution and the artificial seawater in which the spec- imens were cultivated (determined with the help of Knauer Semi-Micro Osmometer). Additional CSOs of Acteon tor- natilis, Aplvsia punctata, Berthella plumula and Haminoea hydatis were fixed in an isoosmotic solution after SHORT and TAMM (1991) containing 2.5% glutaraldehyde, 0.13 M NaCl and 50% seawater, pH 7.2 — 7.4 at room temper- ature (method 2). The investigated CSOs and number of Fig. 1. Photographs of the investigated species and schematic drawings of their cephalic sensory organs (CSOs). A Aplysia puncta- ta. B Berthella plumula. C Acteon tornatilis. al anterior lobe, e eye, gr groove, m mantle, ot oral tentacle, ov oral veil, pl po- sterior lobe, rh rhinophore. Bonner zoologische Beiträge 55 (2006) 225 replicates were as follows (method 1/ method 2): Acteon tornatilis head shield (4/1), Aeolidiella glauca rhinophore (2/-), oral tentacle (2/-), Aplysia punctata rhinophore (4/2), oral tentacle (4/2), Berthella plumula rhinophore (6/2), oral veil (3/1), Haminoea hydatis head shield (4/1), lip or- gan (4/2), Hancock ’s organ (4/2). For SEM, fixed CSOs were dehydrated through a graded acetone series, critical point dried (BAL-TEC, CPD 030), sputtered with gold (Sputter-Coater, Agar Scientific) and examined with a Hitachi S4500 SEM. Photographs were taken with DISS — Digital Image Scanning System (Point Electronic). : Sizes of cilia were determined by average over measure- ment of at least ten cilia. 3. RESULTS The investigated species exhibit several different types of CSOs (Figs. | and 2), like rhinophore, oral tentacle, oral veil, head shield, lip organ and Hancock’s organ. The rhinophores and oral tentacles of Aplysia punctata (Fig. 1A) possess dark pigmented grooves. These grooves con- tain tufts of cilia. A total of four rhinophores fixed with method | were investigated. One of these exhibits paddle Fig. 2. Photographs of the investigated species and schematic drawings of their CSOs. A Aeolidiella glauca. B Haminoea hy- datis — scheme: dorsal (left) and lateral (right) view of the CSOs. e eye, ho Hancock’s organ, hs head shield, lo lip organ, ot oral tentacle, rh rhinophore. cilia (Fig. 3A), whereas the other reveal cilia without pad- dle shaped ends (Fig. 3B). The rhinophore exhibiting pad- dle cilia has a total length of 3 mm, whereas the other in- vestigated rhinophores without paddle cilia are 5 mm long. The rhinophores fixed with method 2 show both cilia with and without paddle-shaped ends. The paddle cilia are al- ways shorter (-12 um) than the other cilia (~16 um). The oral tentacles fixed with method | reveal mainly pad- dle cilia (Fig. 3C) with a length of ~12 um. Cilia without paddles are ~16 um long (Fig. 3D). The oral tentacles fixed with method 2 possess mainly cilia without paddle- shaped ends (Fig. 3E), the fewer paddle cilia occur in sin- gle tufts. Berthella plumula possesses rolled rhinophores and an oral veil with lateral grooves (Fig. 1B). Inside the rhinophores tufts of cilia are arranged regularly from the tip to the base of this CSO. These cilia have straight tips in rhinophores (Fig. 3F) fixed with method 1, whereas they reveal pad- dle-shaped ends in those preserved with method 2. Out- side the rhinophores mainly paddle cilia are found (Fig. 3G) irrespective of fixation methods. Inside the lateral grooves of the oral veils fixed with method 1 tufts of cilia with straight tips were found (Fig. 3H). On the margin of the grooves as well as on the dor- sal and ventral side of the oral veil paddle cilia are pre- dominant (Fig. 31). The isoosmotically fixed oral veil al- so reveals paddle cilia inside the grooves. The head shield of Haminoea hydatis possesses postero- lateral flattened appendages. The lip organ is located an- teriorly beneath the head shield and continues into the Hancock’s organ. The arrangement of the three CSOs is shown in Fig. 2B. The lip organ reveals tufts of paddle cilia as well as tufts of cilia with straight tips and even both types of cilia in one tuft (Fig. 4A) after fixation with method 1. The Hancock’s organ also exhibits cilia with (Fig. 4B) and without paddle-shaped ends when fixed with method 1. In contrast to this, both CSOs showed no pad- dle cilia after fixation with method 2 (Fig. 4C and D). The head shield is densely covered by cilia on the dorsal side. No paddle cilia could be detected after application of both fixation methods. Nevertheless fixation with method 1 re- sulted in cilia whose tips are curved to the axoneme (Fig. 4E), whereas fixation with method 2 revealed cilia with straight tips (Fig. 4F). The head shield of Acteon tornatilis is completely cov- ered by cilia. It is divided into a pair of antero-lateral lobes and a pair of postero-lateral lobes (Fig. IC). A groove runs along the front and lateral side of the anterior lobe. The specimens fixed with method | show numerous cilia with slightly, and some cilia with extremely, swollen ends in- 226 Kathrin GÓBBELER & Annette KLUSSMANN-KOLB: Paddle cilia in Opisthobranchia Fig. 3. Scanning electron microscopy of sensory regions of the CSOs of Aplysia punctata and Berthella plumula fixed with me- thod 1 (except E). A rhinophore of Aplysia punctata — paddle cilia (arrows) inside the groove of a small individual. B rhinophore of Aplysia punctata — cilia inside the groove. C oral tentacle of Aplvsia punctata — paddle cilia inside the groove. D oral tentacle of Aplysia punctata — tuft of cilia without paddle-shaped ends inside the groove. E oral tentacle of Aplysia punctata fixed with me- thod 2 — cilia inside the groove. F rhinophore of Berthella plumula — cilia on the inside. G rhinophore of Berthella plumula — paddle cilia on the tip. H oral veil of Berthella plumula — cilia inside the lateral groove. I oral veil of Berthella plumula — paddle cilia on the outside, beneath the groove. Bonner zoologische Beiträge 55 (2006) 227 side the groove (Fig. 5A). In addition, clusters of paddle cilia could be detected in different parts of the groove (Fig. 5B). These paddle cilia are longer than the surrounding cilia without paddle-like ends. In the region of the mouth below the groove numerous paddle cilia are found (Fig. 5C). These paddle cilia could not be detected in the spec- imen that was fixed with the isoosmotic solution (method 2 / Fig. 5D). Moreover, no paddle cilia occur inside the groove of this specimen. Fig. 4. Scanning electron microscopy of sensory regions of the CSOs of Haminoea hydatis. A lip organ fixed with method I — paddle cilia and cilia with straight tips in one tuft. B Hancock's organ fixed with method | — tuft of paddle cilia. C lip organ fi- xed with method 2 — tuft of cilia with straight tips. D Hancock’s organ — tuft of cilia with straight tips. E head shield fixed with method 1 — cilia with curved tips. F head shield fixed with me- thod 2 — cilia with straight tips. The rhinophores and oral tentacles of Aeolidiella glauca are solid structures (Fig. 2A). Both were only fixed with method 1. Paddle-like structures could be detected on the tip of the rhinophores (Fig. SE). Some of these paddle- like structures are not, as usually, found on the distal end of the cilia, but occur subapically. Paddle cilia are also present laterally on the oral tentacles (Fig. SF). In addi- tion, the cilia on the tip of the oral tentacles reveal swollen ends. 4. DISCUSSION All investigated taxa and organs exhibit cilia with and without paddle-shaped ends. This fact might on the one hand support the assumption that paddle cilia are genuine structures. On the other hand it might be concluded that cilia with variable morphological features react different- ly to special fixation solutions. The only specimen of Aplysia punctata exhibiting paddle cilia inside the groove of the rhinophore when fixed with method 1 was distinctly smaller than the other ones. This might possibly indicate that juveniles possess paddle cil- ia that get stretched during growth. It is also possible that this specimen was not handled with enough care which might be a reason for paddle cilia formation according to NIELSEN (1987). Thus the natural condition would be dis- played by the straight tips. EHLERS & EHLERS (1978) al- so discovered that paddle cilia normally do not appear in the same quantity in all the individuals when several an- imals are fixed. Since paddle cilia are predominant inside the groove of the oral tentacle fixed with method 1, the cilia in this CSO seem to be more sensitive to paddle cil- ia formation than cilia in the rhinophore. The length of the cilia is equal in rhinophore and oral tentacle and cilia with straight tips are 4 um longer than paddle cilia. This dif- ference might be caused by rolling in of the tips of straight cilia resulting in paddle cilia formation. Thus paddle cil- ia would not be a genuine type of cilia but an artefact. The isoosmotic fixed specimens of both CSOs of Aplysia punc- tata reveal paddle cilia as well as cilia without paddle-like structures. This could indicate that paddle cilia are a dis- tinct type of cilia. On the other side, it could also mean that SHORT & Tamm's (1991) fixation method does not work as efficiently for opisthobranchs as it did for bi- valves. The specimens could have also been exposed to another form of stress. Perhaps paddle cilia formation might even be induced by collecting or anaesthetisation of the animals. The findings for Berthella plumula are con- trary to the results of SHORT & Tamm (1991), because isoosmotic fixation results in additional formation of pad- dle cilia, instead of decreasing the amount of these cilia. These results nevertheless reveal that cilia react different- ly to the application of variable fixation solutions, a fact 228 Kathrin GOBBELER & Annette KLUSSMANN-KOLB: Paddle cilia in Opisthobranchia SEEN, Fig. 5. that strengthens the assumption of paddle cilia being arte- facts. The tufts of paddle cilia found inside the groove of the head shield of Acteon tornatilis fixed with method 1 are longer than the surrounding cilia. BONE et al. (1982) in- vestigated the nature of cilia in the endostyl of a tunicate. They argued that paddle cilia are artefacts and particular- ly prone to occur in long cilia. This seems to be true for Acteon tornatilis. The lip and Hancock's organ of Haminoea hydatis fixed with method | reveal cilia with and without paddle-like structures even in the same tuft. EHLERS & EHLERS (1978) described a similar situation for turbellarians. Not all the cilia of a given cell show paddle-like ends in these species, unmodified cilia also occur (EHLERS & EHLERS 1978). Scanning electron microscopy of sensory regions of the CSOs of Acteon tornatilis and Aeolidiella glauca fixed with me- thod 1 (except D). A Acteon tornatilis — cilia with dilated tips inside the groove. B Acteon tornatilis — tuft of paddle cilia inside the groove. € Acteon tornatilis — paddle cilia in the region of the mouth. D Acteon tornatilis — cilia in the region of the mouth. E rhinophore of Aeolidiella glauca — paddle cilia (arrows) on the tip. F oral tentacle of 4eolidiella glauca — paddle cilia (arrows) la- terally. Morphological differences are apparently not the only rea- son for the formation of paddle cilia because it seems un- likely that cilia of one single tuft exhibit variable morpho- logical features. Cilia with tips curved to the axonemes found on the hypoosmotically fixed head shield of Haminoea hydatis might be a developmental stage of pad- dle cilia since swelling of the ciliary membrane would lead to fully developed paddle cilia. The fact that paddle cilia could not be detected on spec- imens of Acteon tornatilis and Haminoea hydatis fixed with method 2 corresponds to the findings of SHORT & TAMM (1991) and further indicates that paddle cilia are in- deed artefacts. The paddle-like structures found on the rhinophores and oral tentacles of Aeolidiella glauca fixed with method 1 Bonner zoologische Beiträge 55 (2006) 229 are partly not located at the distal end of the cilia but are situated subapically. In addition numerous cilia with swollen ends were found on the oral tentacles. HEIMLER (1978) described these special forms as different types of paddle cilia. He distinguished three different types of pad- dle cilia. Type 1 has swollen, bulblike heads with a cen- tral straight axoneme. Type 2 paddle cilia are character- ized by heads with a curved lateral axoneme and in type 3 paddle cilia the axoneme forms a loop. On the CSOs of Aeolidiella glauca type 1 and 2 paddle cilia could be de- tected whereas Aplysia punctata and Berthella plumula re- veal type 3. Furthermore, HEIMLER (1978) supposed that these types represent different developmental stages of these cilia. Thus 4eolidiella glauca does not display the fully developed stage of paddle cilia. MATERA & Davis (1982) investigated the distribution of paddle cilia on the opisthobranch Pleurobranchaea cali- fornica. Since paddle cilia only occurred in areas known to mediate chemoreception, they assumed them to be chemoreceptors. The current study shows that there are lots of other cilia with straight tips on the chemorecep- tive structures of different Opisthobranchia, especially when using an isoosmotic fixation solution. This obser- vation does not correspond to the conclusions of MATERA & Davis (1982), but indicates that paddle cilia might rather be artefacts. EHLERS & EHLERS (1978) equally detected paddle-like structures only in sensory cells of the investigated Turbellaria. But they demonstrated that the number of pad- dles of sensory cilia increases in proportion to increasing osmolarity of the fixation solution and increasing temper- ature during fixation and concluded that paddle cilia are artefacts. Altogether the current study enforces the assumption that paddle cilia are artefacts. Especially the results for Acteon tornatilis and Haminoea hydatis confirm that paddle cil- ia formation can be induced by the application of differ- ent fixation solutions. Nevertheless the results for Aplysia punctata and Berthella plumula leave the possibility that beyond being artefacts paddle cilia might occur natural- ly in a lesser amount. This speculation needs to be proven in living, unstressed animals. Acknowledgements. We would like to thank Marion Basoglu for introduction into dehydration and critical point drying, Man- fred Ruppel for SEM, Sid Staubach for the specimens of Haminoea hydatis and Alen Kristof for the picture of Haminoea hydatis. This work was financially supported by the German Re- search Foundation (KL 1303/3-1). REFERENCES BONE, Q., RYAN, K.P. & PULSFORD, A. 1982. The nature of com- plex discocilia in the endostyle of Ciona (Tunicata: Ascidi- acea). Mikroskopie 39: 149-153. Campos, B. & MANN, R. 1988. Discocilia and paddle cilia in the larvae of Mulinia lateralis and Spisula solidissima (Mollus- ca: Bivalvia). Biological Bulletin 175: 343-348. Davis, W.J. & MATERA, E.M. 1982. Chemoreception in gastro- pod molluscs: electron microscopy of putative receptor cells. Journal of Neurobiology 13 (1): 79-84. DEINER, M., TAMM, S.L. & Tamm, S. 1993. Mechanical proper- ties of ciliary axonemes and membranes as shown by paddle cilia. Journal of Cell Science 104: 1251-1262. EHLERS, U. & EHLERS, B. 1978. Paddle cilia and discocilia — gen- uine structures? Observations on cilia of sensory cells in ma- rine Turbellaria. Cell and Tissue Research 192: 489-501. HEIMLER, W. 1978. Discocilia — a new type of kinocilia in the larvae of Lanice conchilega (Polychaeta, Terebellomorpha). Cell and Tissue Research 187: 271-280. Matera, E.M. & Davis, W.J. 1982. Paddle cilia (discocilia) in chemosensitive structures of the gastropod mollusk Pleuro- branchaea californica, Cell and Tissue Research 222: 25—40. NIELSEN, C. 1987. Structure and function of metazoan ciliary bands and their phylogenetic significance. Acta Zoologica (Stockholm) 68: 205-262. SHORT, G. & Tamm, S.L. 1991. On the nature of paddle cilia and discocilia. Biological Bulletin 180: 466-474. TAMARIN, A., Lewis, P. & Askey, J. 1974. Specialized cilia of the byssus attachment plaque forming region in Mytilus cal- ifornianus. Journal of Morphology 142: 321-328. Authors addresses: Katrin GOBBELER (corresponding aut- hor), Institute of Ecology, Evolution and Diversity — Phy- logeny and Systematics, Johann Wolfgang Goethe-Univer- sity, Siesmayerstr. 70, 60054 Frankfurt/Main, Germany, Email: goebbeler(@bio.uni-frankfurt.de; Annette KLuss- MANN-KOLB, Institute of Ecology, Evolution and Diver- sity — Phylogeny and Systematics, Johann Wolfgang Goe- the-University, Siesmayerstr. 70, 60054 Frankfurt/Main, Germany, Email: klussmann-kolb@bio.uni-frankfurt.de. | Bonner zoologische Beitrage Band 55 (2006) Heft 3/4 Seiten 231—254 Bonn, November 2007 Preliminary study on molecular phylogeny of Sacoglossa and a compilation of their food organisms* Katharina HANDELER!) & Heike WAGELE!)-?) Institut für Evolutionsbiologie und Ökologie, Rheinische Friedrich-Wilhelms-Universitát, Bonn, Germany 2)Zoologisches Forschungsmuseum Alexander Koenig, Bonn, Germany *Paper presented to the 2nd International Workshop on Opisthobranchia, ZFMK, Bonn, Germany, September 20th to 22nd, 2006 Abstract. The first molecular analysis of the Sacoglossa based on the 16Sr DNA gene (partial), with 39 species and 59 specimens analysed, is presented. A saturation of substitution is observed, as well as conflict in the data concerning cer- tain taxa. Therefore the phylogenetic relationship presented here has to be considered as preliminary. Many results are congruent with the analysis of JENSEN (1996) based on morphological characters. Plakobranchacea are monophyletic. Elysiella pusilla is nested within the genus E/ysia and therefore synonymy of Elysiella with Elysia confirmed. The po- sition of Plakobranchus remains unclear, but a closer affinity to Elysia than to Thuridilla seems most likely. Major dif- ferences lie in the paraphyly of the Oxynoacea, but only few members of this taxon have been included. A compilation of available data on food algae is presented and the data are discussed under the light of the phylogenetic relationship presented here. Keywords. Opisthobranchia, molecular systematics, SplitsTree, Chlorophyta, evolution, 16S rDNA. 1. INTRODUCTION Sacoglossa is a small group of Opisthobranchia with about 250 to 300 described species (JENSEN 1997a). Animals are small and species larger than 30 mm are rare. The mono- phyly of this group is well supported (JENSEN 1996) al- though their phylogenetic position relating to other groups of Opisthobranchia is unresolved (DAYRAT et al. 2001; DAYRAT & TILLIER 2002; GRANDE et al. 2004; Von- NEMANN et al. 2005; WAGELE & KLUSSMANN-KOLB 2005). The first and only phylogeny of Sacoglossa based on mor- phological data was presented by JENSEN (1996) applying Hennigian principles. Her analysis was performed on genus level, except for the genus Elysia with four includ- ed species. One of her results was a differentiation into two major clades, the shelled Oxynoacea and the shell- less Plakobranchacea. The latter comprise the monophyle- tic Plakobranchoidea and the monophyletic Limapon- tioidea. Cylindrobulla appeared as the most basal taxon within her analysis. GOSLINER (1995) performed a mor- phologically based analysis concentrating on the genus Thuridilla. An analysis of the family Plakobranchidae based on a genetic marker (COI) was published lately (Bass & KARL 2006). No further phylogenetic analyses have been performed up to now. Sacoglossa is feeding nearly exclusively on siphonalean or siphonocladalean Chlorophyta. They penetrate the cell wall with their uniseriate radula and suck out the cell sap. Noteworthy is the possibility of several sacoglossans to incorporate the chloroplasts from their food intracellular in their digestive gland and partially perform photosyn- thesis. This was first recorded by KAwAGUTI & YAMASU (1965) and most recently analysed by EVERTSEN et al. (2007) (for a review see RUMPHO et al. 2000). For sever- al sacoglossans reception of polysaccharides and amino acides from these chloroplasts is documented (TRENCH 1973; WILLIAMS & WALKER 1999 after Greene & Musca- tine 1972). In addition, many members of the Sacoglos- sa are able to incorporate secondary metabolites from their food and use these as a chemical defense (CIMINO & GHIS- ELIN 1998; CIMINO et al. 1999; GAVAGNIN & FONTANA 2000; MARÍN & Ros 2004; CIMINO & GAVAGNIN 2006). JENSEN (1997a: 313) was the first who discussed a possi- ble correlation between the evolution of morphological characters of the Sacoglossa and the possible evolution of the food plants. She argued that more investigations are needed: “Rigorous application of cladistic methodology in coevolution analysis requires fully resolved phyloge- nies of the ‘hosts’ as well as the "associates'.” Intention ho eS) ho of this study was to fill this gap by analysing the phyloge- ny of Sacoglossa with molecular data and by including many more species. New available data on food organ- isms in Sacoglossa are presented, which might help to elaborate co-evolutionary processes. 2. MATERIALS AND METHODS 2.1. Biological material 58 specimens including 38 species of sacoglossans were used for the phylogenetic reconstruction. Locality, date of collection and accession numbers are indicated in Table 1. According to the results of JENSEN (1996) and unpub- lished data of the senior author, Cylindrobulla beauii (Cylindrobullidae, Opisthobranchia) was chosen as out- group species. Cylindrobulla and Sacoglossa share synapomorphies (presence of a shell, adductor muscle, and a lamellate gill (JENSEN 1996)), but also lack synapomor- phies of the Sacoglossa (e.g. ascus), and show plesiomor- phic characters (e.g. triangular shaped teeth, large shell). 2.2. DNA extraction, amplification and sequencing DNA was extracted from alcohol-preserved specimens by means of NucleoSpin™ Tissue-Kit by Machery-Nagel or Dneasy* Blood and Tissue Kit by Quiagen, guided by the enclosed protocol. A segment of the 16Sr DNA gene was amplified with primer pairs, LR-J-12887 5°-GGA GCT CCG GTT TGA ACT CAG ATC-3* and LR-N-13398 5*- CGG CCG CCT GTT TAT CAA AAA CAT-3*. Amplifi- cation reactions (SOul) consisted of 39.15u1 ddH,0, Sul 10X PCR Buffer without MgCl, (Fermentas), 4ul MgCl, (25mM), 0.15ul Taq-Polymerase (SU/ul), 0.4ul of each Primer (10pmol/ul) and 0.5ul DNA. The PCR was car- ried out in the Gene Amp PCR System 9600 by Perkin Elmer* under following conditions: 95°C for 240s, fol- lowed by 9 Touch-down-cylces of 45s at 94°C, 45s at S6(- 1)°C, 90s at 72°C, followed by 25 amplification-cycles of 45s at 94°C, 45s at 48°C, 90s at 72°C and a final exten- sion at 72°C for 10min followed by cooling down to 4°C. Amplicons were purified by means of NucleoSpin* Ex- trat II by Machery-Nagel, guided by the enclosed proto- col. The mass of the amplicons was estimated by compar- ing ethidium bromide staining intensity of Sul of each pu- rified reaction. Cycle sequencing reactions (Cycle Sequencing Kit BigDye* Terminator v1.1 by Applied Biosystems) were carried out in the Gene Amp PCR System 9600 by Perkin Elmer® under following conditions: 96°C for 120s fol- lowed by 15 cycles of 10s at 96°C, 5s at 50°C, 150s at Katharina HANDELER & Heike WAGELE: Phylogeny of Sacoglossa 60°C followed by cooling down to 4°C. Reaction pro- ducts were purified by adding 40u1 water containing Dex- tranblau, 5ul natriumacetate (3M, pH4.6) and 125ul pure ethanol, centrifuging for 15min, taking supernatant, adding 50011 fresh 70% ethanol, centrifuging for 5min, taking supernatant, and were dried at 37°C for 30-40min. The fluorescently labeled products were size sorted and visualized using an ABI PrismTM 377 DNA Sequencer by Applied Biosystems. The sequences were extracted with Sequencing Analysis 3.0. The sequences of the re- verse primer were complemented with Factura 2.0.1 and sequences of both strands were compared and synchro- nised with Sequence Navigator. The gene fragments were aligned by applying Muscle3.7 (EDGAR 2004a; EDGAR 2004b) and the resulting alignment was verified by eye. Length of the alignment was 478 base pairs. 2.3 Phylogentic reconstruction A Bayesian analysis was performed with MrBayes 3.1.2 (HUELSENBECK & RONQUIST 2001; HUELSENBECK & BOLLBACK 2001). Maximum Parsimony analysis, Neigh- bor-joining analysis and Maximum likelihood analysis have been performed with PAUP* 4.0b10. Modeltest 3.7 (POSADA & BUCKLEY 2004) selected as best-fit model the GTR+G+I-model (AIC). Therefore, neighbour-joining analysis was conducted on a GTR-model and Maximum Likelihood analysis and Bayesian analysis on a GTR+G+I-model. Saturation of substitutions was tested against patristic distances as well as against distances ob- tained with the GTR-model. A posteriori statistical analy- sis could not be performed due to long calculation time of bootstrapping methods. For visualizing conflict in the data set, which seemed obvious after the analyses, Splits- Tree4 (HUSON & BRYANT 2006) was applied. A Neighbour- Net analysis was performed applying the K2P-model. 3. RESULTS Saturation in substitutions of the partial 16Sr DNA ob- tained by using the GTR model is shown in Figure 1. The graph approaches a plateau indicating a saturation of sub- stitutions. When splitting the data set into subgroups (fam- ily or genera), saturation was less evident for most groups, but not e.g. the genus Elysia or Thuridilla. Substitution rates were also less saturated, when using p-distances (not shown here). Bonner zoologische Beiträge 55 (2006) 233 3.1. Phylogenetic reconstruction A maximum parsimony analysis produced 81 shortest topologies with a length of 1401 steps. The 50%-Major- ity-Rule tree is shown in Figure 2. Within the monophyle- tic Plakobranchacea, the family Polybranchiidae and the polybranchiid genus Cyerce are also monophyletic. The monophyletic genus Ercolania appears as the sistertaxon to the Polybranchiidae. The families Plakobranchidae and Limapontiidae are not monophyletic, the genera Elysia and Thuridilla are also paraphyletic. Plakobranchus groups as sistertaxon of several Elysia species. The shelled Oxynoacea are basal. They are paraphyletic as well as the family Juliidae and Oxynoidae. The position of Oxynoe viridis 1s unresolved. In the Neighbor-joining analysis (not presented here), Plakobranchacea are monophyletic, as well as the family Polybranchiidae and the genus Thuridilla. Bosellia mimet- ica (Boselliidae) groups in the family Plakobranchidae, rendering this family paraphyletic. Limapontiidae is pa- raphyletic again, since Ercolania and Stiliger do not group together. The genus E/ysia again comprises Elysiella and Plakobranchus and is polyphyletic in this analysis. Plakobranchus is sistertaxon to the same E/ysia-complex as in the maximum parsiomony analysis. Similar to the maximum parsimony analysis, Elysia benettae is the sis- tertaxon to Elysiella pusilla. Monophyletic Ercolania ap- pears as the most basal genus of the Plakobranchacea. p-distances 0,9 0,8 0,7 0,6 0,5 0,4 0,3 0,2 0,1 4 0 04 0,2 0,3 0,4 Fig. 1. Oxynoacea are basal and paraphyletic. The families Juli- idae and Oxynoidae again are not monophyletic. Similar as in the MP analysis, Tamanovalva is the sistertaxon to the Plakobranchacea. The position of Oxynoe viridis is un- clear. Two identical trees were found in the Maximum-likeli- hood analysis; the resulting tree is shown in Figure 3. Plakobranchacea is monophyletic, as well as the families Plakobranchidae and Polybranchiidae. Boselliidae is sis- tertaxon to the Plakobranchidae. The genus Thuridilla is monophyletic. Elysiella pusilla with Elysia benettae as sis- tertaxon groups in the genus Elysia, rendering this genus paraphyletic. In this analysis Plakobranchus ocellatus 1s the sistertaxon to all Elysia-species (including Elysiella) and monophyletic Thuridilla to this Plakobranchus/Elysia clade. Limapontiidae again ıs paraphyletic. Contrary to the other analyses, Ercolania is not monophyletic. The two species represent the most basal Plakobranchacea. The Oxynoacea and Oxynoidae are paraphyletic again, but contrary to all other analyses, the Juliidae with the gen- era Julia and Tamanovalva represent a monophyletic group. The position of Oxynoe viridis is unresolved. The cladogram based on the analysis with MrBayes is shown in Figure 4. Plakobranchacea is monophyletic, but resolution within that taxon is low. Elysia (without E. benettae), Elysiella/Elysia bennettae complex and Plako- branchus form a monophyletic clade, but their position is 0,5 0,6 0,7 0,8 0,9 d-Distances GTR Saturation of substitution of the 16Sr DNA gene (partial) used in this analysis. The straight line is the bisector. Distances are calculated as patristic distances (y-axis) against d-distances calculated by applying the GTR model (x-axis). 234 Katharina HÁNDELER € Heike WÄGELE: Phylogeny of Sacoglossa 100 - Cyerce spec. 2 Cyerce spec. 1 Cyerce nigricans Cyerce antillensis Cyerce edmundsi Polybranchia cf. orientalis Mourgona osumi Ercolania spec. 1 Ercolania spec. 2 Bosellia mimetica 1 Bosellia mimetica 2 Bosellia mimetica 3 Placida dendrítica 1 Placida dendfritica 2 Placida dendritica 3 Thuridilla carlsoni 1 Thuridilla carisoni 2 Thundilla kathae Thuridilla vatae Thuridilla albopustulosa Thuridilla hoffae Polybranchiidae Limapontiidae Boselliidae Limapontiidae Plakobranchidae Thundilla ratna 2 Thuridilla bayeri Thuridilla lineolata Thuridilla hopei 1 Thuridilla hopei 2 Elysia timida 1 Elysia timida 2 Elysia timida 3 Elysia obtusa Elysia tomentosa 1 Elysia tomentosa 2 Elysia macnaei Elysia spec.5 Elysia spec.1 Elysia ornata 1 Elysia ornata 2 Elysia ornata 3 Elysia amakusana Plakobranchus ocellatus 1 Plakobranchus ocellatus 2 Elysiella pusilla 1 Elysiella pusilla 2 Elysiella pusilla 3 Elysiella pusilla 4 Elysia benettae Elysia viridis 1 Plakobranchidas Plakobranchidae = Plakobranchidae Fig. 2. Elysia viridis 2 Elysia vindis 3 Tamanovalva limax Julia exquisita Volvatella viridis Lobiger viridis 1 Lobiger viridis 2 ——— Lobiger viridis 3 Cylindrobulla beauii Oxynoe viridis Sacoglossa; 16Sr DNA gene: Maximum Parsimony analysis, 50% Majority Rule consensus tree. Thuridilla gracilis Thuridilla ratna 1 Plakobranchidae ] Juliidae ] Juliidae ] Volvatellidae Oxynoidae ] Oxynoidas Bonner zoologische Beiträge 55 (2006) 235 unresolved. This complex and two Thuridilla complexes (representing all Plakobranchidae) and Bosellia form the next higher unresolved monophyletic clade. Polybranchiidae and within that family the genus Cyerce, is monophyletic. The family Limapontiidae and the limapontiid genus Ercolania are paraphyletic. The Oxynoacea and Oxynoidae are paraphyletic. The Juliidae are not resolved and the position of Oxynoe viridis remains again unclear. The analysis of the dataset with SplitsTree visualises the incongruencies in the dataset and the resulting incongru- encies in the different cladograms. Figure 5 shows a net- work connecting all terminal taxa. Parallel edges of same length within the network visualise a split between two groups. Members of each group share same characters (as- sumed apomorphic nucleotides for that group). Fit value of the analysis is 95.06, indicating that actually all con- flicts are visualised in the figure. No long branch taxa can be identified. Very well supported is the split between Oxynoacea + Cylindrobulla and Plakobranchacea, indi- cated by the long edges. Also well supported are splits be- tween species (e.g. Placida dendritica, Elysia timida or Elysia viridis) and the rest of the sequences. No split be- tween outgroup taxon Cy/indrobulla and Sacoglossa ex- ists (Fig. 5 and Fig. 6, the latter showing the analysis with a reduced data set for better visualisation). But a split be- tween Cylindrobulla + Tamanovalva limax + Oxynoe viridis against all other sequences can be observed (Fig. 5). Phylogenetic signal (shown as splits) 1s already low on genus level (e.g. Ercolania), or not existing (e.g. Cy- erce, Elysia or Thuridilla). A separate analysis of all Thuridilla species (Fig. 7) shows the close affinities of 7. ratna and T. gracilis, whereas T. bayeri appears separate. 3.3. Food spectrum in Sacoglossa A compilation of available data on food organisms of sacoglossans is presented in Table 2. Food organisms of Cylindrobulla are not known. All members of Oxynoacae feed on species of Caulerpa. In contrast, species of the Plakobranchacea have a much wider food spectrum: Members of the family Boselliidae seem to be restricted to the order Caulerpales (Bryopsidophyceae), mainly feed- ing on Halimeda. Within Plakobranchidae, Elysia shows the widest range on food organisms, containing Het- erokontophyta (Vaucheria, Padina, Biddulphia) and an- giospermes (seagrasses). Elysia timida and Elysia filicau- da feed on Acetabularia (Dasycladales). KREMER and JANKE (1988) claimed that chloroplasts of the algal genus Codium have been found in Elysia timida as well. Only for one species of Thuridilla, T. hopei, data are available: Derbesia and Cladophora. The only two references for Plakobranchus ocellatus are based on observations in the laboratory (JENSEN 1980a after Switzer-Dunlap 1975; Hı- ROSE 2005 after Adachi 1991). This species feeds on Chlorodesmis and Udotea. Interesting is the food organ- ism of Pattyclaya, which, according to JENSEN (1992), is Caulerpa. Polybranchiidae feed on species of the orders Bryopsidales and Caulerpales (Bryopsidophyceae) except for Mour- gona (Cymopolia (JENSEN 1981; CLARK 1994) and Aceta- bularia (HAMATANI 1994), Dasycladales, Dasyclado- phyceae). Hermaeidae also exhibit a broader spectrum of food organisms: The genus Hermaea 1s characterized by feeding on Rhodophyta (GRAHAM 1955 after Pelseneer 1935; TAYLOR 1968; TAYLOR 1971; KREMER & SCHMITZ 1976; KREMER & SCHMITZ 1976 after Cornet & Marche- Marchad 1951; JENSEN& CLARK 1983; JENSEN 1993a, b; WILLIAMS & WALKER 1999 after JENSEN 1983) except Her- maea vancouverensis consuming Isthmia nervosa, a di- atom (WILLIAMS & GOSLINER 1973). Aplysiopsis feeds on algae belonging to the orders Cladophorales (JENSEN 1980a after Gonor 1961; JENSEN 1980a after Greene 1970; JENSEN 1993a; TROWBRIDGE 1993; JENSEN 1995; WILLIAMS & WALKER 1999 after Jensen 1983) or Ulvales (JENSEN 1980a after Gonor 1961; JENSEN 1980a after Greene 1970; TROWBRIDGE 1993), Costasiella feeds on algae belonging to the orders Bryopsidales (CLARK & Busacca 1978; JENSEN 1980a, 1981; CLARK et al. 1981; CLARK 1984; JENSEN 1993a; CLARK 1994; WILLIAMS & WALKER 1999 after Jensen 1980, 1981, 1983) and Caulerpales (JENSEN 1981; WILLIAMS & WALKER 1999 after Jensen 1980, 1981, 1983) (Bryopsidophyceae), Cladophorales (WILLIAMS & WALKER 1999 after Jensen 1980, 1981, 1983) and Vaucheria (Heterokontophyta) (JENSEN 1990a, b). Limapontiidae show the most extraordinary food range. Species of the genus Limapontia feed on Cladophorales (GRAHAM 1955 after Gascoigne 1952; JENSEN 1975, 1980a after Gascoigne 1956), but also Bryopsidales (JENSEN 1975, 1980a after Gascoigne 1956), Ulvales (GRAHAM 1955 after Gascoigne 1952) and the heterokontophyte Vaucheria (GRAHAM 1955 after Gascoigne 1954; GAS- COIGNE & SARTORY 1974; JENSEN 1980a after Gascoigne 1956; HARTOG 1959). Alderia also feeds on Vaucheria (Evans 1953; GRAHAM 1955 after Gascoigne 1954; CLARK 1975; JENSEN 1980a after Hartog 1959; KruG & MANZI 1999), but there is one reference for the cladophoralean Rhizoclonium (GRAHAM 1955 after Gascoigne 1954). The genus Placida mainly feeds on species of Bryopsidales (BRUEL 1904; GRAHAM 1955 after Gascoigne 1954; BLEAKNEY 1989 after Thompson 1976 and after Millen 1980; JENSEN 1980a; JENSEN 1980a after Monselise & Mienis 1977 and after Schmekel 1982, JENSEN 1981, 1990b; MARZO, DI et al. 1993; TROWBRIDGE 2004 after Willan & Morton 1984, after Burn 1989 and after Trow- 236 Katharına HÄNDELER & Heike WAGELE: Phylogeny of Sacoglossa bridge 1998b) and Codium (BRUEL 1904 after Hecht 1895; GRAHAM 1955 after Gascoigne 1954; CLARK 1975; TROWBRIDGE 2004 after Willan & Morton 1984, after Burn 1989 and after Trowbridge 1998b; BEHRENS 2004) (Bry- opsidophyceae). BRÜEL (1904) referenced to TRINCHESE 1876 that Placida dendritica feeds on Ulva doubting this fact at the same time. The observation that Placida den- dritica ate another sacoglossa (MARCUS DU BoIs-REYMOND 1972) seems to be a particular case. Placida kingstoni ex- tended its diet to Cladophora (JENSEN 1981). The genus Elysia timida 1 Elysia timida 2 Elysia timida 3 Elysia obtusa Elysia macnaei Elysia spec.5 Elysia spec.1 Elysia ornata 1 Elysia ornata 2 Elysia ornata 3 Elysia amakusana Elysia tomentosa 1 Elysia tomentosa 2 Elysia viridis 3 Elysia viridis 1 Elysia viridis 2 Elysiella pusilla 1 Elysiella pusilla 2 Elysiella pusilla 3 Elysiella pusilla 4 779 Elysia benettae Plakobranchus ocellatus 1 Plakobranchus ocellatus 2 Thuridilla gracilis Thuridilla ratna 1 Thuridilla ratna 2 Thuridilla bayerı Thuridilla lineolata Thuridilla hopei 1 Thuridilla hopei 2 Thuridilla vatae Thuridilla albopustulosa Thuridilla carlsoni 1 Thuridilla carlsoni 2 Thuridilla kathae Thuridilla hoffae Bosellia mimetica 1 Bosellia mimetica 2 Bosellia mimetica 3 Cyerce spec.2 Cyerce spec.1 Cyerce nigricans Cyerce antillensis Cyerce edmundsi Polybranchia cf. orientalis Mourgona osumi | Placida dendritica 1 Plakobranchidae | Boselliidae Polybranchiidae Placida dendritica 2 Placida dendritica 3 Ercolania spec.5 Ercolania spec.1 Volvatella viridis | Limapontiidae J Limapontiidae J Limapontiidae ] Volvatellidae Tamanovalva limax | Juliidae Julia exquisita Lobiger viridis 1 Lobiger viridis 2 Oxynoidae Lobiger viridis 3 Cylindrobulla beauii Oxynoe viridis J] Oxynoidae Fig. 3. Sacoglossa; 16Sr DNA gene: Maximum Likelihood tree (GTR+G+I-model, AIC). Ercolania prefers Cladophorales (CLARK 1975; JENSEN 1980a; JENSEN 1980a after Trinchese 1872, after Rao 1937, after Schmekel 1968, after Rasmussen 1973 and after Usu- ki 1977; JENSEN 1981; MARIN & Ros 1992 after Marin & Ros 1988; JENSEN 1993a; MARZO, DI et al. 1993; CLARK 1994; JENSEN 1999; GRZYMBOWSKI et al. 2007), one ref- erence for Caulerpa (JENSEN 1980a after Edmunds 1963) is not confirmed again. Stiliger fuscovittatus feeds on red algae (JENSEN 1980a after Lance 1962). Unusual feeding strategies in the family show Calliopea oophaga, Olea hansineensis and Stiliger vesiculosus, they consume eggs of other Opisthobranchia (JENSEN 1986; LEMCHE 1974; CRANE 1971; JENSEN 1999 after Haefelfinger 1962). In Figure 8, the available data of higher food taxa are mapped onto the cladogram of the Maximum Likelihood analysis. Two references are not included: KREMER and JANKE (1988) determined chloroplasts in Elysia tímida as chloroplasts of Codium neither describing the origin of the included REM-slide nor giving a reference. Elysia timi- da was never described feeding on Codium, a misdeter- mination of the chloroplasts is therefore likely. BRUEL (1904) already doubted Trinchese’s information that Placi- da dendritica feeds on Ulva. Trinchese’s observation was never confirmed over the following years. 4. DISCUSSION 4.1. Phylogeny of Sacoglossa The Oxynoacea, comprising the shelled sacoglossans, are paraphyletic in all phylogenetic analyses. Applying SplitsTree, there exists no information for a split Cylin- drobulla against Sacoglossa. But according to the very long edges, there is much information for a split Cylin- drobulla + Oxynoacea against Plakobranchacea. JENSEN (1996) mentioned an eversible oral tube, a long, rod- shaped preradular tooth and a large female genital papil- la as synapomorphies for the Oxynoacea. These charac- ters are not present in Cylindrobulla. Using a cephalaspid as outgroup and including Cy/indrobulla as part of the in- group is not applicable for this gene, since it shows already a high substitution rate for the present data set. But the position of Cvlindrobulla as member of the Oxynoacea has to be reconsidered by using a more conservative gene, as 16S has proven to be. CO1 is also not apt for this inves- tigation, since it shows high substitution rates on nu- cleotide level and is not informative on amino acid level (unpublished data of the authors). The unresolved posi- tion of Oxynoe viridis at the base of the cladograms is pe- culiar. The genus never groups with Lobiger, both usual- ly united under the Oxynoidae. A contamination is unlike- ly due to high affinity with opisthobranch sequences. SplitsTree visualizes uniting positions with Cylindrobul- Bonner zoologische Beiträge 55 (2006) 237 la and Tamanovalva limax, which might present homo- plastic characters. The Juliidae are monophyletic only in the Maximum Likelihood analysis, but support is present in the SplitsTree analysis. According to JENSEN (1996) the bivalved shell and small, paired pharyngeal pouches are autapomorphies of this family. It is most unlikely that the bivalved shell has evolved several times independently, as several phylogenetic analyses presented here would suggest. 0.psElysia timida 3 na 1.008 Elysia timida 2 048 | Elysia timida 1 Elysia obtusa Elysia macnaei Elysia spec.5 Elysia spec.1 Elysia ornata 1 Elysia ornata 2 Elysia ornata 3 Elysia amakusana 1.00; Elysia tomentosa 1 Elysia tomentos 2 Elysia viridis 3 Elysia viridis 1 Elysia viridis 2 pElysiella pusilla 1 ft Elysiella pusilla 2 Elysiella pusilla 3 Elysiella pusilla 4 Elysia benettae (1.01 Plakobranchus ocellatus 1 Plakobranchus ocellatus 2 | 0.627 huridilla gracilis Plakobranchidae 1.40 Thuridilla ratna 1 A* Thuridilla ratna 2 Thuridilla bayeri Thuridilla lineolata .op Thuridilla hopei 1 Thuridilla hopei 2 Thuridilla vatae Thuridilla albopustulosa 1.00 Thuridilla carlsoni 1 pl. Thuridilla carlsoni 2 Thuridilla kathae | Thuridilla hoffae 6 Bosellia mimetica 1 = Bosellia mimetica 2 Bosellia mimetica 3 Cyerce spec.2 Cyerce spec.1 Cyerce nigricans Cyerce antillensis Cyerce edmundsi Polybranchia cf. orientalis Mourgona osumi 0.5Placida dendritica 1 had Placida dendritica 2 Placida dendritica 3 Ercolania spec.5 Ercolania spec. Plakobranchidae Plakobranchidae Boselliidae Polybranchiidae Limapontiidae ] Limapontiidae ] Limapontiidae | Tamanovalva limax ] Juliidae | Volvatella viridis ] Volvatellidae | Julia exquisita J Juliidae | 0.84Lobiger viridis 1 | 1.00 | obiger viridis 2 | Oxynoidae | Lobiger viridis 3 Oxynoe viridis ] Oxynoidae Cylindrobulla beauii Fig. 4. Sacoglossa; 16Sr DNA gene: Bayesian Analysis, 50%- Majority Rule consensus tree with posterior probabilities (GTR+G+I-model, AIC). In all analyses, the Plakobranchacea sensu JENSEN (1996) are monophyletic and this is visualized very well by the long edges separating this taxon from Oxynoacea + Cylin- drobulla. But this is not the case for the higher ranking taxa Plakobranchoidea and Limapontioidea. Whereas the former taxon was recognized at least in the maximum like- lihood and MrBayes analyses, the Limapontioidea with the genera Placida and Ercolania (both members of the Limapontiidae) never appeared monophyletic. This is of special interest taking into consideration the ongoing dis- cussion on the validity of Placida, Ercolania and Stiliger as three separate genera (e.g. MARCUS DU BoIs-REYMOND 1982; JENSEN 1985). In JENSEN’s (1996) analysis, the Limapontidae and the genus Costasiella (Costasiellidae) form a partly unresolved group. Limapontiidae 1s charac- terized by two visceral ganglia instead of three or more (JENSEN 1996). Contrary to Jensen’s analysis, the Polybranchiidae form a monophyletic group with exactly the same branching pattern of the six species (three genera) in all analyses, al- though no support can be seen in the splits graph. Inter- esting is the sister taxa relationship of the two Atlantic Cy- erce species, C. antillensis and C. edmundsi. MARCUS DU Bots-REYMOND (1982) and HAMATANI (1994) considered Cverce edmundsi Thompson, 1977 as synonymous with Mourgona germaineae Marcus & Marcus, 1970. Assign- ment of Cyerce edmundsi, as described by THOMPSON (1977), to the genus Cyerce contradicts diagnostic features of this genus: ,, The cerata proved to contain abundant lob- ules of the digestive gland“ (THOMPSON 1977: 137). Cy- erce 1s seperated from the other Polybranchiidae by hav- ing no branches of the digestive gland in the cerata (MAR- CUS DU BoIs-REYMOND 1982 after Eliot 1910). According to the results of this study, Cyerce edmundsi is related to Cyerce and not Mourgona. A re-arrangement of the gen- era and species is too early, more taxa of Limapontioidea with the families Polybranchiidae, Hermaeidae, Costasiel- lidae, and Limapontiidae have to be added in future in- vestigations to clarify validity of these taxa. Within the usually monophyletic Plakobranchoidea, two families have been considered, the monogeneric Boselli- idae and the species rich family Plakobranchidae. The genus Bosellia comprising only five species (JENSEN 1997a) was always a taxon to debate. PRUVOT-FOL (1954) settled Bosellia within the family Polybranchudae. She in- terpreted the dorsal apertures on the surface as a reminis- cence of lost cerata: ,,Des «orifices» semés sur la partie dorsale font penser que l'animal decrit est incomplet, c est- a-dire qu'il a perdu ses papilles.* (PRUVOT-FOL 1954: 180). However, PRUVOT-FOL has not examined a specimen of Bosellia by herself but followed the first description of TRINCHESE written in 1891. PORTMANN (1958) re-described the species and denied the existence of pores on the dor- 238 Katharina HANDELER & Heike WAGELE: Phylogeny of Sacoglossa P Bee ER o ly, bra A _ n ch lig e: Polybranchia cf. orientalis OS de Y a 1 BN 1 N c 7 N N Mourgona osumi y Ders sp.2 corer 2 Or Pe e o tneeeeTteetetttttenecs y, o yerce nigricans EE No Cyerce antillensis | Z IC . \ y i Julia exquisita Volvatella viridis .Q Limapontiidae Cyerce edmundsi y un : E = — - Ercolania sp.1 3 ; Placida dendritica 3 = Lobiger vindis 3 Placida dendritica 1 ‘ Lobiger viridis 2 Placida dendritica 2 Efoolayiia SPS Be viridis 1 „77° "Truridilla hopei 1 Año A = Thuridilla lineolata Thuridilla hopei 2 ima? . : 7 Thuridilla bayeri Oxynoe viridis. 2 / Thuridilaratnat \\ N N Hl! 7 Y Fe 7" Thuridilla gracilis — ¿— Cylindrobulla beauii / Thuridilla ratna 2 . \ ÉS AA y) Thuridilla vatae Ss Thuridilla albopustulosa \ Thuridilla carlsoni 2 Thuridilla carlsoni 1 Thuridilla kathae \ Thuridilla hoffae Bosellia mimetica 2 \ Bosellia mimetica 1 B Bosellia mimetica 3 —. Elysia timida 3 — Elysia timida 2 N Elysia timida 1 \ R A Elysia viridis 2 o . \ Elysia viridis 1 Elysia obtusa | Elysia vindis 3 Elysiella pusilla 1 * E. pusilla Elysia tomentosa 2 / S E. pusilla 2 + 3 Elysia tomentosa 1 A Elysia benettae P. océllatus 2 Iysia sp.1 Elysia macnaei / > Plakobranchus ocellatus Elysia sp.5 . > 7 Tas Elysia ornata 1 . SS Elysia ornata 3 RG Plak a Elysia ornata2 \Elysia amakusana ae Obr, an Ch; A A Rn - 'dae —— Fig. 5. Sacoglossa; Neighbor network analysis applying SplitsTree4 to visualize conflict in the whole sacoglossan data set. Par- allel edges with same length indicate a split and visualize the distances between the two groups involved. Note the long edges bet- ween the Plakobranchacea and the Oxynoacea + Cylindrobulla. Fit value: 95,85. sal surface. MARCUS DU BOIS-REYMOND (1980) and later THOMPSON and JAKLIN (1988) positioned Bosellia ın the family Plakobranchidae, but MARCUS DU BOIS-REYMOND (1982) transferred it in its own family Boselliidae based on the difference in number of chromosomes in Bosellia (n=7) and other sacoglossans (n=17), as well as the lack of parapodia. JENSEN (1996) criticized that the number of chromosomes is only known for the species Bosellia mimetica and not for other species of this genus but de- cided to retain the family status until the phylogenetic po- sition of the genus is solved. According to our results, Bosellia seems to represent an own evolutionary line, in- dependent of the family Plakobranchidae (with the gen- era Elysia, Thuridilla and Plakobranchus), but its posi- tion varies within the Plakobranchacea. The result of the Maximum Likelihood analysis places Bosellia as sister- taxon of Plakobranchidae, similar to the results obtained by JENSEN (1996). Bass and KARL (2006) used Bosellia mimetica and Bosellia marcusi in their phylogenetic analy- sis of Plakobranchidae. Similar to our results, B. mimet- ica forms the sister taxon to the further included genera Elysia, Elysiella, Plakobranchus and Thuridilla. But the other included member of Bosellia, B. marcusi, represents the sistertaxon to Elysia timida. The authors considered a wrong determination of their anımal. The family Plakobranchidae appears monophyletic only in the Maximum Likelihood analysis. This is in accor- dance with JENSEN’s analysis (1996). She mentioned as most important autapomorphies the parapodia in which the digestive gland and the reproduction system reach. In the SplitsTree analysis support of the split Plakobranchidae versus rest of Sacoglossa is not found at all. Within the Plakobranchidae, the genus 7huridilla is monophyletic in the NJ- and ML -analysis and branching pattern is identi- cal. The Bayesian analysis produced two of these mono- phyletic Thuridilla complexes but could not resolve their positions in detail. Also in the Splits analysis, no split Thuridilla/rest of Sacoglossa is visible. GOSLINER (1995: 45) mentioned following apomorphic features: „elongate rhinophores, single, thick, darkly pigmented ampulla, non- muscular, unarmed penis, bursa copulatrix with seperate A TR Bonner zoologische Beiträge 55 (2006) 239 gonopore and orange or red eggs.“ A darkly pigmented ampulla could not be confirmed in the Mediterranean Thuridilla hopei (own unpublished data). It is possible that only Indopacific species have a pigmented ampulla, since Gosliner dealt with these species. Elysia viridis also lays reddish yellow“ eggs when feeding on Chaetomorpha (TROWBRIDGE & Topp 2001: 222). One particular clade was found in all analysis: (7! baveri (T. ratna 2 (T. ratna 1 /T! gracilis))). The species are only distinguished by dif- ferent colouration (GOSLINER 1995). JENSEN (1992: 278) found „slight differences in the size of the pharyngeal pouch.“ BRODIE and BRODIE (1990) considered 7! bayeri as synonym with 7. ratna. GOSLINER (1995: 9) mentioned that “Internally, there is little difference between Thuridil- la bayeri [and] Thuridilla ratna*, “... these data strongly suggest that the described differences between 7. baveri and 7. ratna are simply variations within a single species that is extremely variable in its colouration.* Concerning Thuridilla gracilis, GOSLINER (1995) complained about the insufficient description of Risbec in 1928, hampering a correct assignment. RUDMAN (2000) stated that “Contrary to Gosliner (1995), I consider 7. bayeri to be a synonym of Thuridilla gracilis. Our study supports the synonymi- sation of 7. ratna with T. gracilis Risbec, 1928. But syn- onymisation of Thuridilla baveri with Thuridilla gracilis seems preliminary, since we have no information on in- 0.01 Volvatella viridis o Lobiger viridis 3 , Lobiger viridis 2 .. Lobiger viridis 1 o Oxynoe viridis Fig. 6. traspecific variability of the gene. According to our splits analysis, there are affiliations also between 7! bayeri and T. lineolata (Fig. 7). Comparison of GOSLINER's data (1995) with our data is hardly possible due to small over- lap of included species. In agreement is the sister taxa re- lationship of Thuridilla vatae and Thuridilla albopustu- losa. JENSEN (1997a) estimated the number of Plakobranchus- species from 1 to 14. In this study two specimens of Plako- branchus ocellatus were used. Although all analyses pub- lished yet (GOSLINER 1995; JENSEN 1996; Bass & KARL 2006) assign Plakobranchus to other members of the fam- ily Plakobranchidae, its position varies considerably. A unique character of the genus in relation to other mem- bers of Plakobranchidae is the dorsal position of the anus (JENSEN 1992). In all analyses on Plakobranchidae (JENSEN 1996; BASS & KARL 2006; this study), Elysiella pusilla is always lo- cated among Elysia species, E. benettae usually forming its sistertaxon. JENSEN and WELLS (1990: 324) emphasized the validity of an own genus Elvsiella: “this species shows so many differences from other species of Elysia that it definitely belongs to a different genus.” But all phyloge- netic analyses indicate the invalidity of this genus, there- Julia exquisita : Tamanovalva limax o Cylindrobulla beauii Neighbor network analysis applying SplitsTree4 to visualize conflict in the reduced data set, comprising only Oxynoa- cea and the outgroup taxon Cylindrobulla beauii. Note that there is no split separating Cylindrobulla beauii from the Oxynoacea. Similar long edges connecting Volvatella, Julia and Tamanovalva indicate the lack of signal of the taxon Juliidae. Fit value: 98,53. 240 Katharına HÄNDELER & Heike WÄGELE: Phylogeny of Sacoglossa fore Elysiella pusilla is named from hereon as Elysia pusil- la (Bergh, 1872). Several species, described under the genus name Elvsia (Elysia dubia Eliot, 1904, Elysia hal- imedae Macnae, 1954, Elysia latipes Marcus & Marcus, 1960 and Elysia macnaei Marcus, 1982) are mentioned to be synonymous to Elysia pusilla (see JENSEN & WELLS 1990). MARCUS DU BoIs-REYMOND (1980) distinguished material identified as Elysia halimedae by Baba (1957) and Burn (1972) from the original £. halimedae Macnae, 1954 from South Africa by the presence of papillate rhinophores in the latter. She re-named the material misidentified as £. halimedae and established the name E. macnaei Marcus, 1980. Our study includes a specimen from the Zoologische Staatssammlung Munchen, collect- ed in Sulawesi and identified by courtesy by Michael Schrödl as E. macnaei. This specimen does not group with Elysia pusilla indicating that £. macnaei 18 not synony- mous with £. pusilla. 0.01 Thuridilla carlsoni 1 Thuridilla carlsoni 2 * Thuridilla albopustulosa Thuridilla vatae Thuridilla ratna 2 ./ Thuridilla gracilis + Thuridilla ratna 1 Fig. 7. Thuridilla kathae Comparing our results with the only available phyloge- netic hypothesis on Sacoglossa elaborated by JENSEN (1996), the results of the Maximum Likelihood analysis come closest to her results. Both analyses support mono- phyly ofthe Plakobranchacea, Plakobranchoidea, Plako- branchidae as well as the genera Elysia (with Elysiella as a synonym) and Thuridilla. Plakobranchus and Bosellia form distinct evolutionary lines. Juliidae are also mono- phyletic. The two hypotheses contradict in the paraphyly (this study) versus monophyly (JENSEN 1996) of the Oxynoacea, Oxynoidae and Limapontiidae. But it has to be emphasized that these taxa are underrepresented in our study and future analyses have to be performed with more representatives of these groups. Furthermore, other genes have to be used for analysing deeper nodes of Sacoglos- sa, since the 16Sr DNA gene (this study), as well as the COI gene (unpublished results) show saturation on high- er level. Thuridilla hoffae Thuridilla hopei 1 Thuridilla hopei 2 * Thuridilla lineolata Thuridilla bayeri Neighbor network analysis applying SplitsTree4 to visualize similarity of species in a reduced data set, comprising only members of the genus Thuridilla. Note the one edge leading to the 2 specimens of Thuridilla ratna and the one specimen of 7: gracilis. No conflict occurs, indicating the synonymy of these 2 species. Thuridilla bayeri is separate, nevertheless the long edges characterizing the split Thuridilla gracilis /T. ratna /T. bayeri indicate its closer affinity to 7. gracilis, than the shorter edge cha- racterizing the split 7. bayeri/T. lineolata. Fit value: 98,12. Bonner zoologische Beitráge 55 (2006) 241 4.2. Food spectrum of Sacoglossa non-specific feeders, possibly because of the higher diver- sity of (see Table 2) foods in tropical seas.* (CLARK & Several authors emphasized that sacoglossans have a very BusaccA 1978: 281). That food spectrum varies in dif- narrow food spectrum (e.g. THOMPSON 1964; CLARK 1975, ferent taxa is evident in the genus E/ysia with species feed- 1994). However, the contrary was also stated: ,,..., some ing exclusively on one food species (e.g. Elysia translu- species feed on several different food items (...), and food cens on Flabellia (former Udotea) petiolata, see MARIN spectra are not so narrow as previously assumed.” & Ros 2004) or on a high variety of food species belong- (JENSEN 1980a: 73). „Tropical Ascoglossa are relatively — ing to completely different phyla (e.g. Elysia crispata feed- Elysia timida Dasycladales / Cladophorales Elysia obtusa Elysia macnaei Caulerpales Elysia spec. 5 Elysia spec. 1 Elysia ornata E Elysia amakusana Elysia tomentosa Elysia viridis Cladophorales / Bryopsis | Codium T Elysiella pusilla Bryopsidales T Elysia benettae Bryopsidales P. ocellatus ? Bryopsidales / Caulerpales ? Thuridilla gracilis Thuridilla lineolata Thuridilla hopei Cladophorales / Bryopsis | Codium Thuridilla vatae T Thuridilla albopustulosa Thuridilla carlsoni Thuridilla kathae Thuridilla hoffae a Bosellia mimetica Caulerpales Cyerce spec. 2 Cyerce spec. 1 Cyerce nigricans Caulerpales Cyerce antillensis Caulerpales a Cyerce edmundsi Polybranchia cf. orientalis Mourgona osumi Dasycladales _ Placida dendritica Bryopsis | Codium > Ercolania spec. 5 Cladophorales = Ercolania spec. 1 Cladophorales T A B Volvatella viridis Caulerpales Juliidae Caulerpales = Lobiger viridis Caulerpales = Oxynoe viridis Caulerpales Cylindrobulla beauii Fig. 8. Phylogeny of Sacoglossa (redrawn after the Maximum Likelihood phylogram, see Fig. 3) with food (higher taxa level) and tooth form mapped on the tree. T Triangular shaped teeth, S sabot shaped teeth, B blade shaped teeth. 242 Katharina HANDELER & Heike WAGELE: Phylogeny of Sacoglossa ing on the siphonalean heterokontophyte Vaucheria, the siphonocladalean Chaetomorpha (Cladophorales), the siphonalean Batophora (Dasycladales), on different siphonalean Halimedales and members of the taxon Bry- opsidales). In few species, an ontogenetic food switch is described, (e.g. juvenile Elysia timida feeds on Cladopho- ra dalmatica, adults on Acetabularia acetabulum (MARIN & Ros 1993)). Oxynoacea, representing about 20% of Sacoglossa (JENSEN 1997a), exclusively feed on the genus Caulerpa. Members of the Caulerpales with xylan in the cell wall remain the main food source of sacoglossans, even in the highly evolved Plakobranchidae (see Table 2). Some plakobranchids do not feed solely on Caulerpales, but seem to forage additionally on Cladophorales, which have cellulose in their cell wall. Ercolania and other limapon- tiids have switched to the Cladophorales as the main food source, Hermaeidae focus on Rhodophyta, and few sacoglossans are oophagous (consuming eggs of other opisthobranchs). A switch to Dasycladales with mannan in the cell wall is seldom (e.g. Elysia timida, E. filicau- da) and for these species other food sources (Caulerpales) are also recorded. Only species of Mourgona seem to fo- cus on Dasycladales. JENSEN (1997a) assumed that the evolution of the sacoglossans was linked to the structure of cell walls in their food organisms. Within evolution of Plakobran- chacea, members of this taxon could only change their food spectrum from Caulerpales and Udoteacea (contain- ing xylan) to Codiales and Dasycladales (with mannan) or Cladophorales/Siphonocladales (with cellulose) by ex- ploring Derbesiales (actual taxonomy: Bryopsidales) with cell walls of xylan in the haplonts and cell walls of mannan in the diplonts. According to JENSEN (1993a), 12% of the sacoglossans are able to feed on several algal species with different cell wall structure. She described popula- tions of Elysia viridis feeding on Chaetomorpha (Cladophorales) with cell walls containing cellulose and populations on Codium (Bryopsidales) with cell walls con- taining mannan (JENSEN 1989a). In experiments, she trans- ferred Elysia viridis collected from Codium to Chaetomor- pha and vice versa. Some specimens accepted the other algal species but still preferred the one that was their orig- inal food. In choice experiments, TROWBRIDGE and TODD (2001) con- firmed this behaviour of possible food switch. They showed that E. viridis collected on the native food Cladophora rupestris preferred this alga, but were able to switch to introduced species of Codium fragile ssp tomen- tosoides and C. fragile ssp. atlanticum. They also demon- strated that the Fl generation also preferred C. fragile in laboratory experiments, independent whether the parents were cultivated on Cladophora rupestris or Codium frag- ile. Juveniles grew faster on the introduced species. They concluded, that “slug performance is much better on the introduced host Codium fragile than on native Cladopho- ra.” (TROWBRIDGE & Topp 2001: 234). 4.3. Food spectrum and tooth shape BLEAKNEY (1990) detected different types of teeth in Placida dendritica in correlation with different kinds of food. JENSEN (1993a, 1997a) also found a positive corre- lation of tooth shape and algal diet. Within Oxynoacea (feeding exclusively on Caulerpa) the plesiomorphic tri- angular shaped tooth occurs as well as the blade-shaped tooth form. The triangular type ıs only found in those species, which feed on algae with xylan or mannan in the cell wall, but not cellulose. The blade-shaped tooth evolved several times within Sacoglossa, and this tooth type seems to be adapted for feeding on all types of al- gae. Only the sabot-shaped tooth seems to be restricted to those species mainly feeding on algae with cellulose. In an experiment JENSEN (1993a) could show that pheno- plastic variation of teeth underlies environmental factors. Elvsia viridis collected from Codium (Bryopsidales with mannan) was offered Chaetomorpha (Cladophorales with cellulose) as only food source for one year. The new- ly formed teeth in the upper limb of the radula were sim- ilar to those teeth of Elvsia viridis specimens collected from Chaetomorpha. Mapping tooth shape and algal food on the cladogram obtained by the ML analysis (Fig. 8) does not reveal correlation of slug evolution and tooth shape, nor toothshape in correlation with consumed algal taxon. 4.4. Food spectrum and phylogeny Based on her phylogenetic hypothesis, JENSEN (1997a) postulated that the ancestor of the Sacoglossa foraged on a filamentous calcified member of the udoteacean Halime- da (Caulerpales), since all basal Plakobranchacea are feed- ing on this type of algae. The monophyletic Oxynoacea switched to the uncalcified genus Caulerpa, on which the shelled sacoglossans exclusively feed. In this hypothesis, JENSEN (1997a) assumed that Cy/indrobulla also feeds on Halimeda, which would strengthen her hypothesis. But this was never observed. Actually, her note that “Cylin- drobulla is apparently associated with the non-photosyn- thetic, uncalcified rhizoids of Halimeda “ (JENSEN 1997a: 319) could indicate that Cvlindrobulla forages on epiben- thic algae associated with Halimeda. Her description on tooth morphology (JENSEN 1989b), as well as the lack of a suctorial pump (JENSEN 1996, 1997a) also suggests graz- ing as a feeding mode, and not piercing cell walls and Bonner zoologische Beiträge 55 (2006) 243 sucking out the cytoplasmatic contents. According to our analysis and the resulting phylogenetic hypothesis, anoth- er evolutionary scenario can be discussed, in which un- calcified Caulerpa is the original food source of Sacoglos- sa. This conforms to the hyptheses of CLARK & BUSAC- cA (1978), JENSEN (1980a) and CLARK & DE FREESE (1987). Caulerpa seems to be attractive even for sacoglos- sans that do not feed on that genus in situ. In experiments, JENSEN (1988) demonstrated that many slugs chose a cell homogenate of Caulerpa as food. But mapping radula morphology as described by JENSEN (1996) on our clado- gram (Fig. 8) produces incongruencies especially concern- ing the re-evolution of the triangular tooth shape. Fossil records of Sacoglossa are rare, but shelled Juliidae have been recorded from the Oligocene (JANSSEN 1979). JENSEN (1997a) dated back the origin of Sacoglossa in the Cretaceous because the fossil record of the calcified Hal- imeda dates back so far. She further noted that Sacoglos- sa then had to exist before the origin of Halimeda itself. We do not follow this statement since Halimeda could have existed long before the first sacoglossan slug evolved and foraged on this particular food. CLARK & DE FREESE (1987) hypothesized that Halimeda was already calcified when the first sacoglossan appeared and that feeding did not influence the grade of calcification significantly. In contrast, JENSEN (1997a) assumed an increasing calcifi- cation as a mechanical defense of the alga as an answer to predation by Sacoglossa and the adaptation of the buc- cal apparatus of Sacoglossa to deal with this mechanical defense. Calcification does not only appear in algae pre- dated by Sacoglossa but is a wide spread phenomenon in Rhodophyta or Phaeophyceae since it gives mechanical stability in areas with strong current or strong swell (e.g. dictyoalean Padina pavonica, Corallinacea, etc.). The mediterranean Coralligene is produced mainly by the red alga Mesophyllum alternans and Lithophyllum frondosum with a high calcification rate and no recorded predation (BALLESTEROS 2006). 4.5. Conclusions In future analyses, more taxa especially of the Oxynoacea and Limapontioidea have to be included to test the two different hypotheses presented by JENSEN (1996) based on morphology and the one outlined earlier by other authors and the present study based on molecular markers. Since both mitochondrial genes, 16Sr DNA (presented here) and CO1 (not included in this study), show a high substitu- tion rate already on higher taxa level, genes with lower evolutionary rates, e.g. nuclear genes as 18S and 28S, have to be analysed additionally. Further investigation on food organisms is also needed to analyse a possible co-evolu- tion of slugs and algae. This can only be done with a rig- orous identification of ingested algae, e.g. by analysing algal chloroplast genes in the slugs, which would exclude those algae used only as substrat. Acknowledgments. We thank Yvonne Grzymbowski (Bonn), In- go Burghardt (Bochum), Karen and Wolfgang Loch (Ankum), Kristina Stemmer (Bochum), Michael Schrödl (Munich) and An- nette Klussmann-Kolb (Frankfurt) for help in collecting mate- rial. We also thank Anne Hoggett and Lyle Vail from Lizard Is- land Research Station (Australia) for their continuous support. Alicia Cabello (Bonn) helped with SplitsTree analyses. 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Authors’ addresses: Katharina HÄNDELER, Institut für Evolutionsbiologie und Ökologie, An der Immenburg 1, Rheinische Friedrich-Wilhelms-Universität, 53121 Bonn, Germany, E-mail: uzsamk@uni-bonn.de; Heike WÄGELE (corresponding author), Institut für Evolutionsbiologie und Ökologie, An der Immenburg 1, Rheinische Friedrich-Wil- helms-Universität, 53121 Bonn, Germany and Zoologi- sches Forschungsmuseum Alexander Koenig, Adenauer- allee 160, D-53113 Bonn, Germany, E-mail: hwaegele(wevolution.uni-bonn.de Addendum: After receiving the proofs, GRZYMBOWSKI et al. 2007 was published, and a name for Ercolania spec. l is available now. The species is called Ercolania ken- colesi Grzymbowski, Stemmer & Wägele, 2007. Table 1. Specimens used in the phylogenetic analysis with accession numbers. ZSM Zoologische Staatssammlung Munich. Family outgroup OXYNOACEA Volvatellidae Pilsbry 1895 Juliidae E.A. Smith 1885 Oxynoidae H. Adams & A. Adams 1854 PLAKOBRANCHACEA Plakobranchoidea Boselliidae Marcus 1982 Plakobranchidae Rang 1829 Bonner zoologische Beitráge 55 (2006) Taxon Cylindrobullidae Thiele, 1931 P. Fischer, 1856 Volvatella viridis Hamatani, 1976 Julia exquisita (Gould, 1862) Tamanovalva limax Kawaguti & Baba, 1959 Oxynoe viridis (Pease, 1861) Lobiger viridis | Pease, 1863 Lobiger viridis 2 Pease, 1863 Lobiger viridis 3 Pease, 1863 Bosellia mimetica | Trinchese, 1891 Bosellia mimetica 2 Trinchese, 1891 Bosellia mimetica 3 Trinchese, 1891 Elysia amakusana Baba, 1955 Elysia benettae Thompson, 1973 Elysia macnaei Marcus E. duB.-R., 1980 Elysia obtusa Baba, 1938 Elysia ornata | (Swainson, 1840) Elysia ornata 2 (Swainson, 1840) Elysia ornata 3 (Swainson, 1840) Elysia timida | (Risso, 1818) Elysia timida 2 (Risso, 1818) Elysia timida 3 (Risso, 1818) Elysia tomentosa | Jensen, 1997 Elysia tomentosa 2 Jensen, 1997 Origin Cylindrobulla beauii Miama, 03-2005 South Island, Lizard Island/Australia, 12-07-2006 South Island, Lizard Island/Australia, 11-07-2006 Casuarina Beach, Lizard Island/Australia, 13-07-2006 South Island, Lizard Island/Australia, 10-07-2006 Lizard Island/Australia, 10-07-2006 Fiji Island, 26-08-2006 (ZSM 20061650) Coconut Beach, Lizard Island/Australia, 12-08-2006 Baretta del Abre near Matarö/Spain, 23-05-2006 Banyuls-sur-mer/Southern France, 11-05-2006 Tossa/Spain, 29-08-2003 Lizard Island/Australia, 04-03-2005 Samoa, 06-08-2005 (ZSM 20060293) NW-Sulawesi/Indonesia, 23-07-2003 (ZSM 20033821) Samoa, 20-08-2005 (ZSM 20060257) Lizard Island/Australia, 09-2002 North Point, Lizard Island/Australia, 14-07-2006 Lizard Island/Australia, 18-03-2005 Banyuls-sur-mer/Southern France, 02-05-2006 near Roses/Spain, 22-05-2006 near Tossa/Spain, 25-05-2006 Lizard Island/Australia, 13-09-2004 Lizard Island/Australia, 21-03-2005 Accession number EU140897 EU EU EU EU EU EU EU EU EU EU EU EU EU EU EU 140890 140895 140896 140891 140892 140893 140894 140872 140873 140874 140851 140868 140854 140860 140849 EU140850 EU EU EU EU EU EU 140848 140857 140858 140859 140852 140853 248 Katharina HANDELER & Heike WAGELE: Phylogeny of Sacoglossa Family Limapontioidea Polybranchtidae O’ Donoghue 1929 Taxon Elysia viridis | (Montagu, 1804) Elysia viridis 2 (Montagu, 1804) Elysia viridis 3 (Montagu, 1804) Elysia spec. (see WAGELE et al. 2006) Elysia spec. 5 Elysiella pusilla | Bergh, 1872 Elysiella pusilla 2 Bergh, 1872 Elysiella pusilla 3 Bergh, 1872 Elysiella pusilla 4 Bergh, 1872 Plakobranchus ocellatus | Van Hasselt, 1824 Plakobranchus ocellatus 2 Van Hasselt, 1824 Thuridilla albopustulosa Gosliner, 1995 Thuridilla bayeri (Marcus, 1965) Thuridilla carlsoni | Gosliner, 1995 Thuridilla carlsoni 2 Gosliner, 1995 Thuridilla gracilis (Risbec, 1928) Thuridilla hoffae Gosliner, 1995 Thuridilla hopei | (Verany, 1853) Thuridilla hopei 2 (Verany, 1853) Thuridilla kathae Gosliner, 1995 Thuridilla lineolata Bergh, 1905 Thuridilla ratna | (Marcus, 1965) Thuridilla ratna 2 (Marcus, 1965) Thuridilla vatae (Risbec, 1928) Cyerce antillensis Engel, 1927 Cverce edmundsi (Thompson, 1977) Cyerce nigricans (Pease, 1866) Origin Roscoff/Bretagne, Northern France, 2006 Tossa/Spain, 19-05-2006 Banyuls-sur-mer/Southern France, 06-05-2006 Lizard Island/Australia, 25-07-2005 South Island, Lizard Island/Australia, 24-07-2006 Maledives, 04-2006 Maledives, 04-2006 Maledives 04-2006 South Island, Lizard Island/Australia, 10-07-2006 Lizard Island/Australia, 21-03-2005 Lizard Island/Australia, 21-03-2005 NW-Sulawesi/Indonesia, 17-07-2003 (ZSM 20033615) N W-Sulawesi/Indonesia, 17-07-2003 (ZSM 20033612) Lizard Island/Australia, 13-09-2004 Lizard Island/Australia, 25-06-2006 Lizard Island/Australia, 18-03-2005 Samoa, 16-08-2005 (ZSM 20060224) Elba, 07-2001 Baretta del Abre near Matarö/Spain, 24-05-2006 Lizard Island/Australia, 13-09-2004 NW-Sulawesi/Indonesia, 07- 2003 Lizard Island/Australia, 25-06-2005 Lizard Island/Australia, 13-09-2004 Samoa, 11-08-2005 (ZSM 20060088) Azores, 09-2001 Channel, Lizard Island/Australia, 13-07-2006 Accession number EU140861 EU140862 EU140863 EU140856 EU14085 EU140864 EU140866 EU140865 EU140867 EU140875 EU140876 EU140889 EU140886 EU140877 EU140878 EU140883 EU140880 EU140881 EU140882 EU140879 EU140887 EU 140884 EU140885 EU140888 EU140841 EU140842 EU140843 an Family Limapontiidae Gray 1847 Bonner zoologische Beitráge 55 (2006) 249 Taxon Origin Accession number Cverce spec. | Lizard Island/Australia, 25-07-2005 EU140845 (see WAGELE et al. 2006) Cverce spec. 2 Lizard Island/Australia, 13-09-2004 EU140844 (see WAGELE et al. 2006) Polybranchia cf. orientalis Lizard Island/Australia, 01-09-2004 EU140846 (Kelaart, 1858) Mourgona osumi North Point, EU140847 Hamatani, 1994 Lizard Island/Australia, 14-07-2006 Placida dendritica | Tossa/Spain, 20-05-2006 EU140869 (Alder & Hancock, 1843) Placida dendritica 2 Tossa/Spain, 20-05-2006 EU140870 (Alder & Hancock, 1843) Placida dendritica 3 Tossa/Spain, 20-05-2006 EU140871 (Alder & Hancock, 1843) Ercolania spec. Casuarina Beach, EU140840 Lizard Island/Australia, 02-07-2006 Ercolania spec.5 Casuarina Beach, EU140839 Lizard Island/Australia, 07-07-2006 Table 2. Sacoglossan food organisms compiled from literature. Species OXYNOACEA Volvatellidae Ascobulla fischeri Ascobulla fragilis Ascobulla ulla Volvatella australis Volvatella bermudae Volvatella pyriformis Volvatella Juliidae Edenttellina typica Berthelinia caribbea Berthelinia darwini Berthelinia ganapati Berthelinia rottnesti Julia japonica Midorigai australis Tamanovalva babai Tamanovalva limax Food Caulerpa spp. Caulerpa prolifera Caulerpa racemosa Caulerpa racemosa, Laboratory: Caulerpa sertularoides, Caulerpa cupressoides, Caulerpa verticillata Caulerpa racemosa Caulerpa Caulerpa racemosa Caulerpa spp. Caulerpa Caulerpa scalpelliformis Caulerpa brownii Caulerpa scalpelliformis Caulerpa verticillata Caulerpa verticillata Caulerpa verticillata Caulerpa sp. Caulerpa racemosa Caulerpa simpliciuscula Caulerpa racemosa Caulerpa ambigua Caulerpa simpliciuscula Caulerpa simpliciuscula Caulerpa scalpelliformis, Caulerpa geminata Caulerpa scalpelliformis, Caulerpa geminata Caulerpa okamurai Reference GASCOIGNE & SARTORY 1974 after Burn 1972 MARIN & Ros 2004 JENSEN 1980a JENSEN 1981 CLARK 1994 JENSEN 1997b CLARK 1994 GASCOIGNE & SARTORY 1974 after Burn 1972 FONTANA et al. 1999 GONOR 1961 after Burn 1960b Burn 1965 GASCOIGNE & SARTORY 1974 after Burn 1972 EDMUNDS 1963 JENSEN 1980a after Grahame 1969 CLARK 1994 JENSEN 1997b JENSEN 1980a after Sarma 1975 JENSEN 1993b WILLIAMS & WALKER 1999 JENSEN 1980a after Kawaguti & Yamasu 1966 GASCOIGNE & SARTORY 1974 JENSEN 1980a after Burn 1960 GASCOIGNE & SARTORY 1974 after Burn 1972 JENSEN 1980a after Burn 1960, 1965 JENSEN 1980a after Kawaguti & Baba 1959 250 Species Oxynoidae Lobiger serradifalci Lobiger souverbii Oxynoe antillarum Oxynoe azuropunctata Oxynoe olivacea Oxynoe panamensis Oxynoe viridis Roburnella wilsoni PLAKOBRANCHACEA Plakobranchoidea Boselliidae Bosellia corinnae Bosellia marcusi Bosellia mimetica Plakobranchidae Elysia atroviridis Elysia australis Elysia benettae Elysia canguzua Elysia catulus Elysia chilkensis Elysia chlorotica Elysia crispata Elysia degeneri Food Caulerpa prolifera Caulerpa prolifera Caulerpa racemosa Caulerpa racemosa Caulerpa racemosa, Laboratory: Caulerpa sertularioides Caulerpa paspaloides, C. cupressoides, C. sertularioides, Laboratory: C. racemosa Caulerpa paspaloides Caulerpa prolifera Caulerpa sertularioides Caulerpa sertularioides Caulerpa spp. Caulerpa Halimeda Halimeda opuntia Halimeda tuna Udotea, Halimeda Codium fragile Enteromorpha, Cladophora, Chaetomorpha Enteromorpha, Cladophora, Rhizoclonium Acetabularia peniculus, Chaetomorpha, Rhizoclonium, Enteromorpha Chlorodesmis Codium sp. Bryopsis plumosa, Codium sp. Laboratory: Chaetomorpha Zostera marina most probably Enteromorpha Cladophora sp. Cladophora spp., Vaucheria spp. Chaetomorpha sp. Vaucheria litorea Halimeda sp., Bryopsis plumosa, Penicillus sp., Batophora oerstedi Halimeda sp., Brvopsis plumosa, Penicillus sp., Batophora oerstedi, Caulerpa verticillata Laboratory: Caulerpa verticillata, C. racemosa, Halimeda discoidea, Chaetomorpha sp. Vaucheria litorea Penicillus capitatus, Halimeda incrassata, H. monile Bryopsis plumosa, Derbesia tenuissima Bryopsis plumosa Udotea Katharina HÄNDELER & Heike WÄGELE: Phylogeny of Sacoglossa Reference GONOR 1961 MARIN & Ros 2004 JENSEN 1980a JENSEN 1980b JENSEN 1980a; JENSEN 1980a after Marcus & Hughes 1974; CLARK & BUSACCA 1978 JENSEN 1980b CLARK 1994 MARIN & Ros 2004 JENSEN 1980a after Lewin 1970 JENSEN 1993c after Doty & Aguilar-Santos 1970 GASCOIGNE & SARTORY 1974 JENSEN 1993b JENSEN 1993a JENSEN & CLARK 1983 PORTMANN 1958 THOMPSON & JAKLIN 1988 KAWAGUTI & YAMASU 1965 JENSEN 1992 after Jensen 1991 JENSEN 1993a WILLIAMS & WALKER 1999 after Hinde 1983 and after Jensen 1990 JENSEN 1993a JENSEN 1980a after Marcus 1955 JENSEN & CLARK 1983 after Marcus 1955 JENSEN & CLARK 1983 CLARK 1975 JENSEN 1993c FRANZ 1968 CLARK 1975 JENSEN & CLARK 1983 after West 1977 RUMPHO et al. 2000 after Mujer et al. 1996 and after Pierce et al. 1996 CLARK & BUSACCA 1978 JENSEN 1980a THOMPSON & JARMANN 1989 PIERCE et al. 2003 Curtis et al. 2005 after Curtis et al. 2003 Curtis et al. 2005 Curtis et al. 2005 HORGEN et al. 2000 Species Elysia diomedea Elysia evelinae Elysia expansa Elysia filicauda Elysia flava Elysia flavomacula Elysia fusca Elysia gordanae Elysia hedgpethi Elysia japonica Elysia leucolegnote Elysia macnaei Elysia maoria Elysia ornata Elysia papillosa Elysia patagonica Elysia patina Elysia punctata Elysia rufescens Elysia serca Elysia subornata Elysia thompsoni Bonner zoologische Beiträge 55 (2006) 2 Food Padina sp. Biddulphia sp., ? Dictyota Biddulphia Caulerpa Caulerpa racemosa, C. cupressoides ? Acetabularia peniculus Acetabularia Caulerpa racemosa, C. cupressoides cf. Cladophora "Laboratory: Chaetomorpha antennina, Cladophoropsis sp. Codium sp. Cladophora sp. Codium fragile, Bryopsis corticulans Chaetomorpha, Cladophoropsis sp. Boodleopsis pusilla Halimeda cuneata Halimeda macroloba Codium Codium convolutum, Codium fragile ssp. tomentosoides Bryopsis plumosa Bryopsis sp. Bryopsis Sp. Penicillus spp. Halimeda ssp., Penicillus, Udotea flabellum Penicillus, Udotea, Halimeda adult: Codium adult: Bryopsis plumosa, veliger: Nanochloropsis sp. Penicillus, Halimeda Udotea Codium spp. Bryopsis sp. Bryopsis sp. Halodule wrightii, Ulva lactuca Halophila engelmanni, Thalassia testudinum, Halodule wrightii Syringodium filiforme Caulerpa ashmeadii, C. mexicana, C. paspaloides, C. racemosa Caulerpa ashmeadii, C. racemosa, C. sertularioides, Laboratory: C. cupressoides, C. mexicana, C. paspaloides Caulerpa Nn Reference JENSEN 1980a after Bertsch & Smith 1973 JENSEN 1980a; JENSEN 1980a after Marcus 1957 JENSEN 1993a JENSEN 1993b WILLIAMS & WALKER 1999 JENSEN & WELLS 1990 JENSEN 1992 after Schmekel 1968 and after Nuttall 1989 WILLIAMS & WALKER 1999 WAGELE & JOHNSON 2001 after Marin & Ros 1988 JENSEN 1990a JENSEN 1980a after Schmekel 1968 WAGELE & JOHNSON 2001 after Marin & Ros 1988 GREENE 1970 JENSEN 1993a JENSEN 1990a, JENSEN 1990b MACNAE 1954 PAUL & ALSTYNE 1988 JENSEN 1980a after Reid 1964 WILLIAMS & WALKER 1999 after Trowbridge 1995 JENSEN 1981 HORGEN et al. 2000 FONTANA et al. 2001 after Hamann & Scheuer 1993 and after Hamann et al. 1996 JENSEN 1980a CLARK 1984 JENSEN 1993a SCHRODL 1996 MUNIAIN et al. 2001 JENSEN 1993a CLARK 1994 MACNAE 1954 HORGEN et al. 2000 FONTANA et al. 2001 after Hamann « Scheuer 1993 and after Hamann et al. 1996 JENSEN 1980a; JENSEN 1980a after Horsoe 1956 JENSEN 1982 WILLIAMS & WALKER 1999 after Jensen 1983 CLARK & BUSACCA 1978 JENSEN 1980a JENSEN 1993b Species Elysia timida Elysia translucens Elysia trisinuata Elysia tuca Elysia viridis Elysiella pusilla Plakobranchus ocellatus Pattyclaya arena Pattyclaya brycei Thuridilla hopei Limapontioidea Polybranchiidae Caliphylla mediterranea Cverce antillensis Cyerce nigra Cyerce nigricans Mourgona germaineae Mourgona osumi Polybranchia viridis Hermaeidae Aplysiopsis enteromorphae Aplysiopsis formosa Katharina HANDELER & Heike WAGELE: Phylogeny of Sacoglossa Food Codium Acetabularia acetabulum juvenil: Cladophora dalmatica, adult: Acetabularia acetabulum Flabellia (as Udotea) petiolata Codium Halimeda spp., Caulerpa sertularioides, C. mexicana, C. racemosa, Avrainvillea nigricans, Udotea sp. Halimeda discoidea Halimeda discoidea, H. incrassata Halimeda incrassata Halimeda incrassata Codium tomentosum Codium Codium fragile Codium tomentosum Codium, Chaetomorpha Codium spp., Bryopsis plumosa, Chaetomorpha linum Codium tomentosum, Bryopsis spp., Cladophora rupestris Halimeda spp. Laboratory: Udotea, Chlorodesmis Laboratory: Udotea javensis, Bryopsis sp. Caulerpa spp. Caulerpa spp. Derbesia tenuissima Derbesia tenuissima Cladophora vagabunda Bryopsis plumosa Bryopsis Brvopsis Bryopsis plumosa Penicillus spp., Halimeda spp. Penicillus, Udotea, Halimeda Penicillus dumetosus, Halimeda ssp. Chlorodesmis comosa Chlorodesmis fastigiata Chlorodesmis Cymopolia barbata Cymopolia barbata Acetabularia rvukvuensis ? Caulerpa spp. Caulerpa racemosa Chaetomorpha sp., Cladophora, Urospora, Rhizoclonium Cladophora prolifera Reference KREMER & JANKE 1988 MARIN & Ros 1992 MARIN & Ros 1993 MARIN & Ros 2004 JENSEN 1992 CLARK & BUSACCA 1978 STIRTS & CLARK 1980 WAUGH & CLARK 1986 CLARK & DEFREESE 1987 CLARK 1994 TAYLOR 1968 BOUCHET 1984 TRENCH et al. 1973 THOMPSON & JAKLIN 1988 JENSEN 1989a JENSEN 1980a; JENSEN 1980a after Fretter 1941 and after Rasmussen 1973; JENSEN 1994 after Jensen 1990 TROWBRIDGE & Topp 2001 RUDMAN 2003 after Jensen 2003 JENSEN 1980a after Switzer-Dunlap 1975 Hirose 2005 after Adachi 1991 JENSEN 1992 JENSEN 1992 MARIN & Ros 2004 CIMINO & GHISELIN 1998 after Gavagnin et al. 1994b WAGELE & JOHNSON 2001 after Marin & Ros 1988, 1989 BRUEL 1904 CLARK & BUSACCA 1978 GASCOIGNE 1979 MARZO, DI et al. 1993 JENSEN 1980a JENSEN 1993 CLARK 1994 MILLER 1969 Hay et al. 1989 CIMINO & GHISELIN 1998 after Roussis et al. 1990 JENSEN 1981 CLARK 1994 HAMATANI 1994 JENSEN 1980a after Bertsch € Smith 1973 CLARK 1994 TROWBRIDGE 1993 JENSEN 1995 0 Aplysiopsis smithi Aplysiopsis zebra Hermaea bifida Hermaea cruciata Hermaea evelinemarcusae Hermaea vancouverensis Costasiellidae Costasiella nonatoi Costasiella ocellifera Costasiella pallida Limapontiidae Alderia modesta Calliopaea oophaga Ercolania boodleae Ercolania coerulea Ercolania emarginata Ercolania endophytophaga Ercolania fuscata Ercolania gopalai Ercolania nigra Ercolania translucens Ercolania vanellus Bonner zoologische Beiträge 55 (2006) Food Rhizoclonium sp., Cladophora sp., Chaetomorpha sp., Urospora sp. Cladophora Cladophora fuliginosa, Chaetomorpha Griffithsia Griffithsia, Delesseria, Heterosiphonia Griffithsia flosculosa Griffithsia flosculosa Bornetia secundiflora Griffithsia sp. Griffithsia - Griffithsia, Dasya Griffithsia cf. ovalis Isthmia nervosa Avrainvillea Avrainvillea nigricans Avrainvillea nigricans Avrainvillea, Laboratory: Caulerpa fastigiata Avrainvillea nigricans Avrainvillea Avrainvillea nigricans, Caulerpa fastigiata, Cladophora Avrainvillea Avrainvillea nigricans Vaucheria vipera Vaucheria spp. Vaucheria, Rhizoclonium Vaucheria spp. Vaucheria spp. Vaucheria longicaulis Eggs of Philine denticulata, Laboratory: eggs of Polycera quadrilineata, Alderia modesta, Ercolania nigra, Limapontia capitata, Retusa truncatula, Turritella communis Eggs of Philine denticulata and Retusa truncatula Chaetomorpha aerea, C. moniligera, Cladophora spp. Valonia sp. Valonia sp., Dictvosphaeria cavernosa, juvenile: Cladophoropsis sp. Dictyosphaeria, Valonia, Cladophoropsis Chaetomorpha, Cladophoropsis Struvea plumosa (native), Laboratory: Valonia spec. Cladophora spp., Chaetomorpha linum Chaetomorpha sp. Chaetomorpha linum Rhizoclonium ? Caulerpa verticillata Reference JENSEN 1980a after Gonor 1961; JENSEN 1980a after Greene 1970 JENSEN 1993a 233 WILLIAMS & WALKER 1999 after Jensen 1983 GRAHAM 1955 after Pelseneer 1935 TAYLOR 1968 TAYLOR 1971 KREMER & SCHMITZ 1976 KREMER & SCHMITZ 1976 after Cornet & Marche-Marchad 1951 JENSEN & CLARK 1983 JENSEN 1993a WILLIAMS & WALKER 1999 after Jensen 1983 JENSEN 1993b WILLIAMS & GOSLINER 1973 JENSEN 1993a CLARK & BUSACCA 1978 JENSEN 1980a JENSEN 1981 CLARK et al. 1981 CLARK 1984 WILLIAMS & WALKER 1999 after Jensen 1980, 1981, 1983 JENSEN 1993a CLARK 1994 JENSEN 1990a, 1990b Evans 1953 GRAHAM 1955 after Gascoigne 1954 CLARK 1975 JENSEN 1980a after Hartog 1959 KRUG & Manzı 1999 JENSEN 1986 LEMCHE 1974 (Appendix of GASCOIGNE & SARTORY 1974) JENSEN 1980a after Usuki 1977 JENSEN 1980a after Schmekel 1968 JENSEN 1981 JENSEN 1993a JENSEN 1993a JENSEN 1999 CLARK 1975 JENSEN 1980a after Rao 1937 JENSEN 1980a; JENSEN 1980a after Rasmussen 1973 JENSEN 1997b JENSEN 1980a after Edmunds 1963 254 Katharina HANDELER & Heike WAGELE: Phylogeny of Sacoglossa Species Food Reference Ercolania viridis Chaetomorpha sp. JENSEN 1980a after Trinchese 1872 Chaetomorpha spp., Cladophora spp. JENSEN 1980a Chaetomorpha linum MARIN & Ros 1992 after Marin & Ros 1988 Chaetomorpha, Cladophora, Cladophoropsis JENSEN 1993a, Jensen 1981 Chaetomorpha capillaris, Chaetomorpha linum MARZO, DI et al. 1993 Ercolania spec. | Ercolania spec.5 Ercolania sp. (CLARK 1994) Limapontia capitata Limapontia senestra Limapontia depressa Olea hansineensis Boergesenia cf. forbesii Boodlea sp. Cladophoropsis Cladophora arcta, Enteromorpha Cladophora spp., Chaetomorpha linum, Bryopsis plumosa Cladophora spp., Chaetomorpha linum, Laboratory: Bryopsis plumosa Cladophora rupestris Cladophora Cladophora rupestris Vaucheria, Rhizoclonium, Conferva Vaucheria spp., Rhizoclonium riparium Laboratory: Chaetomorpha Vaucheria Opisthobranchia-eggs GRZYMBOWSKI et al. 2007 Unpublished data HW CLARK 1994 GRAHAM 1955 after Gascoigne 1952 JENSEN 1975 JENSEN 1980a after Gascoigne 1956 and after Jensen 1975 GRAHAM 1955 after Gascoigne 1952 and after Gascoigne 1954 GASCOIGNE & SARTORY 1974 JENSEN 1980a after Gascoigne 1956 GRAHAM 1955 after Gascoigne 1954 JENSEN 1980a after Gascoigne 1956 and after Hartog 1959 JENSEN 1994 GASCOIGNE & SARTORY 1974 JENSEN 1999 after Haefelfinger 1962 Eggs of Haminoea virescens, Aglaja diomedea, CRANE 1971 Gastropteron pacificum, Laboratory: Eggs of Archidoris montereyensis, Hermissenda crassicornis, Dendronotus iris Placida aoteana TROWBRIDGE 2004 after Trowbridge 1998b, after Willan & Morton 1984, after Burn 1989 Codium fragile, Codium convolutum, Bryopsis vestita Placida capensis Placida daguilarensis Placida dendritica Placida kingstoni Placida viridis Placida sp. (BEHRENS 2004) Stiliger fuscovittatus Stiliger vesiculosus specimens of Codiaceae Bryopsis sp., Derbesia sp. Bryopsis plumosa Ulva Codium tomentosum Codium tomentosum, Bryopsis Hermaea paucicirra Codium tomentosum Codium sp., Bryopsis plumosa Bryopsis Bryopsis plumosa Codium adhaerens, Codium tomentosum, Codium vermilara, Bryopsis plumosa, Bryopsis hypnoides, (introduced) Codium fragile Codium spp., Bryopsis plumosa, Bryopsis corticulans Bryopsis sp. Bryopsis plumosa, Cladophora Bryopsis plumosa Codium magnum Polysiphonia pacifica eggs of Favorinus branchialis MACNAE 1954 JENSEN 1990b BRUEL 1904 BRUEL 1904 after Trinchese 1876b BrÜEL 1904 after Hecht 1895 GRAHAM 1955 after Gascoigne 1954 MARCUS, E. DuB.-R. 1972 after Salvat 1968 CLARK 1975 JENSEN 1980a after Fretter 1941 and after Greene 1970 BLEAKNEY 1989 after Thompson 1976 and after Millen 1980 MARZO, DI et al. 1993 TROWBRIDGE 2004 after Alder & Hancock 1843 and after Thompson 1976 WILLIAMS & WALKER 1999 after Jensen 1980 and Trowbridge 1991a, 1992, 1995 JENSEN 1980a JENSEN 1981 JENSEN 1980a after Schmekel 1968 and after Monselise & Mienis 1977 BEHRENS 2004 JENSEN 1980a after Lance 1962 JENSEN 1980a after Haefelfinger 1962 Bonner zoologische Beiträge Band 55 (2006) Heft 3/4 Seiten 255-281 Bonn, November 2007 Biogeography of the Sacoglossa (Mollusca, Opisthobranchia)* Kathe R. Jensex!) DZoological Museum, Copenhagen, Denmark *Paper presented to the 2nd International Workshop on Opisthobranchia, ZFMK, Bonn, Germany, September 20th to 22nd, 2006 Abstract. The Sacoglossa (Mollusca, Opisthobranchia) comprise almost 400 nominal species level taxa. Of these 284 are considered valid (1.e., no published synonymies) in this study. About half of the nominal species have been descri- bed before 1950, and the 10 most productive taxonomists have described about half of the species. Distributions of all valid species are reviewed. The highest diversity is found in the islands of the Central Pacific, though species diversity 1s almost as high in the Indo-Malayan sub-province. The Caribbean forms another center of species diversity. These three areas are distinguished by the high number of Plakobranchoidea. Similarity among provinces is generally low. Endemi- city is high in most provinces, but this may be an artifact of collecting activity. The decrease in number of species with latitude is spectacular, and the number of cold-water endemics is very low, indicating that sacoglossans in cold tempe- rate regions are mostly eurythermic warm water/ tropical species. The highest number of species in cold temperate are- as is found in Japan and Southeastern Australia. This coincides with high species diversity of the algal genus Caulerpa, which constitutes the diet of all shelled and many non-shelled sacoglossans. Keywords. Species diversity, endemism. 1. INTRODUCTION Information on biogeography is important for understand- ing speciation and phylogeny as well as for making dec1- sions about conservation. Ideally, combining a phyloge- netic tree with a distributional map should give informa- tion on whether species dispersed from a center of origin or were the result of vicariance events. For most marine invertebrate groups, however, phylogenies are not fully re- solved and/or taxonomy is not yet stable, and even infor- mation on distributions is incomplete. Species are still be- ing split or synonymized, and new and undescribed species are discovered. In a worst case scenario a distribution map would show the activities of taxonomists rather than ac- tual species distributions. In the present study existing dis- trıbutional data for the Sacoglossa (Mollusca: Opistho- branchia) is reviewed and analyzed with regard to differ- ent biogeographic theories as well as activities of taxon- omists over time. Phylogenetic analysis has been per- formed at the genus level (JENSEN 1996a), and for one genus, Thuridilla, at species level (GOSLINER 1995). The relationship of the Sacoglossa to other opisthobranchs has been discussed in several recent publications (JENSEN 19968; MIKKELSEN 1996, 1998; THOLLESON 1999; WAGELE et al. 2003). Sacoglossans are suctorial herbivores; only two or three species are oophagous, feeding on the eggs of other opisthobranchs (JENSEN 1993a, 1997a). This means that they have depth distributions restricted to the photic zone, i.e. generally <100m. Sacoglossans are also dietary spe- cialists, the majority of species feeding on siphonaceous green algae, especially Caulerpa spp. (Jensen 1997a). Hence they only occur in the habitats where these algae are found. The total number of valid species is around 300, but new species are still described and other species are synonymized. 2. MATERIALS AND METHODS Distributional data for all species of Sacoglossa were tak- en from the literature. The study has included most pub- lications of original descriptions to get the type localities. However, in the case of the oldest descriptions, the pub- lications by SCHMEKEL & PORTMANN (1982) and BOUCHET (1984) have been used. Also, national and regional fau- nal checklists have been included, as well as records pub- lished on the Sea Slug Forum (http://www.- seaslugforum.net/). All nominal species listed in Appen- dix | have been included in the first analysis for bias of taxonomic expertise and scientific activity. In the distri- butional analyses, however, only species considered valid in this study have been included. As the present study is not a taxonomic analysis, species identifications and syn- tO onymizations, with a few controversial exceptions, will not be discussed. Only synonymies that have been pub- lished and not subsequently contested are used. Thus species that have only been mentioned once in the liter- ature are, with few exceptions mentioned in the text, con- sidered valid. Biogeographic regions and provinces were taken from BriGGs (1995) (Fig. 1), and sacoglossan distributions among these provinces were recorded. Although it must be assumed that a species occurs continuously between the extreme points of distribution, species were only scored as occurring in a region or province 1f at least one published record existed. The number of endemic species was determined for each province. As some regions were clearly underrepresented with regards to faunistic studies on opisthobranchs, a few regions have been merged or deleted from the analyses. Similarity between biogeo- graphic regions or provinces was analyzed using three in- dices: CJ= Jaccard’s coefficient= 100(a/(N1+N2-a)) (Valentine 1966), SD= Dice coefficient = 100(2a/(2a+b+c)) (LEAL € BOUCHET 1991), and I=index Fig. 1. 56 Kathe R. Jensen: Biogeography of Sacoglossa of inclusion= 100(a/Nmin) (GOLIKOV 1989). These indices differ ın the weight placed on shared species (a) compared to total number of species in the compared regions (Nl, N2), and species found exclusively in one or the other of the compared regions (b, c). 3. RESULTS 3.1. Fossil history After the description of live specimens of bivalved sacoglossan gastropods (KAWAGUTI & BABA 1959), sev- eral papers on fossil species of these sacoglossans ap- peared. The first reviews of fossil sacoglossans were those of BOETTGER (1963) and Kay (1968). There have been ex- tensive discussions about the identity of the Recent Tamanovalva, Edenttellina and Midorigai and the Mid- dle Eocene genus Berthelinia (e.g. EDMUNDS 1963; BURN 1998). KEEN & SMITH (1961) listed several other fossil species and included all in the family Juliidae Dall, 1898, which had previously been located in the Bivalvia. More Map showing biogeographic regions used in the present study. Regions have been modified from BriGGs (1995). 1. Nort- heast Atlantic. 2. Lusitanian. 3. Mediterranean (including the Black Sea). 4. Northwest Atlantic. 5. Caribbean. 6. Southwestern At- lantic tropical and warm temperate. 7. Aleutian. 8. Oregonian. 9. Californian. 10. Mexican-Panamanian. 11. Southeastern Pacific warm temperate. 12. Cold temperate South America. 13. Northwest Pacific cold temperate. 14. Northwest Pacific warm tempera- te. 15. Northern and northwestern Australia. 16. Southwestern and southern Australia. 17. Southeastern Australia. 18. Northeastern Australia. 19. Great Barrier Reef. 20. New Zealand. 21. Southeast Atlantic. 22. Western Indian Ocean. 23. Indo-Polynesian regi- on (including Ryukyu Islands). 24. Hawaii. Bonner zoologische Beiträge 55 (2006) 257 recently several more fossil bivalved sacoglossans have been described (see LE RENARD et al. 1996 for review), and also a single species of Volvatella has been described from the Lower Miocene of France (VALDÉS & LOZOUET 2000). Thus there may be two or five Recent genera of bivalved sacoglossans, whereas there are 9 fossil genera extending from the Lower Eocene to Lower Pliocene. Most fossil species have been found in European locali- ties, but a few are from the Caribbean, and one each from Australia and Indonesia. However, no doubt more fossil species will be described in the future. The temporal and spatial distribution of fossil sacoglos- sans indicates that they arose as part of the Tethys Sea fau- na. As sea level receded and temperatures cooled down, their distribution became more restricted, and today there is only one species of Berthelinia in the Caribbean and one in the Panamanian region; the remaining species are Indo-West Pacific. Julia has one species in the East Pa- cific; the remaining species are Indo-West Pacific. For Vol- vatella there is only one species in the Caribbean, one in warm temperate South Africa, and the remaining species are Indo-West Pacific. The disappearance of a major part of the coral reefs at the end of the Cretaceous (BRIGGS 1995) may have created ideal conditions for speciation of siphonaceous green algae when sea level rose again in ear- ly Eocene. 100 # species Fig. 2. Frequency of species descriptions through time. 3.2. Recent species Slightly more than half (199 of 387) of the nominal species have been described before 1950. There is a distinct peak around the 1860s and 1870s when PEASE and BERGH were most active describing species from the Indo-West Pacif- ic and Costa and Trinchese worked in the Mediterranean (Fig. 2). After 1950, it is especially the MARCUSes (39 species) and K. BABA (30 species) who dominate the num- ber of new species (Table 1). The 10 most productive au- thors or groups of authors have described almost 50 % of the species. Of the 387 nominal species 284 (73%) have been includ- ed in the similarity analyses. The number of species recorded from regions and provinces shown in Fig. | is listed in Table 2. Some regions are distinctly underrepre- sented in regards to number of records. This is true for most of the southern cold temperate zone, but also for trop- ical East Atlantic and southern East Pacific. Most of the biogeographic regions and provinces are supported by the present study as indicated by the percentage of endemic species. The regions and provinces with less than 10% en- demism will be discussed below. The Northeast Atlantic and Mediterranean were the ear- liest studied areas. The number of species described dur- 258 Table 1. Number of species described by different authors (cal- led “group(s)”, when with one or several co-authors) through ti- me. All nominal taxa have been included. Author # species described Marcus (& Marcus), 1955-1982 39 Baba (et al.), 1935-1959 30 Jensen (et al.), 1980-1999 23 Bergh, 1871-1905 20 Pease, 1860-1871 17 Ortea (et al.), 1981-2006 15 Trinchese, 1869-1895 14 Ichikawa, 1993 1 Thompson (et al.), 1973-1988 10 A. Costa, 1862-1876 10 Total for 10 authors (groups) 189 Total # nominal species 387 Percentage described by 10 most productive authors (groups) 49 Total # authors (groups) 104 Average # species per author (group) 3.7 Kathe R. Jensen: Biogeography of Sacoglossa ing the 19th century is high, but many species have sub- sequently been synonymized. New species are still discov- ered (ORTEA & TEMPLADO 1990; PERRONE 1990; CERVERA et al. 1991), old synonyms resurrected (CERVERA & LOPEZ- GONZALEZ 1996; ORTEA & Moro 1998), and the validity of some species, even some of the more recently described ones, is still debated (CERVERA et al. 2006). In addition, species ranges appear to be expanding (THOMPSON 1983; ORTEA et al. 1997; EVERTSEN & BAKKEN 2002). Hence the present analyses represent an ad hoc picture of species dis- tributions and diversity. Three faunal provinces are recognized in the Northeast At- lantic: the warm temperate Mediterranean Sea, including the Black and Azov Seas (no sacoglossans occur in the Caspian and Aral Seas), the warm temperate Lusitanian province and the cold temperate Northeast Atlantic Bo- real region (BRIGGS 1995). Three species appear to be en- demic to the Northeast Atlantic cold-water region. How- ever, two of these may be identical to Lusitanian and/or Mediterranean species. The possible synonymy of Ercola- nia nigra and E. viridis is currently under study by the present author, and it is also likely that Calliopaea oopha- Table 2. Species distribution and endemicity of sacoglossan opisthobranch in biogeographic regions as defined by BRIGGs (1995). Only species considered valid in the present study have been included. n.d. not determined. Region # Species 1. Northeast Atlantic boreal 11 2. Lusitanian 40 3. Mediterranean + Black Sea 37 4. Northwest Atlantic boreal 6 5. Caribbean incl. Florida 40 6. Southwestern Atlantic tropical + warm temperate 19 7. Aleutian 6 8. Oregonian 8 9. Californian 9 10. Mexican-Panamanian 23 11. Southeast Pacific warm temperate 12. Cold temperate South America 4 13. Northwest Pacific cold temperate 29 14. Northwest Pacific warm temperate 41 15. North and Northwestern Australia 23 16. South and Southwestern Australia 22 17. Southeastern Australia 15 18. Northeastern Australia 18 19. Great Barrier Reef 22 20. New Zealand 5 21. Southeastern Atlantıc 9 22. Western Indian Ocean + Red Sea 32 23. Indo-Polynesian, incl. Ryukyu Islands 107 24. Hawaii 25 # endemics (%) *Two of these species may be synonymous with Lusitanian and Mediterranean species. Bonner zoologische Beiträge 55 (2006) 259 Table 3. Similarity between provinces of the Atlantic Ocean. The Southeast Atlantic is excluded due to lack of information. In this and all following tables only species considered valid in the present study have been included. CJ Jaccard’s Coefficient; SD Dice Coefficient; I Index of inclusion; N number of species included; n.d. not determined. a. CJ \SD NE Atl cold NW Atl cold N=11 N=6 NE Atl cold - 23.5 NW Atl cold 13.3 - Lusitanian 15.9 4.55 Mediterranean 16.7 4.76 Caribbean 1.69 7.84 Brazil n.d. n.d. b. I NE Atl cold NW Atl cold NE Atl cold - - NW Atl cold 33:3 - Lusitanian 63.6 33.3 Mediterranean 63.6 33.3 Carıbbean 9.09 66.7 Brazil n.d. n.d. ga Lemche, 1974 is a subtidal variety of C. bellula d’Or- bigny, 1837. Limapontia depressa may be the only endem- ic from this cold-water region, and even that species may occasionally be found further south on the French Atlantic coast (M. PODDUBETSKAIA, Bordeaux, pers. comm. 2006). The other two species of Limapontia only extend into the Lusitanian and/or Mediterranean province (PRUVOT-FOL 1954; SCHMEKEL & PORTMANN 1982); L. capitata occurs in all three provinces and L. senestra occurs in two provinces. L. capitata is also the only sacoglossan recorded from Iceland and the Faroe Islands (PLATTS 1985). Hence the genus Limapontia is probably a cold- water, eurythermal genus, which is endemic to the North- east Atlantic-Mediterranean region. Various records exist of unidentified species of Limapontia from other regions (e.g. ENGEL et al. 1940; BURN 1973; SCHRODL 1996). How- ever, these need confirmation as many species of Ercola- nia lack cerata in juveniles. One species, Hermaea variopicta, has its northern limit along the south coast of the UK (LEMCHE & THOMPSON 1974). Two species, Elysia viridis and Calliopaea bellu- la, have their northern limit around Trondheimfjord (BRAT- TEGARD € HOLTHE 2001). Four species, Alderia modes- ta, Placida dendritica, Limapontia capitata and L. senes- tra, occur in the northernmost part of Norway (VADER 1981; BRATTEGARD & HOLTHE 2001; EVERTSEN & BAKKEN 2002; pers. obs.), and all three species of Limapontia plus Alderia modesta have been recorded from the White Sea (ROGINSKAYA 2000; MARTYNOV et al. 2006). Prior to 1997 Lusitan. Medit Carib Brazil N=40 N=38 N=49 N=19 27.5 28.6 3.33 n.d 8.70 9.09 14.5 n.d. = 66.7 31.5 13.6 50.0 - 23.0 n.d. 18.7 13.0 - 47.1 1.21 n.d. 30.8 - Lusitanian Medit Carib 68.4 = : 35.0 26.3 - 21.1 n.d. 84.2 P. dendritica had not been recorded from Norway north of the Bergen area (EVERTSEN & BAKKEN 2002). Five species appear to be endemic to the Lusitanian province. These have all been described after 1980, so 1t is possible that they will be found in neighboring regions in the future. The Canary Islands has the highest species diversity of this region (27 of 40 species have been record- ed here). A number of Caribbean species have been record- ed from these islands in recent years (ORTEA et al. 1998). Since the sacoglossan fauna of these islands has been well documented over many years (ORTEA 1981; FERNANDEZ- OvIES & ORTEA 1986; CERVERA et al. 1988; ORTEA et al. 1990; TEMPLADO et al. 1990), and only one or a few spec- imens have been collected at one single time far from their native distribution area, these species have most likely been transported by human activities (CHAPMAN CARL- TON 1991). Few species have been recorded from Madeira, Salvage, Cape Verde and Azores Islands (ORTEA 1981; ORTEA & TEMPLADO 1990; ORTEA et al. 1988, 1990, 1998; MALAQUIAS & CALADO 1997; JENSEN 1995, in prep.). There is an old record of the shelled Ascobulla frag- ilis from the Atlantic coast of Spain, which is cited in more recent publications (PILSBRY 1895; PrtwoT-FoL 1954; CERVERA et al. 2006). As this species feeds exclusively on Caulerpa, which does not extend this far north (DOUMENGE 1995), this needs to be re-examined. Eight species are endemic to the Mediterranean. Some of these may be synonyms of other species with wider dis- 260 Kathe R. Jensen: Biogeography of Sacoglossa tributions (SCHMEKEL & PORTMANN 1982; THOMPSON 1988). Eight amphi-Atlantic (one of which may be cos- mopolitan), eight Northeast Atlantic-Lusitanian, and five endemic species do not extend into the eastern basin of the Mediterranean. Only four species found in the east- ern basin of the Mediterrnanean do not occur in the west- ern basin; two of these have been described recently (THOMPSON 1988). The total number of sacoglossans in the eastern basin is only 17 whereas 34 species are known from the western basin (SWENNEN 1961; BARASH & DANIN 1971; SCHMEKEL & PORTMANN 1982; THOMPSON 1983; BOUCHET 1984; THOMPSON et al. 1985; THOMPSON 1988; THOMPSON & JAKLIN 1988; CATTANEO-VIETTI & THOMP- SON 1989; CERVERA et al. 2006). The southern coast of the Mediterranean has been insufficiently studied. It ıs uncer- tain whether two or three species extend into the Black and Azov Seas. Limapontia capitata and Calliopaea bel- lula (as Stiliger bellulus) have been recorded previously (MURINA & ARTEMJEVA 1997), but recent pictures on the Sea Slug Forum (KURAKIN 2002) have shown that Ercola- nia viridis is present, and it remains to be seen whether the species previously identified as C. bellula has been misidentified, or whether both species occur. The Lusitanian and Mediterranean provinces have very high similarity indices (Table 3). The combined number of species amounts to 52, which is very close to the species diversity found in the Caribbean. However, the number of Plakobranchidae is lower in the eastern Atlantic provinces (Table 9). BRIGGS (1995) recognizes one circumpolar Arctic region comprising Spitzbergen, Greenland and the northern coasts of North America and Russia. No sacoglossans have been recorded from Spitzbergen (GULLIKSEN et al. 1999) or the north coast of North America (BLEAKNEY 1996; GODDARD & FOSTER 2002). The four species occurring in the White Sea also occur in the Russian part of the Bar- ents Sea (ROGINSKAYA 2000; MARTYNOV et al. 2006). In addition, MARTYNOV et al. (2006) mentions an old record of a single juvenile specimen of Placida dendritica from Kola Bay. This indicates that this area 1s influenced by the North Atlantic Current and should be included in the bo- real region. The single specimen of Alderia modesta recorded from western Greenland (PLATTS 1985) could be attributed to larvae transported from Canada, which may occasionally be able to find suitable habitats for metamor- phosis in Greenland. The latitude of Disko Fjord is about the same as northern Norway, where the species occurs regularly. It should be mentioned that in Danish waters this species seems to have disappeared from localities where prior to 1997 it was abundant (pers. obs.). Whether this is due to habitat deterioration or increased temperature is unknown. The fauna of the Northwest Atlantic is also very well stud- ied (e.g. MARCUS & Marcus 1970; Marcus 1972a,b; MARCUS & HUGHES 1974; CLARK 1975; JENSEN & CLARK 1983; BLEAKNEY 1996), though new species are still be- ing described from the tropical waters (ORTEA & ESPINOSA 1996, 2000, 2001, 2002; CABALLER et al. 2006; PIERCE et al. 2006). Only 6 species occur in the cold temperate province; one of these, Placida dendritica, may be cos- mopolitan (BLEAKNEY 1989) and one, Alderia modesta, is circum-boreal. Apparently only one species, Elysia catu- lus, ıs endemic to the Northwest Atlantic cold water re- gion (CLARK 1975). This species feeds on the seagrass Zostera marina, which does not occur in Florida. It is pos- sible that £. catulus ıs a dark pigmented variety of the oth- er seagrass feeding species, Elysia serca (JENSEN 1982), in which case there will be no endemic species for the Northwest Atlantic. Ercolania fuscata may occur from No- va Scotia, Canada to Sao Paolo, Brazil, but this distribu- tion 1s based on synonymization with £. vanellus and E. talis (JENSEN & CLARK 1983). Two species, Elysia chlorot- ica and Hermaea cruciata, have their southern limit in Florida (JENSEN & CLARK 1983), 1.e., just south of the cold-water region. The former species also occurs in the northern part of the Gulf of Mexico (BOONE 1982), and the latter has ıts northern limit in Massachusetts (MAR- cus 1972a). The species presently known as Limapontia zonata, and known only from its original collection (GOULD & BINNEY 1870), is probably a flatworm; no sacoglossan has transverse pigment bands. A separate Carolinian province could not be distinguished for sacoglossan opisthobranchs, and no difference is ev- ident between the continental and insular parts of the Caribbean (CLARK & DEFREESE 1987), including Bermu- da (CLARK 1984), although these provinces were consid- ered distinct by BRIGGs (1995). However, the majority of studies involving sacoglossans are from the Caribbean is- lands. The limit between cold-water and tropical faunas appears to be along the coast of Florida (JENSEN & CLARK 1983); 36 of the 49 Caribbean species have been record- ed from Florida (THOMPSON 1977; Marcus 1977, 1980; CLARK 1982; CLARK & DEFREESE 1987; PIERCE et al. 2006; VALDÉS et al. 2006). The degree of endemism in the Caribbean is exceptionally high (Table 2). Especially the number of Plakobranchidae is high (Table 9), indicating that speciation in this family has taken place within the province. Five of the 21 endemic species have been de- seribed after 1990, so they may be found outside this province in the future or be synonymized with other species. As for the Mediteranean and Lusitanian provinces, synonymies are extensively debated and the status of sev- eral species remains uncertain (JENSEN & CLARK 1983; MARCUS 1980; VALDÉS et al. 2006). Bonner zoologische Beiträge 55 (2006) 261 Table 4. Similarity of provinces of East Pacific. Due to the high similarity between Aleutian, Oregonian and Californian provin- ces, these have been merged (Al+Or+Cal). Other abbreviations as in Table 3. a CJ\ SD Aleutian Oregonian N=6 N=8 N=9 Aleutian - 85.7 66.7 Oregonian 75.0 - 82.4 Californian 50.0 70.0 - Al+Or+Cal - - - Mex-Panam 16.0 24.0 33.3 SE Pacific n.d. n.d. n.d. b. I Aleutian Oregonian Aleutian - - - Oregonian 100.0 - - Californian 83.3 87.5 - Mex-Panam 66.7 75.0 88.9 SE Pacific n.d. n.d. n.d. The genus Bosellia appears to be an Atlantic warm-wa- ter genus with one amphi-Atlantic, one Canary Islands en- demic (FERNANDEZ-OVIES & ORTEA 1986), and two Caribbean species (MARCUS 1973). There are some scat- tered reports of Bosellia from the Indo-Pacific region (Marcus 1978; IMAMOTO 2004; PITTMAN 2004; RIEK 2006). However, too few specimens have been recorded to either identify them as one of the described species or decide that they are undescribed species. BriGGs (1995) recognized a tropical Brazilian province ex- ‘tending to just south of Rio de Janeiro. For sacoglossans, most Brazilian species extend south to the area around Sao Paolo (MARCUS (ER.) 1955, 1957; MARCUS & MARCUS 1967; Marcus (Ev.) 1977). This can probably be ex- plained by the extensive collecting activity of the Marcus- es around Sao Paolo; 47% of the Brazilian species have been described by them. Furthermore, more than 90% of the Brazilian species also occur in the Caribbean (Table 3b), and the number of endemic species is low (Table 2). This is most likely also due to the activities of the Mar- cuses in both these regions. TROWBRIDGE (2002) reviewed the Northeast Pacific sacoglossan fauna. She recognized four provinces, but un- fortunately the borders are not exactly the same as sug- gested by BRIGGS (1995). This is especially evident for the Californian province, in which Trowbridge records one species of the bivalved genus Berthelinia plus a couple of unidentified/ undescribed species. The present study found no endemic species in the Aleutian, Oregonian and California provinces and hence these were merged. Also, no difference was obvious between the Mexican and Pana- manian provinces, which have also been merged before Californian Al+Or+Cal Californian Al+Or+Cal Mex-Panam SE Pacific N=10 N=23 N=9 - 27.6 n.d. - 38.7 n.d. - 50.0 n.d. 7 48.5 10.5 32.0 43.8 5.56 28.0 - Mex-Panam 80.0 - 11.1 77.8 comparisons with other provinces (Table 4). In the Mex- ican-Panamanian province four species have been record- ed after the publication of Trowbridge’s paper (BEHRENS & HERMOSILLO 2005; KRUG et al. 2007) and one species, Ascobulla californica (originally described as Cylindrob- ulla californica by HAMATANI (1971)), was not considered a sacoglossan by TROWBRIDGE (2002). The occurrence of Alderia modesta ın the Mexican-Panamanian region is probably the recently described species, Alderia willowi (KRUG et al. 2007), which occurs southwards from cen- tral California. The monotypic genus Olea is endemic to the Northeastern Pacific region, extending from the Aleut- ian to the Californian province (TROWBRIDGE 2002). Her- maea vancouverensis 18 a cold-water species, extending across the Bering Strait to the Kurile province (CHERNY- SHEV 2005). One species, Elysia hedgpethi, occurs from British Columbia, Canada to Chile (SCHRODL 1996; Trow- BRIDGE 2002), though the occurrence in Chile needs ver- ification. The species extending into the warm temperate region of the Southeast Pacific almost all are shared with the trop- ical Mexican-Panamanian region (SCHRÓDL 1996; Trow- BRIDGE 2002; BEHRENS & HERMOSILLO 2005). Julia the- caphora is considered the oldest name for J. equatorialis, which was also described from the tropical East Pacific. Only four species have been recorded from cold-temper- ate South America, one from the Atlantic coast and three from the Pacific (MARCUS 1959; SCHRODL 1996; MUNIAN & ORTEA 1997). Due to the low number of species and sparse collecting activity, the species from the Southeast Pacific coast have been considered as one province for analyses. 262 Kathe R. JENSEN: Biogeography of Sacoglossa Table 5. Similarity of Japanese biogeographic provinces and of Northwest and Northeast Pacific provinces. The Japanese provin- ces have also been compared to the neighboring Central Pacific sub-province. Abbreviations: see Table 3. a CJ \ SD Japan cold Japan warm temp., N=26 temp., N=41 Japanese cold temp. - 39:7 Japanese warm temp. 42.6 - Ryukyu 15.7 23.3 Central Pacific 14.9 22.7 NW Pacific cold temp. - n.d. NE Pacific cold temp. n.d. n.d. b. I Japan cold Japan warm temp. temp. Japanese cold temp. - - Japanese warm temp. 76.9 - Ryukyu 30.8 42.4 Central Pacific 38.5 41.5 NE Pacific cold temp - n.d. Four biogeographic regions or provinces can be distin- guished along the coasts of Japan. Biogeographically the southernmost archipelago of Ryukyu belongs in the vast tropical Indo-Polynesian province (BRIGGS 1995), but be- ing under Japanese jurisdiction, the sacoglossan fauna has been studied mostly by Japanese scientists (e.g. BABA 1936; HAMATANI 1980; ICHIKAWA 1993). Hence Ryukyu has been included in both analyses of the Japanese fauna (Table 5) and of the Indo-West Pacific one (Table 7). The warm-water temperate region comprises southern and eastern Japan, including the well studied Seto Inland Sea and Sagami Bay. The corresponding continental coast of China has been insufficiently studied, and only 14 species have been recorded from southern Korea (KOH 2002a,b, 2003, 2005a,b,c; RUDMAN, Sydney, pers. comm. 2007). The cold-water temperate oriental province ın- cludes the central and western coasts of Honshu, where- as the northernmost island of Hokkaido belongs to the Kurile province (BRIGGS 1995). The cold-temperate fau- na of Japan contains more species (N=26) than any oth- er cold-water fauna. A few shelled species extend into this province, which is also seen in southern Australia, but not in other cold-water provinces. Only three additional sacoglossan species have been recorded from the Kurile province (BABA 1935; CHERNYSHEV & CHABAN 2005; TROWBRIDGE 2006) and these were only included in the comparison of Northeast and Northwest Pacific cold-wa- ter faunas (Table 5). The highest number of species has been recorded from the warm temperate region (e.g. BA- BA 1949, 1952a,b, 1955, 1957, 1959, 1966; 1968; HAMATANI 1968, 1969, 1972, 1976a,b, 1994; KAWAGUTI Ryukyu Central Pacif. NW Pacif. cold NE Pacif. N=33 N=51 temp., N=29 cold temp. | N 27.1 26.0 - n.d. 37.8 37.0 n.d. n.d. - 38.1 n.d. n.d. 23.5 - n.d. n.d. n.d. n.d. - 20.5 n.d. n.d. 11.4 - Ryukyu NW Pacif. cold temp. 48.5 - n.d. 40.0 & BABA 1959; HIRANO et al. 2006); in fact BABA described 59% of the species from this province and 69% of the species from the cold-water region of Japan. A few of the species described by BABA have been synonymized with more widespread Indo-West Pacific species (BABA 1974; JENSEN 1985). On the other hand, many of BABA’s species have been identified outside Japan (JENSEN 1985; CARL- SON & Horr 1978, 2003; BURN 1998, 2006). The Northeast Australian and Great Barrier Reef faunas (BURN 1966b; THOMPSON 1973; MARSHALL & WILLAN 1999; WAGELE & JOHNSON 2001) are so closely related to each other that no endemics have been recorded from the Great Barrier Reef and only one endemic species, Placi- da fralila, has been recorded from Northeast Australia (Table 2). Hence these two provinces were merged for similarity analyses, and the combined province has two endemic species (Elysia bennettae, P. fralila). For the Great Barrier Reef 30% of the species were listed as unidentified and/or undescribed (MARSHALL & WILLAN 1999). This part of Australia has been included in the In- do-Polynesian province by BRIGGS (1995). This 1s support- ed by the low endemicity, and also, the similarity with the fauna of the South Pacific islands is higher than with any of the other Australian provinces (Tables 6 and 7). The North and Northwestern Australian fauna has rather high similarity to the Western Indian Ocean fauna (Table 7). This is in spite of the fact that almost 40% of the species have been described by the present author within the last 20 years (JENSEN 1993b, 1997b,c; JENSEN & WELLS 1990). The fauna of the South and Southwestern Australia has rm =-7__ Bonner zoologische Beitráge 55 (2006) 263 Table 6. Similarity of provinces of the Australian continent. Due to the high similarity between the Northeast Australian provin- ce and the Great Barrier Reef, these two provinces have been merged (NE Aus+GBR). Abbreviations as in Tab. 3. a. CJ1SD NE Austr GBR NE Aus+ GBR N+NW Austr SW+S Austr SE Austr N=18 N=22 N=28 N=23 N=22 N=15 NE Australia - 60.0 - 34.1 30.0 30.3 GBR 42.9 - - 26.7 18.2 16.2 NE Aus+GBR - - - 31.4 28.0 27.9 N+NW Austr 20.6 15.4 19.0 7 48.8 26.3 SW+S Austr 17.6 10.0 16.3 32.4 - 43.2 SE Austr 17.9 8.82 16.2 15.2 27.6 - b. Meee NE Austr GBR NE Aus+GBR N+NW Austr SW+S Austr NE Australia - - - - - GBR 66.7 - - 2 - NE Aus+GBR - - - - - N+NW Austr 38.9 27.3 34.8 - SW+S Austr 33:3 18.2 31.8 3 - 333 20.0 40.0 3.3 53.3 SE Austr the highest affınity to the fauna of North and Northwest- ern Australia, and the other way around, whereas the fau- na of Southeastern Australia (Burn 1958, 1960, 1965, 1974, 1998, 2006) has a higher affınity to the fauna of South and Southwestern Australia than to that of North- eastern Australia and the Great Barrier Reef (Table 6). This may change when the 50% unidentified/undescribed species listed for Southeastern Australia (BURN 2006) are properly named. Very few sacoglossans have been recorded from New Zealand (POWELL 1937; WILLAN & MORTON 1984; TROW- BRIDGE 1995a; SPENCER & WILLAN 1995). The fauna con- sists of widespread species and one endemic (Table 2). Hence this fauna has not been further analyzed in the pres- ent study. The sacoglossan fauna of the Red Sea (ELIOT 1908; O”- DONOGHUE 1929; HELLER & THOMPSON 1983) has about the same affinity to the fauna of the Indian subcontinent as to the Western Indian Ocean, and the index of inclu- siveness (1) for the Red Sea and India sensu lato (s.l., see below) is twice that for the Red Sea and the Western In- dian Ocean (Table 7b). Also, two out of the ten recorded species are endemic to the Red Sea. Hence the Red Sea should be considered a separate province. In other groups of invertebrates Lessepsian migrants are common. For sacoglossans this appears to be a small and recent problem (YOKES 2001, 2002; RUDMAN 2002). Except for these few species, the Red Sea does not share any species with the Mediterranean. The sacoglossans found in southwestern Africa are more closely related to those found in southeastern Africa than to any other region or province (GOSLINER 1987a); in fact no species are shared with the Brazilian fauna and only one species, Placida dendritica, ıs shared with the Lusi- tanian province. Hence this province has been considered in connection with the Western Indian Ocean province. Endemicity 1s high, but this could be due to the poor knowledge of tropical western Africa. Only one species has been described from this region (MARCUS & MARCUS 1966), so this was not included in the present study. The East Atlantic species Elysia viridis apparently occurs from central Norway (BRATTEGARD & HOLTHE 2001) to South Africa (GOSLINER 1987a), though no records exist between the Senegal (PRUVOT-FOL 1953) and South Africa, and GOSLINER (1998) has subsequently changed the identifi- cation to Elvsia sp. The species was originally identified in South Africa as the Indian species E. punctata by MAC- NAE (1954). GOSLINER (1987a) found a distinct faunal sep- aration for opisthobranchs at Port Elizabeth, whereas BriGGs (1995) considers the coast between the Cape of Good Hope and north of Durban one province. In the pres- ent study Port Elizabeth has been used to separate the fau- nas of southwestern Africa and the Western Indian Ocean. The Western Indian Ocean is considered a separate province by BrıGGs (1995). The sacoglossan fauna of this province has a high similarity with the South Pacific and North and Northwestern Australia (Table 7). The affinity with the fauna of India s./. (India, Sri Lanka and Maldives) is considerably lower, and also the affinity with the Indo- 264 Kathe R. JENSEN: Biogeography of Sacoglossa Table 7. Similarity of provinces of the Australian continent. Due to the high similarity between the Northeast Australian provin- ce and the Great Barrier Reef, these two provinces have been merged (NE Aus+GBR). Abbreviations as in Tab. 3. a. CJ \ SD SW Afr. WIO Red Sea Indias. /. Indo-Malay N+NW Centr. Pac. NEAus S Pacific Hawaii Ryukyu N=8 N=27 N=10 N=20 N=48 Austr — isl. +GBR N=33 N=25 N=33 N=23 N=51 N=28 E SW Afr. - 28.6 11.1 7.14 n.d. 12.9 n.d. n.d. 9.76 n.d. n.d. WIO 16.7 - 27.0 17.0 24.0 32.0 35.9 25:5 36.7 n.d. n.d. Red Sea 5.88 15.6 - 26.7 n.d. 18.2 n.d. n.d. 18.6 n.d. n.d. India s. /. 3.70 930 15.4 - 20.6 14.0 19.7 12.5 22.6 17.8 133 Indo-Malay n.d. 13.6 nd. 11.5 - 22.5 50.5 28.9 37.0 19.2 24.7 N+NW Austr 6.90 190 10.0 7.50 12.7 - 29.7 31.4 32.1 n.d. n.d Centr. Pac. isl. n.d. 21.9 n.d. 10.9 33.8 lig - 43.0 50.0 42.1 38.1 NE Aus+GBR n.d. 146 nd. 6.67 16.9 19.0 27.4 - 42.6 n.d. n.d. S Pacific 5.13 224 10.3 12.8 22.7 19.1 33.3 27.1 - 41.2 24.2 Hawaii n.d. n.d. n.d. 9.76 10.6 n.d. 26.7 n.d. 26.1 - 13.8 Ryukyu n.d. n.d. n.d. 3.92 14.1 n.d. 23.5 n.d. 13.8 7.41 - b. I SW Afr. WIO Red Sea Indias. Indo-Malay N+NW Centr. Pac. NEAus S Pacific Hawai Austr. isl. + GBR SW Afr. - - - - - - - - - - - WIO 62.5 - - - - - - - - - - Red Sea 12.5 50.0 - - - - - - - - - India s. /. 12.5 20.0 40.0 - - - - - - - - Indo-Malay n.d. 333 nd. 35.0 - - - - - - N+NW Austr 25.0 349 30.0 15.0 34.8 - - - - - Centr. Pac. ısl. n.d. 519 nd 35:0 52.1 47.8 - - - - NE Aus+GBR © n.d. 25.9 n.d. 15.0 39.3 34.8 60.7 - - - S Pacific 25.0 40.7 40.0 30.0 45.5 39.1 63.6 46.4 - - Hawaii n.d. n.d. n.d. 20.0 28.0 n.d. 64.0 n.d. 48.0 - Ryukyu n.d. n.d. n.d. 10.0 30.3 n.d. 48.5 n.d. 24.2 16.0 Malayan sub-province is lower. Endemicity 1s high (5 of 27 species), though two of these species have sometimes been synonymized with widespread Indo-West Pacific species (GOSLINER 1987b). The majority of species recorded from the Western Indian Ocean have been col- lected from the southern part of the region, 1.e. South Africa (THOMPSON 1979; GOSLINER 1987a,b, 1995), Tan- zania (ELIOT 1903, 1904; GOSLINER 1995), Madagascar (GOSLINER 1995) and Mauritius (BERGH 1888; GOSLINER 1995). A total of 107 species of sacoglossans have been record- ed from the vast Indo-Polynesian province. Most of these species are only distributed in part of the province and hence 1t was subdivided into five sub-provinces: The In- dian subcontinent, including Sri Lanka and the Maldive Islands was considered one sub-province (10 endemic species); the Andaman Sea, the South China Sea, Indone- sia and the Philippines form an Indo-Malayan sub- province (9 endemic species); the Mariana and Marshall Islands together with Micronesia were considered a Cen- tral Pacific sub-province (4 endemic species); Papua New Guinea, Solomon Islands, Fiji, Vanuatu, New Caledonia, Samoa and the Polynesian islands form a South Pacific province (6 endemic species); and as mentioned above the Ryukyu Islands form a separate sub-province (10 endem- ic species). Many of the endemic species have been de- scribed within the last 20 years, so they may actually have wider distributions. Of the 107 species recorded from the Indo-Polynesian province only 12 have distributions from the Western In- dian Ocean and/or the Red Sea to the Central and/or South Pacific islands, two species, Elysia ornata and Ercolania coerulea, are circum-tropical and one, Placida dendriti- ca, may be cosmopolitan. Two of the 12 widespread In- do-West Pacific species are shelled (Oxynoe viridis and Berthelinia schlumbergeri), seven are plakobranchoids (5 Bonner zoologische Beitráge 55 (2006) 265 species of Thuridilla, Elvsiella pusilla and Plakobranchus ocellatus) and three are limapontioids (Cyerce elegans, C. nigricans and Polvbranchia orientalis). The widest lati- tudinal distributions along the West Pacific rim are found in the four species that occur from the cold-temperate part of Japan to the cold-temperate part of Australia. Two of these species are the circum-tropical Elysia ornata and the questionably cosmopolitan Placida dendritica; the other two are Elysia obtusa and Stiliger smaragdinus. P. den- dritica has been synonymized with a number of species described from different places in the Indo-West Pacific (BLEAKNEY 1989), but the synonymy has been doubted (e.g. TROWBRIDGE 1995b). Two species, Oxynoe viridis and Elysiella pusilla, extend from warm-temperate Japan to southern Australia. Two further species, Thuridilla splendens and Polybranchia orientalis, extend from warm-temperate Japan to Northeast Australia and the Great Barrier Reef. Four species, Plakobranchus ocella- tus, Thuridilla vatae, T. hoffae and Cyerce nigricans, ex- tend from the Ryukyu Islands to tropical Australia. Most of these species also have wide longitudinal distributions. Plakobranchus ocellatus occurs in a number of colour va- rieties, and it is possible that a complex of sibling species is involved (see discussion on the Sea Slug Forum: http://www.seaslugforum.net/find.cfm?id=13970) (last access 12th of August 2007). The Indian sub-province has a high proportion of endem- ic species (50%). This is probably an artifact due to the activities of a few taxonomists who have worked only within this sub-province (KELAART 1858; NEVILL & NEVILL 1869; RAO 1937; Rao & RAO 1963; SARMA 1975). Most of G. & H. NEVILL’s species have been synonymized, though not consistently with the same species. Although doubtful, they have been considered valid in the present study. The species described by KELAART have been re- examined several times (ELIOT 1906; O’ DONOGHUE 1932) and they are still recognized as valid, mostly widespread Indo-West Pacific species. In spite of the high endemici- ty, India s./. shows higher similarity to the Indo-Malayan, Central and South Pacific sub-provinces than to the West- ern Indian Ocean province (Table 7). The highest simi- larity is found between India s./. and the Red Sea, but this is caused by the low number of species found in the Red Sea, and the high proportion of widespread Indo-West Pa- cific species. The Indo-Malayan and Central Pacific sub-provinces have twice the number of species occurring in tropical Australia and the Western Indian Ocean and 50% more species than the South Pacific and Ryukyu Islands. This could be seen as evidence for being a center of origin for evolution of new species. Looking at species composition in the cen- tral Pacific islands (data from MARCUS 1965; JOHNSON & BOUCHER 1983; CARLSON & HOFF 2003), it seems more likely that they are “traps”, where species dispersing from the Japanese warm-temperate and from tropical and pos- sibly even warm-temperate Australia can find suitable habitats. The Central Pacific sub-province is the only one with less than 10% endemics, but many unidentified and/or undescribed species are known from this sub- province (CARLSON & Horr 2003; own obs.). Most bio- geographic studies indicate that the triangle consisting of Indonesia, Malaysia and the Philippines and sometimes including Papua New Guinea has the highest number of species (EKMAN 1953; BriGGs 1995, 2005). Information on sacoglossan distributions in the Indo-Malayan sub- province has been collected from numerous sources (e.g. BERGH 1871, 1872, 1905; ELiot 1917; Lin 1986, 1990; GOSLINER 1995; GOSLINER et al. 1996; DEBELIUS 1996; JENSEN 1998a,b, 2003; SWENNEN et al. 2001). The num- ber of sacoglossan species is slightly lower in this sub- province than in the islands of the Central West Pacific, i.e. Marianas, Marshall Islands and Micronesia. This may be a collecting artifact caused by the activities of Carlson and Hoff in Guam and neighboring islands; they record- ed 91 species of which 48 (53%) were identified to species level (CARLSON & Horr 2003). However, the present sim- ilarity analyses also indicate that these islands constitute the center of species diversity. Species described from northern Australia, the South China Sea and southern Japan also occur in the Mariana Islands. The genus Gas- coignella ıs endemic to the Indo-Malayan province (JENSEN 1985; SWENNEN 2001), and the genus Sohgenia is endemic to the Central Pacific islands (HAMATANI 1991). However, with one exception, these species have been de- seribed within the last 20 years, so they may have wider distributions. BRIGGS (1995) considered the Hawaiian islands a separate region. The Hawaiian islands have a high endemicity (Os- TERGAARD 1955; Kay 1967; present study), but the pres- ent study has shown that they also have a rather high sim- ilarity to the other islands of the Central and South Pacif- ıc (Table 7). The islands of the South Pacific have been rather sporad- ically studied (PEASE 1861, 1866, 1868, 1871; ELtoT 1899; RIsBEC 1928, 1953; BURN 1966a; MILLER 1969; BRODIE & BRODIE 1990; GOSLINER 1995). Hence the total num- ber of species as well as the number of endemics may be considerably higher. GOSLINER & DRAHEIM (1996) estimat- ed that more than 40% of the opisthobranch species from Papua New Guinea are undescribed. For the Fiji Islands 30% of the species have not been identified to species lev- el (BRODIE & BRODIE 1990). For the sacoglossans most of the old species are poorly described and need re-exam- ination. 266 Kathe R. JENSEN: Biogeography of Sacoglossa 4. DISCUSSION Sacoglossans are small, often cryptically colored species and they are therefore difficult to collect. This means that only a few biogeographic provinces have been thorough- ly studied. In recent years international workshops and biodiversity programs have greatly increased the number of sacoglossans known from more remote localities, es- pecially in tropical waters (JENSEN 1985, 1990, 1993b, 1997b,c, 1999, 2003; JENSEN & WELLS 1990; ICHIKAWA 1993; GOSLINER 1995; SWENNEN 1997, 2001; ORTEA & TEMPLADO 1988; ORTEA & ESPINOSA 1996, 2000, 2001, 2002; CABALLER et al. 2006; MUNIAN & ORTEA 1997). The present study has reviewed the existing information about sacoglossan distributions and analyzed the data for en- demicity and similarity among provinces. Although the da- ta are biased with regards to collecting efforts of a few highly active scientists, several patterns have emerged from these analyses. The collection bias is most obvious in the areas where few other observations have been made, e.g. the Brazilian region where basically all collections have been made by the Marcuses. However, the species richness and endemicity does not differ from that of oth- er tropical regions with no collection bias. The same seems to be true for the Japanese fauna, where Baba has de- scribed most of the species. The sacoglossan fauna of the Indian sub-province has been studied by several taxono- mists, but in this case endemicity seems remarkably high. One reason for this is that descriptions have been pub- lished in local journals and therefore overlooked by oth- er taxonomists. Overall the distributions of sacoglossans correspond to the biogeographic regions and provinces identified by BRIGGS (1995). The exceptions have been pointed out above. Some of the provinces identified contain less than 10 species, which means that just one endemic species will Table 8. Number of species and endemicity of the tropical pro- vinces and sub-provinces of the Indo-West Pacific region. Province/ subprovince # species # endemics (%) Western Indian Ocean 27 5 (19) Red Sea 10 2 (20) India s./. 20 10 (50) Indo-Malayan 48 9 (19) Central Pacific 51 4 (8) Ryukyu 33 10 (30) Hawai 25 6 (24) S Pacific 33 6 (18) Northeast Australia 28 2 (7) + Great Barrier Reef North and Northwestern 23 4 (17) Australia yield more than 10% endemicity. Obviously this cannot be used to make conclusions about their distinctiveness. The records listed in existing literature do not usually have longitude and latitude, and many locality names are am- biguous or oblivious. Consequently it has not been pos- sible to construct a “degree-by-degree” plot of species dis- tributions for analysis. Nevertheless, the latitudinal de- crease in species diversity from warm to cold temperate provinces is spectacular. Sacoglossans are dietary special- ists and the majority of species feed on siphonaceous green algae, which are much more abundant and diverse in trop- ical and warm temperate waters. The cold temperate sacoglossan faunas of the North Atlantic as well as the Northeast Pacific Ocean are mostly eurythermic species with wide latitudinal distributions, extending well into neighboring warm water regions; indices of inclusion are over 60% (Tables 3b, 4b). This 1s also seen in Japan (Table 5b) and southern Africa between the tropical western In- dian Ocean and the warm temperate southwestern Africa (Table 7b), but not nearly as pronounced in Australia (Table 6b). This apparently supports Rapoport's rule that species ranges in high latitudes are larger than in low lat- itudes. However, there are great differences between the provinces. The Northeast Pacific coldwater fauna is 43% of the tropical fauna, the Northeast Atlantic boreal fauna is 21% of the combined Lusitanian and Mediterranean fau- na, and the Northwest Atlantic boreal fauna is only 12% of the Caribbean fauna. The number of amphi-Atlantic species is relatively high, especially for the warm-water/tropical faunas (ORTEA et al. 1997; present study): 16 species (about 30%) occur in the Caribbean and the Lusitanian and/or Mediterranean provinces. However, amphi-Atlantic distributions for most of these species have only been recognized in recent years (TEMPLADO et al. 1990; ORTEA et al. 1988, 1998), and it cannot be excluded that human introductions are involved. Contrary to this, only one cold-temperate species, Her- maea vancouverensis, has an amphi-Pacific distribution. Alderia modesta has a circum-boreal/arctic distribution. Elysiella pusilla has recently been recorded from the Mex- ican-Panamanian province (BEHRENS & HERMOSILLO 2005). However, it cannot be excluded that this is a hu- man introduction. The same is true for Ercolania boodleae, a Japanese species, which has been recorded from California (BEHRENS 1991); this may also be a misidentification (TROWBRIDGE 2002). The so-called cir- cum-tropical species, Elysia ornata and Ercolania coerulea, are conspicuously absent from tropical eastern Pacific; they extend from the Caribbean to the South and/or Central Pacific islands. Lobiger souverbii, which has been synonymized with several Indo-West Pacific species (KAY 1964; BABA 1974; GOSLINER 1987a; GOSLIN- ER etal. 1996), and which occurs in the Caribbean as well Bonner zoologische Beitráge 55 (2006) 267 Table 9. Distribution of species of the Oxynoacea, Plakobranchoidea and Limapontioidea in provinces supported by present stu- dy. Region Oxynoacea NE Atlantic Lusitanian Mediterranean NW Atlantic Caribbean NE Pacif. warm + cold Mexican-Panamanian SE Pacific Hawaii Japan cold temp. Japan warm temp. Ryukyu Islands Australia, tropical Australia, warm + cold W Indian Ocean India, Srı Lanka, Maldı. Indo-Malaya Central Pacific SW Pacific — — MOO 0000 Y PUNOOANOWRO no as Mexican-Panamanian province (BEHRENS 1991; Trow- BRIDGE 2002), may be the only truly circum-tropical species. Very few species are shared between the Caribbean and Mexican-Panamanian provinces, but there is probably a sister-taxon relationship between the East Pa- cific Elysia diomedea (formerly Tridachiella diomedea) and the Caribbean £. crispata (formerly Tridachia crispa- ta) and E. clarki. Thus the Central American land bridge is as effective a geographical separation as the natural land barrier between the Red Sea and the Mediterranean. Al- so the few species occurring on both sides of the Isth- mus of Panama may have arrived on the Pacific side as hitch-hikers on ships travelling through the Panama Canal. The subdivision of the Indo-Polynesian province used in the present study may seem rather arbitrary. In many stud- ies Papua New Guinea is considered a part of the Indo- Malayan sub-province, whereas it has been included in the South Pacific sub-province here. Only 14 species have been identified from Papua New Guinea, which is obvi- ously an underrepresentation; GOSLINER (1992) gives 61 species for this area, but most are undescribed or have not been identified to species. Only sacoglossans that have been identified to species level can be used for calculat- ing similarities between different sub-provinces. The 14 species are almost all widespread species found in the In- do-Malayan as well as the Central Pacific sub-provinces. The one species that does not occur in these sub-provinces, Elysia expansa, has its main distribution in tropical Aus- tralia. Geologically Papua New Guinea has been associ- ated with the Australian continent (BRIGGS 1995). Plakobranchoidea Limapontioidea | 10 10 26 1] 24 2 4 25 23 | 9 6 12 2 3 13 5 11 12 18 14 15 11 20 14 10 11 18 8 8 4 29 14 25 16 17 4 The number of species decreases with longitude to the west of the Central Pacific islands, from 51 through 48 in the Indo-Malayan sub-province to 20 in the Indian sub- province and 27 in the Western Indian Ocean province. The majority of species in the latter province have wide- spread distributions, whereas less than half the species in the Indian sub-province belong in this category. A less pro- nounced longitudinal decrease is seen from 33 species in the South Pacific islands through 28 in northeastern Aus- tralia to 23 in northwestern Australia. The number of species of the Central and South Pacific islands, including Ryukyu, is at least 50% higher than the number of species in tropical East Pacific. This is in spite of the fact that the combined area of these islands is rel- atively small. On the other hand the endemicity of these islands 1s relatively low except in the Ryukyu Islands (Table 8). This means that insular isolation has not result- ed in extensive speciation (or that subsequent dispersal has obscured such speciation). The high endemicity of Ryukyu is almost entirely attributable to one fairly recent publication: ICHIKAWA (1993). Looking at distribution within subordinal taxa of the Sacoglossa, it is obvious that the shelled suborder Oxynoacea, which all feed exclusively on algae of the genus Caulerpa, 1s restricted by the distribution of this al- ga. In the North Atlantic shelled sacoglossans occur in Bermuda, but there is a questionable record of Ascobul- la fragilis from the northern part of Spain (see above). In the East Pacific shelled species are found in the Panaman- 268 Kathe R. JENSEN: Biogeography of Sacoglossa ian region, but also in the southernmost part of the Gulf of California. In the West Pacific shelled species occur in the warm temperate region. The only places where shelled species occur in cold temperate waters are Victoria, south- eastern Australia and the west coast of central Japan. These places also have species diversity of Caulerpa rivaling many tropical and warm temperate provinces (DOUMENGE 1995; PRUD” HOMME VAN REINE et al. 1996). The two non- shelled superfamilies, Plakobranchoidea and Limapon- tioidea, occur in all provinces where sacoglossans are found. At the family level, the bivalved Juliidae are absent from the eastern Atlantic. Since they are common as fossils from this region, this must be due to extinction. Juliidae and Vol- vatellidae are rare in the Atlantic and eastern Pacific. On the other hand, Limapontioidea are highly diverse in the northeastern Atlantic and Mediterranean (Table 9). The least studied areas have the fewest Limapontioidae. This is probably due to the very small size (<10 mm) of most of these species. In tropical regions the number of Plako- branchioidea is always higher than the number of species in the other superfamilies. Plakobranchoidea are most di- verse in the Caribbean region and in the Indo-Malayan and Central Pacific islands sub-provinces; tropical Australia also has a high proportion of this superfamily (Table 9). Only rarely are more than two or three species of one genus found in one area. However, for the genera Elysia and Thuridilla many places have more than 5 species, and in a few cases more than 10 species may co-occur, though on different food algae. Whether this ıs due to sym- or parapatric speciation or dispersal following allopatric spe- ciation, as suggested by GOSLINER (1995), cannot be de- ducted from existing information. Phylogenetic analysis, preferably including molecular data should be applied to these genera!). Some genera are restricted to one or a few neighboring biogeographic provinces. Limapontia only occurs in the Northeast Atlantic and Mediterranean region. This may al- so be the case for the genus Calliopaea, though the Japan- ese Stiliger pusillus has been tentatively assigned to this genus (BABA & HAMATANI 1970). The monotypic Platy- hedyle only occurs in the Mediterranean (WAWRA 1979), whereas the genus Gascoignella, which has been assigned to the same family, Platyhedylidae (JENSEN 1996a), seems restricted to the South China Sea (JENSEN 1985; SWENNEN 2001). The monotypic genus Olea ıs restricted to the Northeast Pacific (TROWBRIDGE 2002). The genus Bosel- lia may have its natural distribution restricted to tropical and warm temperate Atlantic and Mediterranean (MAR- cus 1973; FERNANDEZ-OVIES & ORTEA 1986), and the monotypic genus Sohgenia has only been found in the Central Pacific islands (HAMATANI 1991). Common to all of the above genera is that they have morphological char- acters that appear to be reduced compared to other gen- era in the same superfamilies; they either lack or have very reduced rhinophores, cerata or parapodia (or a combina- tion of these). The genera Roburnella, Plakobranchus and Pattyclaya are Indo-west Pacific endemics, the former fur- thermore restricted to southern Australia, and Julia and Elysiella also occur in the eastern Pacific. Within the more speciose genera, many have few (1-3) representatives in the Atlantic Ocean, zero in the East Pacific, and the re- maining species are Indo-West Pacific. This is seen in Vol- vatella, Thuridilla and Costasiella, indicating that the At- lantic fauna is a Tethyan relict fauna with little subsequent speciation, except in the genus E/ysia in the Caribbean. However, the high number of limapontioid species seems to contradict this. It is possible that the lack of plakobran- choids first spurred a burst of speciation of limapontioids in the East Atlantic/Mediterranean, whereas the speciation in Caribbean Elysía may be a more recent phenomenon. The present study is a preliminary analysis of biogeo- graphic affinities of sacoglossan opisthobranchs. Never- theless several patterns have been identified, which should be further investigated using phylogenetic analyses. Al- so, some obvious gaps in the existing knowledge as well as conflicting and/or questionable records have been iden- tified. There is an urgent need to describe the undescribed species known from the Indo-West Pacific region and to collect sacoglossans from poorly studied regions. 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Marine Molluscs, part 2, Opisthobranchia. University of Auckland, Leigh Marine Lab- oratory: Leigh, 106pp. Yokes, B. 2001. Elysia tomentosa in the Mediterranean? Mes- sage in Sea Slug Forum (http://www.seaslugforum.- net/find.cfm?id=5676) (last access 12th of August 2007). YOKES, B. 2002. Elysia ornata? from Turkey. Message in Sea Slug Forum Re a O22) (last access 12" of August 2007). Author’s address: Zoological Museum, Univer- sitetsparken 15, DK-2100 Copenhagen ©, Denmark. E- mail: krjensen@@snm.ku.dk. Bonner zoologische Beitráge 55 (2006) 27 in APPENDIX List of nominal species of recent Sacoglossa, authorship and type localities. Species are arranged alphabetically within families and superfamilies. Valid species (and species for which synonymy has been contested) are listed with their current generic name. Species considered invalid in the present study are marked with an *. Species Oxynoacea Volvatellidae Ascobulla californica Ascobulla fischeri Ascobulla fragilis Ascobulla japonica Ascobulla? pusilla Ascobulla souverbiei Ascobulla? systremma* Ascobulla ulla Ascobulla? turtoni* Volvatella angeliniana Volvatella australis Volvatella ayakii Volvatella bermudae Volvatella candida Volvatella cincta Volvatella elioti Volvatella evansi Volvatella ficula Volvatella fragilis Volvatella kawamurai* Volvatella laguncula Volvatella omega Volvatella pyriformis Volvatella ventricosa Volvatella vigourouxi Volvatella viridis _ Juliidae Berthelinia (Midorigai) australis Berthelinia (Tamanovalva) babai Berthelinia caribbea Berthelinia chloris Berthelinia corallensis* Berthelinia darwini Berthelinia (Tamanovalva) fijiensis Berthelinia ganapati Berthelinia (Tamanovalva) limax Berthelinia pseudochloris Berthelinia rottnesti Berthelinia schlumbergeri Berthelinia (Edenttellina) typica Berthelinia waltairensis Julia borbonica* Julia burni Julia cornuta* Julia equatorialis* Julia exquisita Julia japonica Julia mishimaensis Julia thecaphora Julia zebra _ Oxynoidae Icarus gravesii* Lobiger corneus* Author (Hamatani, 1971) (Adams & Angas, 1864) (Jeffreys, 1856) (Hamatani, 1969) (Nevill & Nevill, 1869) (Montrouzier in Souverbie & Montrouzier, 1874) (Melvill, 1918) (Marcus & Marcus, 1970) (Bartsch, 1915) Ichikawa, 1993 Jensen, 1997 Hamatani, 1972 Clark, 1982 Pease, 1868 Nevill & Nevill, 1869 (Evans, 1950) (Kay, 1961) Burn, 1966 Pease, 1860 Habe, 1946 Sowerby, 1894 (Melvill, 1918) Pease, 1868 Jensen & Wells, 1990 (Montrouzier, 1861) Hamatani, 1976 (Burn, 1960) (Burn, 1965) Edmunds, 1963 (Dall, 1918) Hedley, 1920 Jensen, 1997 (Burn, 1966) Sarma, 1975 (Kawaguti & Baba, 1959) Kay, 1964 Jensen, 1993 Dautzenberg, 1895 (Gatliff & Gabriel, 1911) Sarma, 1975 (Deshayes, 1863) Sarma, 1975 (De Folin, 1867) Pilsbry & Olsson, 1944 Gould, 1862 Kuroda & Habe, 1951 Kawaguti & Yamasu, 1982 (Carpenter, 1857) Kawaguti 1981 Forbes, 1844 Mörch, 1863 Type locality Gulf of California "South Australia" Mediterranean Kaui, Middle Japan Sri Lanka New Caledonia Gulf of Oman E of Santos, Brazil South Africa Sesoko Isl., Ryukyu Darwin Harbour, N Australia Kaui, Middle Japan Bermuda French Polynesia Sri Lanka Zanzibar Oahu, Hawaii Fiji Sandwich Islands Japan Port Elizabeth, S Africa Gulf of Oman Huaheine, French Polynesia Albany, SW Australia New Caledonia Amami Islands, Japan Torquay, Victoria, Australia Torquay, Victoria, Australia Port Royal, Jamaica Baja California, W Mexico Queensland, Australia Darwin Harbour, N Australia Fiji SE India Seto, Japan Kauai, Hawaii Rottnest Island, SW Australia Madagascar Port Phillip, Victoria, Australia SE India Reunion Andaman Islands, India Mauritius N of Mancara, Peru Hawaiian Islands Wakayama, Honshu, Japan Yamaguchi Pref. Japan Mazatlan, Mexico(?) Yamaguchi Pref. Japan Aegean Sea ? (Cuming collection) 276 Lobiger cumingi* Lobiger nevilli Lobiger pellucidus* Lobiger philippi* Lobiger pilsbryi* Lobiger picta* Lobiger sagamiensis Lobiger serradifalci Lobiger souverbii Lobiger viridis Lobiger viridis* Lophocercus krohnii* Lophocercus sieboldii* Lophopleurella capensis Oxynoe aguayoi* Oxynoe antillarum Oxynoe azuropunctata Oxynoe benchijigua Oxynoe brachycephalus* Oxynoe delicatula Oxynoe hargravesi* Oxynoe kabirensis Oxynoe natalensis* Oxynoe olivacea Oxynoe panamensis Oxynoe viridis Roburnella wilsoni Plakobranchacea Plakobranchidae Actaeon elegans* Aplysiopterus neapolitanus* Elisia marmorata* Elysia abei Elysia albomarginata* Elvsia amakusana Elysia atroviridis Elysia australis Elysia babai Elysia bangtawaensis Elysia bedeckta* Elysia bella? Elysia bennettae Elysia canguzua Elysia catulus Elysia cauze* Elysia chilkensis Elvsia chitwa Elysia chlorotica Elvsia clarki Elvsia clena* Elysia coodgeensis Elysia cornigera* Elysia crispata Elysia cyanea* Elysia degeneri Elysia diomedea Elysia dubia* Elysia duis* Elysia elsiae* Elysia eugeniae Elvsia evelinae Elysia kushimotoensis Elysia expansa Elysia faustula* Kathe R. Jensen: Biogeography of Sacoglossa (A. Adams, 1850) Pilsbry, 1896 A. Adams, 1854 Krohn, 1847 Schwengel, 1941 Pease, 1868 Baba, 1952 (Calcara, 1840) Fischer, 1856 Pease, 1863 Nevill & Nevill, 1869 A. Adams, 1854 Krohn, 1847 (Thiele, 1912) Jaume, 1945 Mörch, 1863 Jensen, 1980 Ortea, Moro & Espinosa, 1999 Mörch, 1863 Nevill & Nevill, 1869 Adams, 1872 Hamatanı, 1980 Smith, 1903 Rafinesque, 1814 Pilsbry & Olsson, 1943 (Pease, 1861) (Tate, 1889) de Quatrefages, 1844 delle Chiaje, 1830 Cantraine, 1835 Baba, 1955 Trinchese, 1869 Baba, 1955 Baba, 1955 (Quoy & Gaimard, 1832) Pruvot-Fol, 1946 Swennen, 1997 MacFarland, 1966 (Pease, 1860) Thompson, 1973 Marcus, 1955 (Gould, 1870) Marcus, 1957 Eliot, 1916 Marcus, 1955 Gould, 1870 Pierce et al., 2006 Marcus & Marcus, 1970 (Angas, 1864) Nuttall, 1989 Mörch, 1863 Mamo ın Caruana, 1867 Ostergaard, 1955 (Bergh, 1894) Eliot, 1904 Marcus & Marcus, 1967 Ostergaard, 1955 Ortea & Espinosa, 2002 Marcus, 1957 Baba, 1957 (O'Donoghue, 1924) Bergh, 1872 Puerto St. Elena, W Colombia n.n. for L. viridis G. &. H. Nevill, 1869 unknown (Cuming collection) Sicily, Italy Sanibel Isl., Florida Huaheine, French Polynesia Sagamı Bay, Japan Sicily, Italy Guadeloupe Huahine Sri Lanka Hawaiian Islands (Sandwich Islands) Messina, Italy S Africa Cuba St. Thomas, USVI Florida Canary Islands Based on figure Sri Lanka New Hebrides Ishigaki Isl., Ryukyu Islands Port Elizabeth, S Africa Sicily, Italy Bocas Isl., Panama Sandwich Islands Port Phillip Bay, Victoria, Australia St. Vaast, France Napoli, Italy Livorno, italy Sagami Bay, Japan Italy Sagami Bay, Japan Sagami Bay, Japan Port Jackson, Sydney, NSW Ryukyu Islands Pattani, Gulf of Thailand Monterey Bay, California Hawaii Heron Island, GBR NE of Santos, Brazil Massachusetts, USA NE of Santos, Brazil Chilka Lake, India NE of Santos, Brazil Massachusetts, USA Florida ?Curacao/? Barbados Port Jackson, Sydney, NSW Spanish Harbor Key, FL St. Croix, USVI Malta Oahu, Hawaii Lower California Zanzibar Biscayne Bay, Florida Waikiki, Hawaii Manzanillo, Costa Rica (Carib) NE of Santos, Brazil Kushimoto, Kai, Japan Abrolhos Islands, NW Australia Masoloc, Philippines Elysia fezi Elysia filicauda Elysia flava Elysia flavipunctata Elysia flavomacula Elysia furvacauda Elysia fusca* Elysia gordanae Elysia grandifolia Elysia grandis? Elysia haingsisiana? Elysia halimedae* Elysia hamatanii Elysia hedgpethi Elysia hendersoni Elysia hetta Elysia hirasei Elysia japonica Elysia latipes* Elysia leucolegnote Elysia lobata Elysia macnaei* Elysia maoria Elysia margaritae Elysia marginata* Elysia minima Elysia minuta* Elysia nealae Elysia nigrocapitata Elysia nigropunctata Elysia nisbeti Elysia obtusa Elysia ocellata* Elysia oerstedii Elysia ornata Elysia ornata* Elysia pagenstecheri* Elysia papillosa Elysia patagonica Elysia patina Elysia pilosa Elysia pratensis Elysia pruvotae* Elysia punctata Elysia purchoni Elysia rufescens Elysia serca Elysia setoensis Elysia siamensis Elysia slimora Elysia splendida* Elysia subornata Elysia sugashimae Elysia thompsoni Elysia timida Elysia tokarensis Elysia tomentosa Elysia translucens Elysia trilobata Elysia trisinuata Elysia tuca Elysia (Elysiella) verrilli* Elysia verrucosa Bonner zoologische Beiträge 55 (2006) Vilella, 1968 Jensen & Wells, 1990 Verrill, 1901 Ichikawa, 1993 Jensen, 1990 Burn, 1958 Philippi, 1844 Thompson & Jaklın, 1988 Kelaart, 1857 Bergh, 1872 Bergh, 1905 Macnae, 1954 Baba, 1957 Marcus, 1961 Eliot, 1899 Perrone, 1990 Baba, 1955 Eliot, 1913 Marcus, Er. & Marcus, Ev. 1960 Jensen, 1990 Gould, 1852 Ev. Marcus, 1980 Powell, 1937 Fez, 1962 (Pease, 1871) Ichikawa, 1993 (Sars, 1835) Ostergaard, 1955 Baba, 1957 (Pease, 1871) Thompson, 1977 Baba, 1938 Pease, 1860 Morch, 1859 (Swainson, 1840) (Pease, 1860) Marcus, Ev., 1982 Verrill, 1901 Munian & Ortea, 1997 Marcus, 1980 Risbec, 1928 Ortea & Espinosa, 1996 Risbec, 1953 Kelaart, 1857 Thompson, 1977 (Pease, 1871) Marcus, 1955 Hamatanı, 1968 Swennen, 1997 Marcus & Marcus, 1966 Grube, 1861 Verrill, 1901 Baba, 1955 Jensen, 1993 Risso, 1818 Baba, 1957 Jensen, 1997 Pruvot-Fol, 1957 Heller & Thompson, 1983 Baba, 1949 Marcus & Marcus, 1967 Thiele, 1931 Jensen, 1985 277 Barcelona, Spain Albany, SW Australia Bermuda Ishigakı Isl., Ryukyu Hong Kong Torquay, Victoria, Australia Palermo, Italy Rovinj, Yugoslavia Sri Lanka Palau? (Pelew) Haingsisi Port Alfred, S Africa Seto, Japan Tomales Bay, California Samoa Gulf of Taranto, Italy Sagami Bay, Japan Japan Maldives Hong Kong Hawaii Japan?/Australia? Auckland, NZ Valencia, Spain Huaheine, French Polynesia Kuro Isl., Ryukyu Bergensund, Norway Waikiki, Hawaii ?Osaka/?Seto/?Tsurugu/? Toyama Bay Japan Tahiti Jamaica Seto, Japan Hawaii Puntarenas, Central America West Indies Hawaii Sete, Mediterranean Bermuda San Jorge Gulf, Argentina (45d58'S; 67d34'W) Florida Keys New Caledonia Eastern part of Yucatan, Mexico New Caledonia Sri Lanka Jamaica Tahiti NE of Santos, Brazil Seto, Japan Pattani, Gulf of Thailand near Sao Tome, W Africa Cherso, N Adriatic Bermuda Sagami Bay, Japan Rottnest Island, SW Australia Nice, France Mediterr. Tokara Islands, Kyushu, Japan Abrolhos Islands, NW Australia Banyuls, France Mediterr. Red Sea Sagami Bay, Japan Biscayne Bay, Florida n.n. for Elysia (Elysiella catula Verrill Hong Kong 278 Elysia viridis Elysia viridissima* Elysia vreelandae Elysia yaevamana Elysia zuleicae Elysiella pusilla Elysiella stylifera Elysiobranchus mercieri Elysiobranchus ryukyuensis Pattyclaya arena Pattyclaya brycei Placobranchus argus* Placobranchus caminguinus* Placobranchus chlorophagus* Placobranchus gracilis* Placobranchus guttatus* Plakobranchus ianthobapsus* Placobranchus laetus* Plakobranchus ocellatus Placobranchus priapinus* Placobranchus punctulatus* Placobranchus variegatus* Thuridilla albopustulosa Thuridilla bayeri Thuridilla carlsoni Thuridilla coerulea Thuridilla decorata Thuridilla flavomaculata Thuridilla gracilis Thuridilla hoffae Thuridilla hopei Thuridilla indopacifica Thuridilla kathae Thuridilla lineolata Thuridilla livida Thuridilla mazda Thuridilla moebii Thuridilla multimarginata Thuridilla neona Thuridilla picta Thuridilla ratna Thuridilla splendens Thuridilla thysanopoda> Thuridilla undula Thuridilla vatae Thuridilla virgata Tridachia schrammi* Boselliidae Bosellia cohellia? Bosellia corinneae Bosellia leve Bosellia marcusi Bosellia mimetica Platyhedylidae Gascoignella aprica Gascoignella jabae Gascoignella nukuli Platyhedyle denudata Limapontioidea Kathe R. Jensen: Biogeography of Sacoglossa (Montagu, 1804) Trinchese, 1869 Marcus & Marcus, 1970 Baba, 1936 Ortea & Espinosa, 2002 Bergh, 1872 Jensen, 1997 Pruvot-Fol, 1930 Ichikawa, 1993 Carlson & Hoff, 1978 Jensen & Wells, 1990 Bergh, 1872 Bergh, 1872 Bergh, 1878 Pease, 1871 Stimpson, 1858 Gould, 1852 Bergh, 1872 van Hasselt, 1824 Bergh, 1872 Bergh, 1872 Pease, 1871 Gosliner, 1995 (Marcus, 1965) Gosliner, 1995 (Kelaart, 1857) (Heller & Thompson, 1983) Gosliner, 1995 (Risbec, 1928) Gosliner, 1995 (Verany, 1853) Gosliner, 1995 Gosliner, 1995 (Bergh, 1905) (Baba, 1955) Ortea & Espinosa, 2000 (Bergh, 1888) Gosliner, 1995 Gosliner, 1995 (Verrill, 1901) (Marcus, 1965) (Baba, 1949) (Bergh, 1905) Gosliner, 1995 (Risbec, 1928) (Bergh, 1888) Mörch, 1863 Marcus, 1978 Marcus, 1973 Fernandez-Ovies & Ortea, 1986 Marcus, 1972 Trinchese, 1890 Jensen, 1985 Swennen, 2001 Swennen, 2001 Salvini-Plawen, 1973 Polybranchidae (FCaliphyllidae) Caliphvlla mediterranea Caliphylla tricolor* Cyerce antillensis (A. Costa, 1867) Trinchese, 1879 Engel, 1927 Devonshire, UK Italy W Mexico Ishigakishima, Ryukyu Cuba Aibukit, ?Palau (Palaos) Darwin Harbour, N Australia New Caledonia Sesoko Isl., Ryukyu Guam Albany, SW Australia Honolulu, Hawaii Luzon, Philippines Huaheine Tahiti Ryukyu Islands Honolulu, Hawaii Masoloc, Philippines Sunda Strait Bohol, Philippines Masoloc, Philippines Huaheine, French Polynesia Madang, PNG Marshall Islands Madang, PNG Sri Lanka Red Sea Luzon, Philippines New Caledonia Madang, PNG Nice, France Mediterr. Aldabra Atoll Madagascar Saleyer Sagami Bay, Japan Mauritius Midway Atoll Lanai, Hawaii Bermuda Palau Sagami Bay, Japan Tual, Kei Islands, Indonesia Madang, PNG New Caledonia Mauritius Guadeloupe ?Red Sea/?Mediterranean Key Biscayne, Florida Lanzarote, Canary Islands Grassy Key & Miami, FL Capri, Italy Hong Kong Pattani, Gulf of Thailand Pattani, Gulf of Thailand Livorno, italy Napoli, Italy Mediterranean Tobago(?), Westindien Manzanillo, Costa Rica (Carib) Cyerce cristallina Cyerce? edmundsi Cyerce elegans Cyerce graeca Cyerce habanensis Cyerce jheringi* Cyerce kikutarobabai Cyerce nigra Cyerce nigricans Cyerce orteai Cyerce pavonina Cyerce verdensis Mourgona germaineae Mourgona murca Mourgona osumi Polybranchia borgnini Polybranchia orientalis Polybranchia pallens Polybranchia papillosa Polybranchia pellucida Polybranchia prasinus Polybranchia rubicundus* Polybranchia viridis Polybranchia westralis Sohgenia palauensis _ Hermaeidae Aplysiopsis brattstroemi Aplysiopsis elegans Aplysiopsis enteromorphae Aplysiopsis formosa Aplysiopsis maculosa* Aplysiopsis minor Aplysiopsis nigra Aplysiopsis orientalis Aplysiopsis sinusmensalis Aplysiopsis smithi* Aplysiopsis toyamana Aplysiopsis zebra* Hermaea bifida Hermaea boucheti Hermaea carminis* Hermaea coirala Hermaea cruciata Hermaea evelinemarcusae Hermaea hillae Hermaea lutescens* Hermaea minor* Hermaea noto Hermaea oliviae Hermaea paucicirra Hermaea polychroma* Hermaea vancouverensis Hermaea variopicta Hermaea venosa* Hermaea wrangeliae Hermaea zosterae Physopneumon carneum* Limapontiidae Alderella comosa Alderia harvardiensis* Alderia modesta Alderia scaldiana* Alderia uda Alderia willowi Bonner zoologische Beitráge 55 (2006) (Trinchese, 1881) Thompson, 1977 Bergh, 1871 Thompson, 1988 Ortea & Templado, 1988 Pelseneer, 1892 Hamatani, 1976 Bergh, 1871 (Pease, 1866) Valdes & Camacho, 2000) Bergh, 1888 Ortea & Templado 1990 Marcus & Marcus, 1970 Marcus & Marcus, 1970 Hamatani, 1994 (Trinchese, 1895) (Kelaart, 1858) (Burn, 1957) (Pease, 1866) Pease, 1860 (Bergh, 1871) (Bergh, 1871) (Deshayes, 1857) Jensen, 1993 Hamatani, 1991 (Marcus, 1959) (Deshayes, 1835) (Cockerell & Eliot, 1905) Pruvot-Fol, 1953 (Trinchese, 1874) (Baba, 1959) (Baba, 1949) (Baba, 1949) (Macnae, 1954) (Marcus, 1961) (Baba, 1959) Clark, 1982 (Montagu, 1816) Cervera, Garcia-Gomez & Ortea, 1991 Fez, 1962 Marcus, 1955 Gould, 1870 Jensen, 1993 Marcus & Marcus, 1967 A. Costa, 1866 Bergh, 1888 (Baba, 1959) (MacFarland, 1966) Pruvot-Fol, 1953 (Hesse, 1873) O'Donoghue, 1924 (A. Costa, 1869) Lovén, 1845 (Ichikawa, 1993) (Baba, 1959) A. Costa, 1862 (Costa, 1867) Gould, 1870 (Lovén, 1844) Nyst, 1855 Marcus, Ev. & Marcus, Er., 1956) Krug, Ellingson, Burton & Valdés, 2007 279 Napolı, Italy Jamaica Palau? (Palaos) Greek Ionian Sea Cuba Napoli, Italy Amami Islands, Japan Palau? (Palaos) Pacific Islands Puntarenas, Costa Rica Mauritius Cape Verde Puerto Rico (aquarium) Curacao Amami-Oshima Island, SW Japan Mediterranean Sri Lanka Queenscliff, Victoria, Australia Pacific Islands Hawaii Luzon, Philippines Tor, Red Sea Guadeloupe Rottnest Island, SW Australia Palau Chile (23d39'S; 70d25'W) Banyuls, France Mediterr. San Pedro, California(?) Temara, Marocco Genova, Italy Toyama Bay, Japan Sagami Bay, Japan Sagami Bay, Japan Table Bay, S Africa Tomales Bay, California Toyama Bay, Japan Florida Devonshire, UK Pontevedra, SW Spain Valencia, Spain NE of Santos, Brazil Massachusetts, USA Rottnest Island, SW Australia Sonora, W Mexico Napoli, Italy Mauritius ?Noto Peninsula/? Toyama Bay Japan Monterey Bay, California Marocco Brest Vancouver Isl., Canada Napoli, Italy Bohuslan, Sweden Kuro Isl., Ryukyu Amakusa, Japan Mediterranean Napolı, Italy Massachusetts, USA Bohuslän, Sweden Scheldt estuary, Netherlands SW of Santos, Brazil San Pedro, California 280 Alderiopsis garfio Alderiopsis nigra Calliopaea bellula Calliopaea oophaga Calliopaea souleyeti* Cenia cocksii* Cenia corrugata* Chalidis coeruleus* Chalidis nigricans* Costasiella formicaria Costasiella illa Costasiella iridophora Costasiella kuroshimae Costasiella lilianae* Costasiella mandurahae Costasiella nonatoi Costasiella ocellifera Costasiella pallida Costasiella paweli Costasiella rubrolineata Costasiella usagi Costasiella vegae Costasiella virescens Custiphorus vesiculosus* Embletonia mariae* Ercolania boodleae Ercolania coerulea Ercolania costai* Ercolania cricetae* Ercolania evelinae Ercolania emarginata* Ercolania endophytophaga Ercolania erbsus Ercolania felina Ercolania funerea* Ercolania fuscata Ercolania gopalai Ercolania irregularis Ercolania lozanoi Ercolania margaritae Ercolania nigra Ercolania nigrovittata* Ercolania nigrevittata* Ercolania pancerii* Ercolania pica Ercolania raorum Ercolania selva Ercolania siottii Ercolania subviridis Ercolania talis* Ercolania tentaculata Ercolania translucens Ercolania trinchesii* Ercolania uziellii* Ercolania vanellus* Ercolania varians Ercolania viridis Ercolania zanzibarica Limapontia capitata Limapontia cornuta* Limapontia depressa Limapontia nigra* Limapontia senestra Limapontia zonata! Olea hansineensis Kathe R. JENSEN: Biogeography of Sacoglossa Caballer, Ortea & Espinosa, 2006 (Baba, 1937) d'Orbigny, 1837 Lemche, 1974 Verany, 1853 Alder & Hancock, 1848 Alder & Hancock, 1848 de Quatrefages, 1844 Alder & Hancock, 1847 (Baba, 1959) (Marcus, 1965) Ichikawa, 1993 Ichikawa, 1993 (Marcus, Ev. & Marcus, Er., 1969 Jensen, 1997 Marcus & Marcus, 1960 (Simroth, 1895) Jensen, 1985 Ichikawa, 1993 Ichikawa, 1993 Ichikawa, 1993 Ichikawa, 1993 Pruvot-Fol, 1951 Deshayes, 1853 Meyer & Mobius, 1865 (Baba, 1938) Trinchese, 1892 Pruvot-Fol, 1951 (Marcus & Marcus, 1970) (Marcus, 1959) Jensen, 1985 Jensen, 1999 (Marcus & Marcus, 1970) (Hutton, 1882) (Costa, 1867) (Gould, 1870) (Rao, 1937) (Eliot, 1904) Ortea, 1981 Burn, 1974 (Lemche, 1936) (A. Costa, 1866) (Rao & Rao, 1963) Trinchese, 1872 (Annandale & Prashad, 1922) (Marcus & Marcus, 1970) Ortea « Espinosa, 2001 Trinchese, 1872 (Baba, 1959) (Marcus & Marcus, 1956) (Eliot, 1917) Jensen, 1993 Pruvot-Fol, 1951 Trinchese, 1872 Marcus, 1957 (Eliot, 1904) (A. Costa, 1866) Elıot, 1903 (Múller, 1773) Giard, 1873 Alder & Hancock, 1862 Johnston, 1836 (de Quatrefages, 1844) (Girard, 1852) Agersborg, 1923 Cuba Amakusa, Japan France Samso, Kattegat, DK Nice. France Mediterr. Falmouth, UK Falmouth, UK Ile de Bréhat, France Falmouth, UK Amakusa, Japan Caroline Islands, Micronesica Kuro Isl., Ryukyu Kuro Isl., Ryukyu NE of Santos, Brazil Darwin Harbour, N Australia NE of Santos, Brazil Bermuda Hong Kong Miyako Isl., Ryukyu Ishigaki Isl., Ryukyu Ishigaki Isl., Ryukyu Ishigaki Isl., Ryukyu Banyuls, France Mediterr. 0) Kieler Bucht Seto, Japan Napoli, Italy ?Banyuls/?Monaco, Mediterranean Curacao Chile (53d22'S; 70d57'W) Hong Kong Rottnest Island, SW Australia Madagascar New Zealand Napoli, Italy Massachusetts, USA Madras, E India Zanzıbar Tenerife, Canary Islands Point Lonsdale, Victoria, Australia Nyborg Fjord, DK Napoli, Italy Gulf of Mannar, SE Indıa Genova, Italy Chilka Lake, India Gulf of Mannar, SE India Manzanillo, Costa Rica (Carib) Genova, Italy Toyama Bay, Japan SW of Santos, Brazil Singgora, SE Thailand Rottnest Island, SW Australia ?Banyuls/?Monaco, Mediterranean Genova, Italy E of Santos, Brazil Zanzibar Napoli, Italy E Zanzibar Baltic Sea 9 Sunderland, UK Berwick Bay, UK Ile de Bréhat, France Massachusetts, USA Friday Harbor, Washington Bonner zoologische Beiträge 55 (2006) (Powell, 1937) Marcus, Ev., 1982 (A. Costa, 1867) Macnae, 1954 (Trinchese, 1892) Jensen, 1990 (Pruvot-Fol, 1953) (Alder & Hancock, 1843) Burn, 1966 Thompson, 1977 (MacFarland, 1966) (Thompson, 1988) (Trinchese, 1873) Placida aoteana* Placida babai* Placida brevicornis Placida capensis* Placida cremoniana Placida daguilarensis Placida dakariensis Placida dendritica Placida fralila Placida kingstoni Placida ornata* Placida saronica Placida tardyi Placida verticillata Ortea, 1981 Placida viridis (Trinchese, 1873) Stiliger akkeshiensis Baba, 1935 Stiliger aureomarginatus Jensen, 1993 Stiliger berghi Baba, 1937 Lance, 1962 Ortea, 1981 Baba, 1959 Baba, 1949 Ehrenberg, 1828 Marcus € Marcus, 1960 (Kelaart, 1858) Stiliger fuscovittatus Stiliger llerai Stiliger pusillus Stiliger smaragdinus Stiliger ornatus Stiliger vossi Stiliger? viridis? IThis species is probably a flatworm rather than a sacoglossan. “The taxonomic status of this species is so uncertain that it has been omitted from the analyses. 3This species has been omitted from the analyses due to lack of information. 4Since the type locality is uncertain, this species has been omitted from the analyses. Auckland, NZ Seto, Japan Napoli, Italy Cape Province, S Africa Napolı, Italy Hong Kong Dakar, Senegal Torbay, UK Queensland, Australia Jamaica Monterey Bay, California Greek Aegean Sca Genova, Italy Tenerife, Canary Islands Genova, Italy Akkeshi Bay, Hokkaido, Japan Rottnest Island, SW Australia Tomioka, Amakusa, Japan San Diego, California Tenerife, Canary Islands Osaka Bay, Japan Sagamı Bay, Japan Red Sea Upper Florida Keys Sri Lanka Bonner zoologische Beiträge Band 55 (2006) Heft 3/4 Seiten 283-290 | Bonn, November 2007 Re-description of a female Pontohedyle brasilensis (Rankin, 1979), a junior synonym of the Mediterranean P. milaschewitchii (Kowalevsky, 1901) (Acochlidia, Gastropoda)* Katharina M. JÖRGER, Timea P. NEUSSER & Michael SCHRODL!) DZoologische Staatssammlung Múnchen, Munich, Germany *Paper presented to the 2nd International Workshop on Opisthobranchia, ZFMK, Bonn, Germany, September 20th to 22nd, 2006 Abstract. Currently 27 species are considered to be valid in the still enigmatic opisthobranch group of the Acochlidia. The taxonomic status of the acochlidian species, Pontohedyle brasilensis (Rankin, 1979), remained unclear due to a lack of primary data. The present study provides the first structural and some histological data on a female P. brasilensis from northern Brazil. The female genital system 1s reconstructed 3-dimensionally from serial semithin sections using AMIRA software. Our new results are compared with published data on a male P brasilensis from southern Brazil and on P mi- laschewitchii (Kowalevsky, 1901) from the Mediterranean and Black Sea; in absence of morphological differences we consider P. brasilensis as a junior synonym. The genus Pontohedyle thus comprises two valid species, the Atlantic/Me- diterranean P. milaschewitchii and the tropical Indopacific P verrucosa (Challis, 1970). Both possess unique bow-sha- ped, flattened oral tentacles, which are diagnostic for the genus and, thus, a probable autapomorphy. Keywords. Mollusca, Opisthobranchia, taxonomy, morphology, anatomy, 3D reconstruction. 1. INTRODUCTION Currently, 27 nominal acochlidian species are considered to be valid (SOMMERFELDT & SCHRÖDL 2005) which were conventionally classified into 12 different genera in 6 fam- ilies (WAWRA 1987). All acochlidians have a characteris- tic shell-less body with a head-foot complex that can be at least partly retracted into a more or less elongate vis- ceral hump. Most species belong to tiny members of worldwide coastal mesopsammic communities, while oth- ers are inhabitants of brackish waters or even limnic (see NEUSSER & SCHRÓDL 2007). Uniquely within the usually hermaphroditic opisthobranchs, microhedylid acochlidi- an species have separate sexes. While most acochlidians have two pairs of cephalic tentacles, a few gonochoristic species lack any rhinophores, 1.e. of the genera Ganitus Marcus, 1953 and Pontohedyle Golikov & Starobogatov, 1972. There are three nominal Pontohedyle species: the tropical Indopacific P verrucosa (Challis, 1970), the At- lantic/ Mediterranean species P milaschewitchii (Kowalevsky, 1901), and the Atlantic P. brasilensis (Rankin, 1979) with uncertain taxonomic status. Pontohedyle milaschewitchii was originally described from the Black Sea (KOWALEVSKY 1901) and later found throughout the Mediterranean (see HADL et al. 1969; JÖRG- ER et al. in press: POIZAT 1984; WAWRA 1986). Addition- ally, MARCUS & MARCUS (1954) described one single male specimen of P milaschewitchii from Ilhabela (Sao Paulo State), the coast of southern Brazil. Solely based on that literature information, RANKIN (1979) established the new genus and species Gastrohedyle brasilensis and separat- ed it from the Mediterranean P milaschewitchii (as Man- cohedyle); her diagnosis of P. milaschewitchii then was limited to the original description by KOWALEVSKY (1901). ARNAUD et al. (1986) listed Gastrohedvle brasilen- sis as Pontohedyle brasilensis with a question mark, and WAWRA (1987) regarded it as a probable synonym of P. milaschewitchii, however without giving any discussion on an entire set of putative external and internal morpho- logical differences that were raised by RANKIN (1979). Unfortunately, anatomical information of P. brasilensis is restricted to a single male specimen. This type specimen of P. brasilensis has not been discovered in the Marcus’ collection of the Museu de Zoologia da Universidade de Sao Paulo (C. Magenta, Sao Paulo, pers. comm. 2006), and thus appears to be lost. Specimens of Pentohedyle from Brazil remain very rare. Even after exhaustive search at the original location, MARCUS & MARCUS (1954) were not able to rediscover further specimens. We conducted collections at Ilhabela, the type locality of P. brasilensis, along the coast of Santa Catarina, Paraná and Sao Paulo State, southern Brasil, and at many sites in Pernambuco 284 Katharma M. JÓRGER et al.: Re-description of Pontohedyle brasilensis and Paraíba, northern Brazil. This search only resulted in two specimens, one of them usable for histological analy- sis. The present study provides the first structural and histo- logical data on a female Pontohedyle from northern Brazil. The taxonomy of P. brasilensis is revised by critically comparing our results with the published data on P brasilensis from Brazil and P milaschewitchii from the Mediterranean. 2. MATERIAL AND METHODS Two Pontohedyle specimens were extracted from sand samples (see SCHRÓDL 2006 for method of extraction), col- lected by scuba diving on the northern coast of Brazil (ap- prox. 5 km off Porto de Galinhas, at 20 m depth) in Jan- uary 2004. One retracted and damaged specimen was used for molecular analysis. The posterior part of the visceral hump of the second specimen was also damaged. The specimens were slowly anaesthetised using 7 % isotonic MgCl, solution and fixed in 75 % ethanol. The preserved specimen used for histological analysis in the present study was transferred into Bouin solution for decalcification and afterwards stained with 0.5 % safranin. Then it was de- hydrated by a graded acetone series and embedded in Spurr’s low viscosity epoxy resin (SPURR 1969) for sec- tioning. The epoxy resin block was cut at 1.5 um with a rotation-microtome (Microtom HM 360; Zeiss), using glass knives and contact cement at the lower cutting edge (HENRY 1977) to receive ribboned serial sections. The sec- tions were stained with methylene blue-azure II (see RICHARDSON et al. 1960). Computer based 3D reconstruc- tion of the female genital system was performed with the software AMIRA 3.0 (TGS Template Graphics Software, Inc., USA). The section series was deposited in the Zoo- logische Staatssammlung München (ZSM), Mollusca Sec- tion (ZSM Mol 20041037). For morphological and anatomical comparison, serial sections and 3D reconstruc- tions of five individuals of P milaschewitchii from the Mediterranean were used (ZSM Mol 20060522- 20060525). 3. RESULTS 3.1. External morphology and spicules Our examined living Brazilian specimen used for struc- tural analysis, showed the usual body shape of marine in- terstitial acochlidians, with a cylindrical anterior head-foot complex that is completely retractable into a broadened and elongated visceral hump. The crawling individual measured approximately 2 mm, but the visceral hump was damaged. The overall body coloration was whitish, with the brownish digestive gland shining through the tissue. The oral tentacles were bow-shaped and curved, rhinophores were lacking (see Fig. 3A). The ciliated foot was short, 1.e. there was no free tail extending behind the head-foot complex, and its posterior edge was rounded. Monoaxone (1.e. needle shaped) spicules (about 25 um length) were found randomly distributed over the head- foot complex and visceral hump. Additionally, an accu- mulation of parallel orientated monoaxone spicules was detected between the oral tentacles. Light microscopic in- vestigation of the sectioned head region indicates cilia on the anterior border of the head and oral tentacles. Fig. 1. Semithin sections of the female Pontohedyle from Brazil. (A) Cross-section of the visceral hump, showing the epidermis and the epidermal gland cells. (B) Horizontal section of the central nervous system. Abbreviations: ag accessory ganglia, cg ce- rebral ganglia, dg digestive gland, egl epidermal gland type I, eg2 epidermal gland type Il, ey eyes, mug mucous gland. Bonner zoologische Beiträge 55 (2006) 28 3.2. Microanatomy The bad condition of the only Brazilian specimen avail- able for structural analysis did not allow detailed histo- logical investigations of all major organ systems. Never- theless, a brief treatise is given on the recognisable organs, focusing on the female genital system which could be re- constructed from serial sections. 3.2.1. Epidermal glands The epidermis contains large spherical glandular cells (5- 10 um in diameter). They form a sub-epidermal sac, which is more or less filled with a homogenous whitish secre- tion (= epidermal glands type I, see Fig. 1A). Smaller vac- uoles could be detected in the epidermis containing pink- ish to violet stained granular material. These vacuoles oc- cur in large numbers (= epidermal glands type II, see Fig. 1A). 3.2.2. Central nervous system Praepharyngeal, large oval cerebral ganglia (approximate- ly 50 um), smaller pedal ganglia (approximately 30 um) and groups of accessory ganglia could be detected. The cerebral ganglia are connected by a very strong and short commissure. A pair of dark pigmented eyes (diameter about 12 um) nestles on the anterior side of the cerebral ganglia (see Fig. 1B). Groups of accessory ganglia are lo- cated anteriorly and laterally of the cerebral ganglia. Dif- ferent from true ganglia true ganglia, accessory ganglia are well defined cell groups with a homogenous distribu- tion of nuclei and without subdivision into cortex and medulla (NEUSSER et al. 2006). Here the accessory gan- glia are more or less spherically shaped and grouped to- gether like pearls on a chain (Fig. 1B). 3.2.3. Digestive system The mouth opening is located subterminally between the oral tentacles. The thin walled oral tube is collapsed. The muscular pharynx extends in the posterodorsal part of the head-foot complex and contains the radula in its posteri- or region. The salivary glands form one mass on the left side of the head-foot complex, slightly extending into the visceral sac. The cells of the salivary glands contain dark blue stained granules. The tube-like oesophagus leaves the pharynx posterodorsally and connects to the digestive gland in the anterior region of the visceral hump. There is no histologically or anatomically detectable stomach. The digestive gland extends over the length of the remain- ing visceral hump and extrudes through the ruptured epi- dermis. It is sac-like in shape and its cells contain small dark blue and red stained granules. The epithelium of the digestive gland bears a series of small whitish and oval in vacuoles. Neither the intestine nor the anal opening could be detected due to the bad condition of the animal. 3.2.4. Excretory and circulatory systems Only the kidney could be detected. It is triangular in shape and squeezed in between the digestive gland and the body wall on the right side of the anterior region of the viscer- al hump. The epithelium of the kidney is characterized by its usual vacuolated structure. 3.2.5. Female genital system The examined individual is a mature female, recognisable by the presence of vitellogenic oocytes in the ovary. The female genital system is composed of the ovary, the nida- mental glands and the oviduct (Fig. 2A, B). The ovary ex- truded through the ruptured epidermis of the visceral sac and was partially falling apart. Nevertheless, seven large vitellogenic oocytes are still in situ (Fig. 2F). The oocytes are comprised of a nucleus containing one nucleolus and yolk (characterised by dense aggregations of blue stained granules). The oocytes reach a diameter of about 50-60 um. The albumen gland is tube-like in shape and its se- cretory cells are stained dark blue to dark violet (Fig. 2E). The secretory cells are alternated by supporting cells, which bear cilia. The membrane gland is comparably large and tube-like in shape. Its secretory cells are stained pink- ish with glandular appearance and containing vacuoles (Fig. 2D). The supporting cells bear cilia. The long tube- like mucous gland runs parallel to the digestive gland in the anterior region of the visceral sac. The supporting cells of the mucous gland are also ciliated and the secretory cells are stained dark violet. The three nidamental glands connect directly to each other (i.e. without any defined proximal oviduct or adhesive region, see Fig. 2A). The dis- tal ciliated oviduct (Fig. 2C) ventrally passes the diges- tive gland and leads to the right anterior region of the vis- ceral hump. The genital opening is located on the right side of the body, at the transition from the head-foot complex to the visceral hump. A short ciliated band originates at the genital opening. It has a diameter of about 10 um and runs anteriorly along the right side of the head-foot com- plex. 4. DISCUSSION According to WAWRA (1987), acochlidians belonging to the genus Pontohedyle share microhedylid features such as having separate sexes and lacking copulatory organs. Pontohedyle species were characterized by the absence of rhinophores and a radula formula of 1.1.1. The combina- tion of these features is unique among acochlidians, but may refer to plesiomorphies. The special shape of 286 Katharina M. JÖRGER et al.: Re-description of Pontohedvle brasilensis Fig. 2. Female genital system of the Pontohedvle from Brazil. (A) Schematic overview, lateral view. (B) 3D reconstruction, la- teral right view, specimen retracted, approximately posterior half of visceral sac and gonad missing (ruptured and therefore not re- constructed). (C) Semithin cross-section of oviduct and genital opening. (D) Semithin cross-section of membrane gland. (E) Se- mithin cross-section of the transition from albumen gland to membrane gland. (F) Semithin cross-section of mature oocytes. Ab- breviations: alg albumen gland, cb ciliary band, dg digestive gland, f foot, go genital opening, meg membrane gland, mug mucous gland, n nucleus, nu nucleolus, o oocyte, od oviduct, ot oral tentacle, ov ovary, y yolk. Bonner zoologische Beitráge 55 (2006) 287 acochlidian oral tentacles may provide more phylogenet- ic information. Apart from species of the genus Ponto- hedyle, flat oral tentacles only occur in the genera Hedy- lopsis and Ganitus. While the oral tentacles of the Hedy- lopsis are much broader than those of Pontohedyle (see Fig. 3F), the ones of Ganitus appear similar. Ganitus can be differentiated since the oral tentacles are never tapered towards the end (see Fig. 3E). In fact, the flat, elongated to bow-shaped oral tentacles of Pontohedyle, which are tapered towards the end (see Fig. 3A-D), are unique and diagnostic, and thus, a probable autapomorphy of Ponto- hedyle. WAWRA (1987) regarded two Pontohedyle species as be- ing valid, the tropical Indopacific P. verrucosa (Challis, 1970) and the temperate P milaschewitchii (Kowalevsky, 1901). RANKIN (1979) however, established an addition- al species P. brasilensis on the basis of a literature descrip- tion of a single male specimen from Brazil. Table 1 com- pares potential distinguishing features of all three nomi- nal Pontohedyle species, including the results of the pres- ent study on the female Brazilian specimen and the spec- imens of P milaschewitchii from the Mediterranean used for comparison. P. milaschewitchii P. verrucosa P. milaschewitchii Ganitus evelinae 4.1. External morphology and spicules Externally, the investigated specimen from Brazil confirms with the general acochlidian characters (e.g. visceral hump in which the head-foot complex can be at least partially retracted; see WAWRA 1987) and those of the genus Pon- tohedyle (lack of rhinophores). Using external characters, RANKIN (1979) differentiated P. brasilensis from P. mi- laschewitchii by referring to the flat triangular versus bow- shaped oral tentacles, and the absence or presence of cil- ia on head and oral tentacles. However, the shape of the oral tentacles ıs variable within specimens of Mediter- ranean P milaschewitchii (see JÖRGER et al. in press). They vary from bow-shaped to elongated triangular, including the flat and triangular form described by Marcus & MAR- cus (1954) for the Brazilian specimen (see Fig. 3B). Al- ready MARCUS & MARCUS (1954) illustrated that the ten- tacles can have a more rounded tip (see fig. 13, 14). This character clearly varies for one individual, depending on the contraction of the animal (see Fig. 3D: P verrucosa with supposedly slightly retracted tentacles). The variabil- ity of this character between individuals is underlined by the observation of our northern Brazilian specimen that had bow-shaped oral tentacles in living condition (see Fig. 3A). P. milaschewitchii Hedylopsis spiculifera Fig. 3. Different types of flattened oral tentacles in acochlidians. (A) Pontohedyle from northern Brazil (present study), as P. mi- laschewitchii. (B) P. milaschewitchii from southern Brazil after MARCUS & MARCUS (1954; fig. 10). (C) P. milaschewitchii from the Mediterranean after KOWALEVSKY (1901; fig. 46). (D) P. verrucosa after CHALLIS (1970; fig. SA). (E) Ganitus evelinae after Marcus & Marcus (1954; fig. 19). (F) Hedylopsis spiculifera (juvenile) after KOWALEVSKY (1901; fig. 49). 288 Katharina M. JÖRGER et al.: Re-description of Pontohedyle brasilensis Table 1 . Comparison of characters previously used for species delineation in the genus Pontohedyle. ? = no data available; * MAR- cus & Marcus (1954) originally described the radula as 44 (2.1.2) misinterpreting the central denticle of the lateral plate as in- complete cleavage (CHALLIS 1970; JÓRGER et al. in press). Data source Collecting site Collecting habitat Foot Oral tentacles Spicules Eyes Radula Digestive system Male genital system Female genital system P. milaschewitchii (Kowalevsky, 1901) KOWALEVSKY (1901) WAWRA (1986) JÖRGER et al. (in press) Sebastopol (Black Sea) Princess Islands, Lesbos Island, Istria (Mediterranean Sea) coarse and fine sands, sublittoral (2-9m) very short, posterior tip rounded bow-shaped to triangular/elongated - aggregatıon of needle-shaped parallel orientated spicules between the tentacles - randomly distributed needle-shaped spicules throughout the body present - 41-54 (1.1.1) - rhachidian tooth with I central cusp and 3 lateral denticles - lateral plate with 1 central denticle - no stomach detectable - salivary glands form one mass on the left side of the body, discharging into the oesophagus close to the transition of the pharynx ciliated vas deferens, extending to the level anterior to the oral tentacles, genital opening dorsal to the mouth opening ciliary band extending from the P. brasilensis (Rankin, 1979) Marcus & MARCUS (1954) present study Sao Paulo, Porto de Galinhas, Brazil (Atlantic Ocean) coarse and shell sands, intertidal and 20m depth very short, posterior tip rounded bow-shaped to triangular/elongated - aggregation of needle-shaped parallel orientated spicules between the tentacles - randomly distributed needle-shaped spicules throughout the body present - 44 (1.1.1)* - rhachidian tooth with l central cusp and 3 lateral denticles - lateral plate with 1 central denticle - “large, spherical stomach” according to MARCUS & MARCUS (1954), but no stomach detectable in the present study - salivary glands form one mass on the left side of the body genital opening on the posterior end of the head-foot complex short ciliary band extending genital pore to about one third of from the genital pore (present study) the head-food complex P. verrucosa (Challis, 1970) CHALLIS (1970) Maraunibina Island, Solomon Islands (Pacific Ocean) coarse, shell sand, intertidal very short, posterior tip free and pointed bow-shaped absent (7?) absent - 43 (11.1) - rhachidian tooth with l central cusp and 3 lateral denticles - lateral plate without denticle - no stomach described - salivary glands paired, discharging into the oesophagus “near its posterior end” Bonner zoologische Beiträge 55 (2006) 289 RANKIN (1979) claimed cilia to be absent from the head and oral tentacles of P milaschewitchii, in contrast to P. brasilensis. However, a constant pattern of cilia could be detected on the oral tentacles of Mediterranean P. mi- laschewitchii (see JÓRGER et al. in press). Similar cilia were described for the Brazilian specimen by MARCUS & MAR- cus (1954) and were also observed for the northern Brazil- ian specimen herein. Therefore, these external characters cannot be further used for separating species. In contrast, the Brazilian specimen described by MARCUS & MARCUS (1954) and the one studied herein resemble specimens of P. milaschewitchii (see JÖRGER et al. in press; KOWALEVSKY 1901) in all examined details, e.g. 1) body size and coloration, 2) shape of oral tentacles, 3) foot (short, posterior end rounded), 4) type (monoaxone) and position (accumulation between oral tentacles and ran- domly distributed all over the body) of spicules, and 4) presence of cilia on head and oral tentacles. 4.2. Microanatomy Anatomically, RANKIN (1979) saw differences between P. milaschewitchii and P. brasilensis regarding fused versus separated cerebral and pleural ganglia, the radula formu- la, the presence/absence of a well developed stomach, and the development of the salivary glands. 4.2.1. Central nervous system Probably based on small semi-schematic drawings of an entire specimen of P. milaschewitchii by KOWALEVSKY (1901; fig. 46, 48), RANKIN (1979) claimed the cerebral ganglia to be fused with the pleural ganglia in P milasche- witchii, while they were described to be separated in P brasilensis. We could not clearly detect pleural ganglia in our damaged northern Brazilian specimen. However, JORGER et al. (in press) showed that the cerebral and pleu- ral ganglia in Mediterranean P. milaschewitchii specimens are clearly separated, as usual for Acochlidia (HUBER 1993; SOMMERFELDT & SCHRODL 2005; WAwRA 1987). 4.2.2. Digestive system Marcus & MARCUS (1954) described a radula formula of 44 x 2.1.2 for their Brazilian specimen. However, CHAL- LIS (1970) suggested that the denticle in the lateral tooth might have been misinterpreted as a gap that appears to separate one broad lateral tooth into two. This explana- tion was accepted by WAwRA (1987) and is indeed very convincing. The radula of Mediterranean P. milasche- witchii closely resembles the one described by MARCUS & Marcus (1954) for the Brazilian specimen: there is a triangular rhachidian tooth with one central cusp that is bordered by three lateral denticles, and just one broad lateral plate on each side with one central denticle, thus with the formula 1.1.1 (JORGER et al. in press). Marcus & MARCUS (1954) saw an unusual large, spher- ical stomach in their Brazilian specimen, which was re- flected in Rankin’s generic name Gastrohedyle. No spe- cial stomach was detected in our Brazilian specimen, but an oesophagus passing into a moderately developed di- gestive gland cavity which was filled with particles. This reflects the normal condition found in Mediterranean P. milaschewitchii (see JORGER et al. in press), and all oth- er marine acochlidians. The large “stomach” described by Marcus & Marcus (1954) maybe easily explained as re- ferring to a digestive gland cavity filled with particles or artificially swollen by gases due to decomposition. RANKIN (1979) declared the salivary glands of P. milasche- witchii as “paired, well separated, long, thin, and taper- ing” in contrast to the large spherical salivary glands of P. brasilensis forming one mass on the left side of the body. However, the salivary glands in Mediterranean P. milaschewitchii are just like those described by MARCUS & Marcus (1954) for the Brazilian specimen and also those observed herein (JORGER et al. in press). 4.2.3. Genital system Marcus & Marcus (1954) described their male Brazil- ian specimen as having a genital opening located on the right side of the head-foot complex close to the transition to the visceral hump. This is the usual position for the fe- male genital pore P. milaschewitchii and of other male and female genital pores in microhedylid acochlidians. How- ever, Mediterranean male P. milaschewitchii show a male genital pore in an unusual cephalic position dorsal to the mouth opening (JORGER et al. in press; WAWRA 1986). Marcus & MArcus (1954) used the anterior part of the head-foot complex of their specimen for radula prepara- tion and were therefore unable to detect a male genital opening in an anterior position. The putative posterior opening in the male Brazilian specimen maybe thus ex- plained by generalization and misinterpretation or maybe due to different ontogenetic stages. If additional male Brazilian specimens in different ontogenetic stages did not show any ciliated duct leading anterior to a cephalic male genital opening but a posterior genital opening, this would be the first serious indication for a specific separation of P. brasilensis from P. milaschewitchii. The female genital system of our Brazilian specimen closely resembles the one observed for P. milaschewitchii (JORGER et al. in press; WAWRA 1986) in 1) presence of a ciliary band originating from the genital opening; 2) po- sition of the genital opening; 3) development and histo- 290 Katharına M. JÓRGER et al.: Re-description of Pontohedyle brasilensis logy of the nidamental glands; 4) comparably small size of mature oocytes (around 60 um). No differentiating fea- tures between the Brazilian specimen and its Mediter- ranean counterparts could be detected concerning the fe- male genital system. 4.3. Taxonomy All the differences between P milaschewitchii and P. brasilensis claimed by RANKIN (1979) are non existent (cilia pattern, radula formula, shape of salivary glands, fu- sion of cerebral and pleural ganglia), variable (shape of oral tentacles) or can be easily explained by biological fac- tors and artefacts (presence of large ““stomach”). Morpho- logical knowledge available at present (Table 1) strong- ly supports WAWRA (1987) in considering P. brasilensis as a junior synonym of P. milaschewitchii. However, the considerable geographical distance between the Mediter- ranean and the northern and southern Brazilian popula- tions of an interstitial species and the hydrographic dif- ferences between warm temperate and tropical waters re- quire molecular investigation as soon as abundant Brazil- ian populations can be found. Pontohedvle milaschewitchii as defined above is a Mediterranean and Atlantic species, while P verrucosa was described from the Solomon Islands in the tropical Indopacific (CHALLIS 1970). Main differences to P. mi- laschewitchii are the absence of spicules, eyes and later- al radula denticles (Table 1). However, at least the lack of spicules might be due to a preservation artefact; P. ver- rucosa urgently needs redescription and comparison with some other potentially undescribed Pontohedvle species found in the tropical Indopacific (see SCHRÓDL et al. 2003). Acknowledgements. We wish to thank Eva Lodde (ZSM, Mu- nich) for assistance in preparing the histological sections. Luis Simone and Carlo Magenta (Sáo Paulo) are domg great recov- ery work with the remainders of the Marcus” collection. AICA- Diving kindly provided SCUBA facilities in northern Brazil. Rosana Carvalho Schródl (ZSM) greatly helped with analysing innumerable sand samples. We also thank Heike Wágele (Bonn) and an anonymous referee for valuable suggestions and helpful comments on the manuscript. This study was supported by a DFG grant (SCHR 667-3) to MS. REFERENCES ARNAUD, P. M., POIZAT, C. & SALVINI-PLAWEN, L. v. 1986. Ma- rine-interstitial Gastropoda (including one freshwater inter- stitial species). Pp. 153-161 in: BOTOSANEANU, L. (ed) Sty- gofauna Mundi. Brill/Backhuys, Leiden. CHALLIS, D. A. 1970. Hedvlopsis cornuta and Microhedyle ver- rucosa, two new Acochlidiacea (Mollusca: Opisthobranchia) from the Solomon Islands Protectorate. Transactions of the Royal Society New Zealand (Biol. Sci.) 12: 29-40. HADL, G., KOTHBAUER, H., PETER, R. & Wawra, E. 1969. Sub- stratwahlversuche mit Microhedyle milaschewitchii Kowalevsky (Gastropoda, Opisthobranchia: Acochlidiacea). Oecologia 4: 74-82. Henry, E. C. 1977. A method for obtaining ribbons of serial sec- tions of plastic embedded specimens. Stain Technology 52: 59-60. HUBER, G. 1993. On the cerebral nervous system of marine Het- erobranchia (Gastropoda). Journal of Molluscan Studies 59: 381-420. JÖRGER, K. M., NEUSSER, T. P., HASZPRUNAR, G. & SCHRÖDL, M. In Press. Undersized and underestimated: 3D-visualization of the Mediterranean interstitial acochlidian gastropod Pontohedyle milaschewitchii (Kowalevsky, 1901). Organ- isms, Diversity & Evolution. KOWALEVSKY, A. 1901. Les Hédylidés, étude anatomique. Me- moires de l’Academie Imperiale des Sciences de St. Peters- bourg 12: 1-32. Marcus, E. & Marcus, E. 1954. Uber Philinoglossacea und Acochlidiacea. Kieler Meeresforschungen 10: 215-223. NEUSSER, T. P., HEß, M., HASZPRUNAR, G. & SCHRÖDL, M. 2006. Computer-based three-dimensional reconstruction of the anatomy of Microhedyle remanei (Marcus, 1953), an inter- stitial acochlidian gastropod from Bermuda. Journal of Mor- phology 267: 231-247. NEUSSER, T. P. & SCHRÓDL, M. 2007. Tantulum elegans reloaded: a computer-based 3D-visualization of the anatomy of a Caribbean freshwater acochlidian gastropod. Invertebrate Bi- ology 126: 18-39. PoizaT, C. 1984. Seasonal variations of Mediterranean intersti- tial opisthobranch assemblages. Hydrobiologia 118: 83-94. RANKIN, J. J. 1979. A freshwater shell-less Molluse from the Caribbean: structure, biotics and contribution to a new un- derstanding of the Acochlidioidea. Royal Ontario Museum Life Sciences Contributions 116: 1-123. RICHARDSON, K. C., JARETT, L. & FINKE, E. H. 1960. Embed- ding in epoxy resins for ultrathin sectioning in electron mi- croscopy. Stain Technology 35: 313-323. SCHRÖDL, M. 2006. Techniques for collecting interstitial opistho- branchs. http://www.seaslugforum.net/factsheet.cfm?base =inteextr, Sea Slug Forum. Australian Museum, Sydney. SCHRODL, M., EHEBERG, D. & BURGHARDT, I. 2003. Sulawesi: Solargetriebene Meeresnacktschnecken und wurmgleiche Sand-Hinterkiemer. Biologie in unserer Zeit 33: 18-19. SOMMERFELDT, N. & SCHRODL, M. 2005. Microanatomy of Hedy- lopsis ballantinei, a new interstitial acochlidian gastropod from the Red Sea, and its significance for phylogeny. Jour- nal of Molluscan Studies 71: 153-165. Spurr, A. R. 1969. A low-viscosity epoxy resin embedding medi- um for electron microscopy. Journal of Ultrastructural Re- search 26: 31-43. Wawra, E. 1986. Geschlechtsdimorphismus bei Pontohedyle mi- laschewitchii (Kowalevsky, 1901) (Gastropoda: Opistho- branchia). Annalen des Naturhistorischen Museums in Wien 87: 325-329. Wawra, E. 1987. Zur Anatomie einiger Acochlidia (Gastropo- da, Opisthobranchia) mit einer vorláufigen Revision des Sys- tems und einem Anhang úber Platyhedylidae (Opistho- branchia, Ascoglossa). Dissertation. Universitát Wien. Authors” addresses: Katharina M. JÖRGER (corresponding author), Zoologische Staatssammlung München, Münch- hausenstr. 21, D-81247 München, Germany, E-mail: k_jo- erger(vhotmail.com; Timea P. NEUSSER, E-mail: timea- neusser@gmx.de; Michael SCHRÓDL, E-mail: schroedl(@zi.biologie.uni-muenchen.de. Bonner zoologische Beitráge Band 55 (2006) Heft 3/4 Seiten 291-300 Bonn, November 2007 Pleurobranchaea inconspicua Bergh, 1897 (Opisthobranchia: Pleurobranchidae): E Redescription and distribution from Argentina and Colombia” Claudia MUNIAIND, Néstor E. ARDILa2) & Juan Lucas CERVERA?) Museo Argentino de Ciencias Naturales “Bernardino Rivadavia”, Buenos Aires, Argentina 2)Universidad de los Andes, Laboratorio de Biologia Molecular Marina, Laboratorio de Botánica y Siste- mática, Bogotá, Colombia 3)Universidad de Cádiz, Departamento de Biología, Cádiz, Spain *Paper presented to the 2nd International Workshop on Opisthobranchia, ZFMK, Bonn, Germany, September 20th to 22nd, 2006 Abstract. A taxonomic study based on fresh specimens from Argentina and Colombia, and voucher material from the collections of the Museo Argentino de Ciencias Naturales “Bernardino Rivadavia” (MACN) and Museo de Historia Na- tural Marina de Colombia (MHNMC-INVEMAR) was conducted. The first study by scanning electron microscopy (SEM) of the radular, jaw elements and reproductive anatomy, paying special attention to the internal cuticular stylet of the pe- nis, is presented. Previous descriptions of P. inconspicua and its junior synonyms, P. hamva, P. hedgpethi, and P. gela, are compared with the present results; P bonnieae is synonymized and a checklist of the distinguishing features of rele- vant species within the genus Pleurobranchaea is provided. This work aims at discussing the proposed amphiatlantic distribution of P inconspicua and at solving the problems of synonymies. It provides a starting point for a comparative study of specimens of the genus Pleurobranchaea from both sides of the Atlantic Ocean and the Mediterranean Sea that will be complemented with molecular studies in future times. Keywords. Taxonomic redescription, Colombian Caribbean, Argentina, amphiatlantic distribution. 1. INTRODUCTION The genus Pleurobranchaea Meckel in Leue, 1813 is rep- resented in temperate-warm waters, by several species with wide geographical distribution ranges. Pleurobran- chaea inconspicua Bergh, 1897 was described from north- ern Brazil (type locality: Sergipe, 10° 54°S, 37° 7 W), and its distribution was later extended under the same name or under junior synonyms of species and subspecies to sev- eral localities in the Western Atlantic Ocean (USA, Caribbean, Brazil, Argentina) and to localities more dis- tant from the type locality, such as the Mediterranean Sea (Israel) and West Africa (Ivory Coast to Nigeria). MARCUS & GOSLINER (1984) gave the most recent revi- sion of the subfamily Pleurobranchaeinae, which includ- ed 14 species of the genus Pleurobranchaea. However, these authors did not include the species P. morosa Bergh, 1892, P morula Bergh, 1905, P. melanopus Bergh, 1907, P. algoensis Thiele, 1925, P. japonica Thiele, 1925, and P. dorsali Allan, 1933, because they were poorly de- scribed. Nevertheless, P. japonica was redescribed by TSUBOKAWA et al. (1992), confirming the validity of the species and P. morosa has been recently mentioned from Azorean archipelago by MALAQUÍAS (2001) and CERVERA et al. (2006), although both works refer to the original description only. In MARCUS & GOSLINER’s revision (1984), the names P hedgpethi Abbott, 1952, P. hamva Marcus and Marcus, 1955 (mispelled hamwa) and P. gela Marcus and Marcus, 1966 were considered junior synonyms of P inconspicua. ABBOTT (1952) described P. hedgpethi from Port Aransas (Gulf of Mexico); this species became the most frequent- ly mentioned junior synonym of P. inconspicua for the Western Atlantic, even more frequently than the senior synonym. However, in the same revision, these authors erected the species P. bonnieae for one specimen from Florida region. According to VALDES et al. (2006) this name could be another junior synonym of P inconspicua. Several authors considered the anatomy of the reproduc- tive system and the penial morphology as the most use- ful characters to discriminate species of the genus Pleu- robranchaea compared to the classical hard structures of radulae and jaws (MARCUS 1961; MARCUS & GOSLINER 1984; WILLAN 1987). 292 Claudia MUNIAIN et al.: Pleurobranchaea inconspicua from Argentina and Colombia Fig. 1. Pleurobranchaea inconspicua Bergh 1897. A. Dorsal view of living specimen from Colombia (Photo: N. Ardila). B. Dor- sal view of head. C. Detail of the gill and the pairs of pinnules. D. Foot showing a conspicuous spur. E. Living specimen from Pa- tagonia (80 mm length) (Photo: C. Muniain). F. Detail of rhinophores and sensory papillae on the oral veil (Photo. C. Muniain). Bonner zoologische Beiträge 55 (2006) 293 Unlike most of the previous descriptions of P. inconspicua and of other species of the genus conducted from pre- served material, often in bad conditions, this work re- describes the characteristics of recently collected speci- mens of P inconspicua from two distant localities. This is the first study using SEM for hard structures of P. inconspicua, especially the examination of the penial stylet. Detailed descriptions of the external appearance and the hard structures, as well as the reproductive system, of specimens from distant localities help to understand the intraspecific variability in this species, and allow to dis- cuss the possible amphiatlantic distribution of P. incon- spicua by comparison with material from other Atlantic localities. Here, synonymization of other Pleurobranchaea species with P. inconspicua is outlined. 2. MATERIALS AND METHODS Specimens were collected over a wide range of depths from shallow intertidal areas down to 150 m in depth. A total of 16 specimens were preserved in 70% ethanol. Pho- tographs and videos were obtained with a digital camera from living specimens. Part of this material is deposited in the Museo de Historia Natural Marina de Colombia (MHNMC-INVEMAR) as well as in the Museo Argenti- no de Ciencias Naturales “Bernardino Rivadavia” (MACN). Colombia: Four specimens collected with a bottom trawl (9 x 1 m opening, 16 m in length) during the research cruises INVEMAR-MACROFAUNA II, 2001 aboard the R/V Ancon, in the Caribbean coast of Colombia at 70 and 150 m depth; two specimens off Dibulla (11° 25” 34N, 73° 27° 40” W), E-100, 150 m depth (MHNMC-INVE- MAR: 3872, 3873), two specimens off Buritaca (11° 18° 28” N, 73° 46’ 50” W) E-108, 109, 70 m depth (MHNMC- INVEMAR: 3874, 3875). One specimen was collected by SCUBA from Neguanje Bay, 5-10 m depth in a culture of bivalves (Ardila priv. coll.). Argentina: One specimen from Puerto Quequén (38° 34°S, 58° 38°W) Buenos Aires, 12/1928, MACN-in: 18234; one specimen from Puerto Quequén, Buenos Aires, 2/1929, MACN-in: 18312; two specimens from Puerto Quequén, Buenos Aires, 1/1934, MACN-in: 412; six spec- imens from Creek Bay (41° 6'S, 63° 56°W), San Matías Gulf, Rio Negro, MACN-in: 36985; one specimen from Larralde (42° 25°S, 64° 19°O), San José Gulf, Chubut, 10/1/2006, MACN-in: 37064. Two specimens were dissected, and their jaws and radu- lae were removed and mounted for Scanning Electron Mi- croscopy (SEM) examination (MACN-in: 36985, 37064). Two penises and their respective cuticular stylets were pre- viously critical-point-dried for SEM (MACN-in: 36985, 37064). 3. RESULTS 3.1. Systematic description Nudipleura Wágele & Willan, 2000 Pleurobranchoidea Gray, 1827 Pleurobranchidae Gray, 1827 Pleurobranchaeinae Pilsbry, 1896 Pleurobranchaea Meckel ın Leue, 1813 (type Pleuro- branchidium meckelii Blainville, 1826) Pleurobranchaea inconspicua Bergh, 1897 (Figs 1-5) Pleurobranchaea inconspicua Bergh, 1897 (BERGH 1897: 49-51, pl. 8, figs 2-10. MARCUS & GOSLINER 1984: 24-27, figs 1B, 10-11. Ríos 1984: 206-207, pl. 69, fig. 987. Mu- NIAIN 1997: 4, 17, 34. ARDILA & RACHELLO 2004: 62, fig.3). Pleurobranchaea hedgpethi Abbott, 1952 (ABBOTT 1952: 1-2, pl. 1, figs 1-8. MARCUS & Marcus 1959: 253 fig. 6. Marcus & Marcus 1960: 253-254, fig. 6. NIJSSEN- MEYER 1965: 143-145, figs 1-2. MARCUS & MARCUS 1967a: fig. 55C. MARCUS & Marcus 1967b: 200. Mar- cus & Marcus 1969: 18. ABotT 1974: 348, fig. 4201. Pleurobranchaea hamva Marcus & Marcus, 1957 (MAR- cus & Marcus 1957: 21-27, fıgs 40-52). Pleurobranchaea hedepethi hamva Marcus, 1961 (MARCUS ER. 1961: 141. MARCUS & MARCUS 1967a: 48). Pleurobranchaea gela Marcus & Marcus, 1966 (MARCUS & MARCUS 1966: 174, figs 35-37. Pleurobranchaea bonnieae Marcus & Gosliner, 1984 (MARCUS & GOSLINER 1984: 29-32, figs 1D, 13-14.). 3.1.1. External features Length of the living animals ranges between 13 and 80 mm.The body ıs oval and elongate. The ground colour is pale cream, with a dense dark brown reticulate along the mantle dorsal region, being lighter and less obvious in the foot dorsal region and absent in the well developed rhinophores (Fig. 1A, B, E). Bright white spots are scat- tered all over the mantle and more conspicuous on the dor- sal foot surface. The oral veil is broad with sensory papil- lae along the anterior edge. Rinophores are held erect in the living animals (Fig. 1B, F), whereas in the preserved ones they are in a lateral position. 294 Claudia MUNIAIN et al.: Pleurobranchaea inconspicua from Argentina and Colombia The mantle is reduced and fused with the oral veil; a large gill is visible on the right side of the body. The gill is at- tached along most of the body with about 26 pairs of pin- nules per gill (Fig. 1 C, E). The anal papilla lies over the seventh pinnule. The foot is rounded in front. The soft mantle, not covering the foot, 1s visible to most of its part (Fig. 1 A, E). The coloration in the ventral foot region is translucent, and the internal organs can be observed by transparency. At the posterior end, the foot ends in a round- ed shape, with a large metapodial gland (yellowish) ex- tending longitudinally from the posterior mantle towards the central foot sole (Fig. 1 E). Dorsally to the metapodi- al gland, the foot has a brown or black tiny spur (Fig. 1A, D). 3.1.2. Anatomy A shell is lacking. The pharyngeal bulb frequently protrud- ed in preserved specimens through the eversion of the very extensible oral tube. Jaws consist of two paired and elon- gate chitinous plates on the lateral walls of the pharyn- geal bulb. The radular formula of one specimen of 40 mm in life is 30 X 64.0.64 (MACN-in: 36985). The radula lacks a rachidian tooth, but a difference among the first four laterals is noticeable in some radular rows (Fig. 2A). Lateral teeth are bicuspid, largest in the middle of the half- row and smaller towards the outer border (Fig. 2B). The jaw elements (rodlets) bear 4 to 8 denticles from the out- ermost and innermost region (Figs 2C, D). The reproductive system arrangement is similar to that of other Pleurobranchaea species (Fig. 3). The tubular am- pulla forks near its outlet into two branches, the shorter branch is the oviduct that enters the capsule gland, and the other branch is the efferent duct that splits in two. One branch enters the prostate and then continues within the penial sac (Fig. 3, number 4). The second branch leads to the bilobed seminal receptacle, which is connected to the bursa copulatrix by a long and thin convoluted duct. The bursa copulatrix is large and round. Fig. 2. tral tooth. B. Outermost teeth on the same radula side (right). C, D. Jaw showing platelets with 4-8 denticles from outermost re- gion. D, Jaw showmg platelets from innermost region. A-D. Scanning electron photographs of radula of P. inconspicua MACN-in: 36985. A. Radula lacking a rachidian cen- Bonner zoologische Beiträge 55 (2006) 295 Fig. 3. Reproductive system of P inconspicua MACN-in: 36985. Scale bar: Imm. Abbreviations: AM ampulla, BC bursa copu- latrix, CG capsule gland, ED efferent duct, LPST penis loop containing the cuticular stylet, OV oviduct, PP penial papilla, PR pro- state, PS penial sac, PST penis (cuticular stylet), SR seminal receptacle, VA vagina. For numbers see text. The penis is an elastic cutilar stylet, that coils in numer- ous helically loops (Fig. 3, number 3), that finally enters in the atrium through a penial papilla (Fig. 3, numbers 2, 1). It is difficult to find the point where the efferent duct and the penis are connected. When stretched out, the pe- nis reaches a length of 16 and 20 cm in specimens of 45 and 80 mm length respectively. Observations of the pe- nis under SEM reveal an external cuticule and internal cu- ticular stylet throughout its length (Fig. 4 A-E). A transversal section of the penis shows that the chitinous stylet is not rounded and has a central cylindrical orifice. The shape and size of these penial stylets from two spec- imens examined under SEM are similar (Figs A, B, C). The width of those stylets is about 80 um. The penis has more than 15 roundish loops (Fig.3, LPST, number 3), and contains the cuticular stylet all along its length (Fig. 4D). The penial stylet runs only along one fifth of the penial cuticule (Fig. 4E). Apparently, as observed from the de- tail of the final portion of the efferent duct, in its portion connecting with the prostate, a central cuticular stylet ın- side it would be lacking (Fig. 4F). 3.2. Geographic distribution The geographical distribution cited for P. inconspicua is from Cape Hatteras, North Carolina 35° N 80° W (USA) to San José Gulf 42° S 64° W (Argentina) (See Table 1, Fig. 5). Regarding the Caribbean distribution of this species, the first record from Colombia was given by ARDI- LA & RACHELLO (2004), with specimens from 70-150 m and an individual collected from 8-10 m, present study. MARCUS & MARCUS (1969) recorded specimens for the first time (as P. hedgpethi) for Argentina (40° 32° S, 60° 19° W, 19 specimens at 57 m) and (40° 11° S, 60° 27° W, 296 Claudia MUNIAIN et al.: Pleurobranchaea inconspicua from Argentina and Colombia Fig. 4. SEM (critical-point-dried) of reproductive system of P. inconspicua (MACN-in: 36985; 37064). A. Penis, showing the ex- ternal cuticule and internal chitinous stylet (MACN-in: 36985); B and C. Detail of the tranverse section of the cuticular stylet of two different specimens (B, MACN-in: 36985; C, MACN-in: 37064). D. Portion of penial loop with a length of 20 cm from spe- cimen of 80 mm in length (MACN-in: 37064). E. Dorsal view of the external cuticule, showing a portion of the cuticular stylet in- side it (MACN-in: 36985). F. Detail of the region of the efferent duct that leaves the prostata (MACN-in: 37064). 7 specimens at 44 m), and their geographical range has been extended to northern Patagonia some years later for the intertidal to 8 m (MUNIAIN, 1997, present study). MARCUS & GOSLINER (1984) cited localities from the Mediterranean Sea (Israel) and West Africa (Ivory Coast to Nigeria). Further investigations will allow testing the hypothesis of an amphiatlantic distribution. 4. DISCUSSION As mentioned above, this work intends to serve as a start- ing point for further taxonomic and molecular studies in- volving a larger number of fresh material to verify whether this species is also present along the western coasts of Africa and the eastern Mediterranean (Israel) (amphiat- lantic species), as MARCUS & GOSLINER (1984) stated, or Bonner zoologische Beiträge 55 (2006) 297 o . Caribbean Sea 2 ¿o ATLANTIC OCEAN Fig. 5. Known distribution of Pleurobranchaea inconspicua Bergh, 1897, including its synonyms. to confirm the validity of other species of the genus (e.g. P. tarda from Western and South Eastern Atlantic, Table 1, Fig. 5). Thus, VALDÉS et al. (2006) suggest that P bonnieae Mar- cus and Gosliner, 1984 could be a junior synonym of P. inconspicua. We agree with this opinion, since the descrip- tion of this species is based on a single preserved and, very probably, juvenile specimen and the features suggested to differentiate it from other nominal species (P. inconspicua or P. vayssierei) are weak (presence of a penial cuticle or diameter of the base of the penis as they can be artifacts of preservation). There is no description of the living an- imal. Besides all the facts stated above, MARCUS & GOSLINER (1984) provide unclear information about the collecting localities of their material of P. inconspicua, but they give a geographical range in the western Atlantic that would include the type locality of P bonnieae. Another species of Pleurobranchaea, P. agassizi Bergh, 1897, would fit the geographical range for P. inconspicua. How- ever, the colour of living specimens of P. agassizi, its ba- thymetric range and some anatomical differences concern- ing the radular teeth and the reproductive system make us consider this species as valid until additional fresh mate- rial can be studied. 298 Claudia MUNIAIN et al.: Pleurobranchaea inconspicua from Argentina and Colombia External and internal characteristics of P inconspicua have been described from fresh collected specimens from two distant localities of 1ts wide distribution, as well as from preserved material. The number of pinnae of the gill and the presence of a conspicuous caudal spur are shared by all dissected specimens. All Patagonian living specimens showed the caudal spur; but it ıs possible that after preser- vatıon the spur ıs not evident, and for this reason in some descriptions 1t might have not been mentioned. The same hypothesis can be drawn for the posterior region of the foot sole, in which a yellow metapodial gland forms a lon- gitudinal furrow, often visible by transparency through the skin of the dorsal side of the tail. The internal anatomy, the radula, the jaw elements and the arrangement of the reproductive organs fully agree with previous descriptions. The morphology of the internal cuticular cylindrical stylet is similar to that illustrated by MARCUS & GOSLINER (1984, fig. 1B) by optical transverse section. An updated review of the species of Pleurobranchaea with details of the radular and jaw elements using SEM is need- ed. Although at present these characters are not consid- ered for specific identifications, they can provide further information that can be used in the comparison of species of dubious identity. Regarding the reproductive system, the presence of a penis with cuticular stylet is an impor- tant character to separate species, but its occurrence in many species is not clear, as in P. californica, MacFarland, 1966 and P. meckelii Meckel in Leue, 1813. Some authors indicate its presence while others do not. CERVERA & GAR- CIA-GÖMEZ (1988) suggest that P notmec Marcus and Gosliner, 1984 and P. vayssierei Marcus and Gosliner, 1984 are junior synonyms of P. meckelii, which agrees with a personal communication by Richard Willan. The P. notmec’s holotype is from eastern Mediterranean (off Turkey) but the paratypes are from Israel (MARCUS & GOSLINER 1984), as well as one specimen attributed to P inconspicua by the same authors. On the other hand, P. vayssierei Marcus and Gosliner, 1984 was described from a single specimen from Algiers (Western Mediterranean) among the Vayssiere’s material attributed to P. meckelii. The diagnostic features used to erect P notmec and P. vayssiere seem weak to us since these species are de- scribed from few specimens and the characters vary in- traspecifically (e.g. ontogeny) or could be artifacts due to preservation. Even MARCUS & GOSLINER (1984) over- looked, or were not able, to place P. notmec within their key to the species of Pleurobranchidae. Overall, we think that a final decision on the identity of this species will be possible only after a review of the type material and af- ter additional fresh material is studied from the type lo- calities areas based on both methods, morphological and molecular analyses. A future revision of type material of P. notmec and P. vayssierei, of further specimens from the type localities, as well as of P inconspicua individuals from the Mediterranean Sea (Israel), will allow to eluci- date the existence of more than one species of this genus in the Mediterranean or confirm that P meckelii is the on- ly species present in this region. Acknowledgements. The authors are grateful to the staff of Museo de la Plata, Patricia Sarmiento and Museo Argentino de Ciencias Naturales “Bernardino Rivadavia”, Fabian Tricarico, for their valuable assistance during SEM sessions. And to Nés- tor Cazzaniga (Universidad del Sur, Bahia Blanca) for kindly providing specimens for this study from Rio Negro. Two anony- mous reviewers provided helpful comments on the manuscript. This study was funded by Argentinian grants from GEF-BIRF 28385/AR (A-CB-51) and ANPCyT (PICT 34111) and Spanish grant GCL2006-05182/BOS (DGI of the Education and Science Ministry). C.M. is Research Member of the National Research Council of Argentina (CONICET). REFERENCES ABBOTT, R. 1952. Two new opisthobranchs mollusks from the Gulf of Mexico belonging to the genera Pleurobranchaea and Polycera. Florida Station University Studies 7: 1-7. ABBOTT, R. 1974. American Sea Shells. 2"4 ed. New York. D. Van Nostrand Reinhold. 663 pp. ARDILA, N. & RACHELLO, P. 2004. Opisthobranchs (Mollusca: Gastropoda) collected by the cruises Invemar-Macrofauna II in the Colombian Caribbean (20-150 m). Avicennia 17: 57-66. BERGH, E. S. R. 1892. Opisthobranches provenant des cam- pagnes du Yacht I’ Hirondelle. Resultats des campagnes par son yacht par Albert I de Monaco 4: 1-35. BERGH, L.S.R. 1897. Malacologische Untersuchungen 5. Pp. 1-115 in: Semper, C. (ed.) Reisen im Archipel der Philippi- nen 7, 4 Abt., | Absch., Die Pleurobranchiden 1-2. Wiesba- den Kreidel’s Verlag. Burn, R. 1989. Opisthobranchs (Subclass Opisthobranchia). Pp. 725-788 in: SHEPHERD, S. A. & THOMAS, I. M. (eds.). Marine Invertebrates of Southern Australia. Part II. South Australian Govermment Printing Division, Adelaide. Burn, R. 2006. A checklist and bibliography of the Opistho- branchia (Mollusca: Gastropoda) of Victoria and the Bass Strait area, South eastern Australia. Museum Victoria Science Reports 10: 1-42. CERVERA, J. L. & GARCÍA GÓMEZ, J. C. 1988. Estudio anatömi- co de Pleurobranchaea meckelii Blainville, 1825 (Mollusca: Opisthobranchia: Notaspidea). Arquivos do Museu Bocage 1: 71-70. CERVERA, J. L., CALADO, G., GAVAIA, C., MALAQUÍAS, M.A. E., TEMPLADO, J. BALLESTEROS, M., GARCÍA GÓMEZ, J. C. & MEGINA, C. 2006 (for 2004). An annotated and updated check- list of the opisthobranchs (Mollusca: Gastropoda) from Spain and Portugal (including islands and archipelagos). Boletin del Instituto Espanol de Oceanografía 20(1-4): 5-111. Dayrat, B. 2000. Indo-Pacific deep water Pleurobranchaeidae (Gastropoda, Opisthobranchia, Notaspidea): New record and new species. Pp. 321-330 in: BOUCHET, P. £ MARSHALL, A. (eds.), Tropical Deep-Sea Benthos, volume 22, Mémoires du Museum nationale d'Histoire naturelle, 185. GOSLINER, T. M. 1985. Redescription and systematic position of Pleurobranchaea obesa (Verrill, 1882) (Opisthobranchia: Pleurobranchaeidae). The Veliger 28: 109-114. Bonner zoologische Beiträge 55 (2006) 299 JENSEN, K. R. 1997. Cannibalism in an opisthobranch mollusc, Pleurobranchaea brockii Bergh, 1897 from Hong Kong wa- ters. Phuket Marine Biological Center Special Publication 17(1): 41-46. MACFARLAND, F. 1966. Studies of the opisthobranchiate mol- lusks of the Pacific Coast of North America. Memoires of the California Academy of Science 6: I-XIV, 1-546. MALAQUIAS, M. A. E. 2001. Updated and annotated checklist of the opisthobranch molluscs (excluding Thecosomata and Gymnosomata), from the Azores archipelago (North Atlantic Ocean, Portugal). Iberus 19: 37-48. Marcus, Er. 1961. Opisthobranchs from North Carolina. Jour- nal of the Elisha Mitchell Scientific Society 77: 141-151. MARCUS, Ev. & MARCUS, ER. 1957. Sea-hares and side gilled slugs from Brazil. Boletim do Instituto Oceanográfico de Sao Paulo 6: 3-49. MARCUS, Ev. & MARCUS, Er. 1959. Some opisthobranchs from the north-western Gulf of Mexico. Publications of the Insti- tute of Marine Science of University Texas 6: 251-264. MARCUS, Ev. £ MARCUS, ER. 1960. Some opisthobranchs from the northwestern Gulf of Mexico. Publications of the Institute of Marine Science, University Texas 6: 251-264. MARCUS, Ev. & MARCUS, ER. 1966. Opisthobranchia from trop- ical West Africa. Studies in Tropical Oceanography 4: 152-208. MARCUS, Ev. & MARCUS, Er. 1967a. American opisthobranch molluscs. Studies in Tropical Oceanography 6: I-VII, 1-256. MARCUS, Ev. & MARCUS, ER. 1967b. Some opisthobranchs from Sapelo Island, Georgia, U.S.A. Malacologia 6: 199-222. MARCUS, Ev. & MARCUS, ER. 1969. Opisthobranchian and lamel- larian gastropods collected by the “Vema”. American Muse- um Novitates 2368: 1-33. MARCUS, Ev. & GOSLINER, T. 1984. Review of the family Pleu- robranchaeidae (Mollusca, Opisthobranchia). Annals of the South African Museum 93: 1-52. MUNIAIN, C. 1997. Moluscos Opistobranquios de Argentina: Re- visión Taxonómica y relación de Ecología Química en algu- nas especies patagónicas. PhD Thesis, Unpublished. Univer- sity of Oviedo, Spain. NIJSSEN-MEYER, J. 1965. Notes on a few opisthobranch Mollus- ca from Surinam (Guianas). Zoologische Mededelingen 40: 144-150. Ríos, E. 1984. SeaShells of Brazil. 2"4 Edition. Fundacao Uni- versidade do Río Grande. Brazil. 368 pp. Thompson, T. E. 1970. Eastern Australian Pleurobranchomor- pha (Gastropoda, Opisthobranchia). Journal of Zoology 160: 173-198. TSUBOKAWA, R., WILLAN, R. C. & OKUTANI, T. 1992. Taxono- my of the two species of Pleurobranchaea in Japan (Gastropo- da: Notaspidea: Pleurobranchidae). Venus, The Japanese Jour- nal of Malacology 50: 249-263. VALDÉS, A., HAMMAN, J., BEHRENS, D.W. & Dupont, A. 2006. Caribbean Sea Slugs. A field guide to the opisthobranch mol- lusks from the tropical northwestern Atlantic. Sea Challengers Natural History Books, Gig Harbour (WA), USA. VAYSSIERE, A.J.B.M. 1885. Recherches zoologiques et anatomiques sur les Mollusques Opisthobranches du Golfe de Marseille, I. Annales du Musee d Histoire Naturelle de Mar- seille (Zoologie) 2: 1-181. VAYSSIERE, A.J.B.M. 1901. Monographie de la famille des Pleu- robranchidés, I. Annales des Sciences Naturelles Zoologie (8) 12: 1-85. VAYSSIERE, A.J.B.M. 1902. Opisthobranches et Prosobranches. Pp. 221-270 In: Expéditions scientifiques du “Travailleur’ et “Talisman”: Pendant les années 1880-1883: Ouvrage publié sous les auspices du ministére de instruction publique. Pa- ris. WILLAN, R. 1983. New Zealand side-gilled sea slugs (Opistho- branchia: Notapidea: Pleurobranchidae). Malacologia 23: 221-270. WILLAN, R. 1987. Phylogenetic systematics of the Notaspidea (Opisthobranchia) with a reappraisal of families and genera. American Malacological Bulletin 5: 215—241. Author's address: Dr. Claudia MUNIAIN (corresponding author), Museo Argentino de Ciencias Naturales “Bernardino Rivadavia”, Avda. Angel Gallardo 470 (C1405DJR) Buenos Aires, Argentina, E-mail: cmuni- ain(@macn.gov.ar; Néstor E. ARDILA, Universidad de los Andes, Laboratorio de Biología Molecular Marina-BIO- MMAR, Lab. Botánica y Sistemática, Bogotá, Colombia, E-mail: ne.ardila23@uniandes.edu.co; Juan Lucas CERVERA, Universidad de Cádiz. Polígono del Río San Pe- dro s/n, Apdo. 40, 11510 Puerto Real (Cádiz), Spain, E- mail: lucas.cervera(Wuca.es. Claudia MUNIAIN et al.: Pleurobranchaea inconspicua from Argentina and Colombia 300 Table 1.Diagnostic features of species of the genus Pleurobranchaea Leue, 1813. Species Pleurobranchaea meckelii (Blainv P. maculata (Quoy & Gaimard, 1832) P. tarda Verril, 1880 P. obesa (Verril, 1882) P. morosa (Bergh, 1892) P. agassizii Bergh, 1897 P. brockii Bergh, 1897 P. inconspicua Bergh, 1897 P. japonica Thiele, 1925 P californica MacFarland, 1966 P. augusta Marcus & Gosliner, P. vayssierei Marcus & Gosliner, P bubala Marcus & Gosliner, P. notmec Marcus & Gosliner, P. catherinae Dayrat, 2000 1984 1984 1984 1984 Geographical Distribution Radular formula Type Locality: ? Mediterranean Other localities: Several Mediterranean localities (Naples, Sicily, Marseille, Genoa), Atlantic and Mediterranean Iberian Peninsula, Canary Islands, Azores Islands,Cape Verde Islands. 0-150 m. Type Locality: ? Southern Australia Other localities: Jarvis Bay (New South West, Australia), New Zealand, Japan ?, South Africa ?, Indo West Pacific, 0-300 m. 46 X 71-0-71 50-60 X 65-55.1.55-65 ?X 70.1.70 ? X 70.0.70 40-50 X 60-70.0.70-60 25 X 50-0-50 40-49 X 80-0-80 40-50 X 70-80.1.80-70 Type Locality: ? Martha’s Vineyard (Masschussets) ? X 70.0.70 Other localities: West Atlantic from Masschussets to south of Cuba. East Atlantic from Ghana & Cape of Good Hope, South Africa. 0-1450 m. Type Locality: ? 39°53°00° N, 69°50° 30 °W Other localities: East Coast of North America. 353-571 m. Type Locality: Strait of Pico-Fayal (Azores Islands). 130 m. Type Locality Straits of Florida, Great Bahama Bank (NW Atlantic), 200-620 m. Type Locality: ? Amboine (Indian Ocean) Other localities: Indo West Pacific, Japan, Hong Kong, South Africa, Australia. 0-60 m. Type Locality: Sergipe (Brazil) Other localities: North Carolina, Florida, Belize, Honduras, Venezuela, Virgin Islands, Martinique, St. Vicent & the Grenadines, Grenada, Surinam, Florida, Colombia, Argentina, Tropical West Africa (Ivory Coast to Nigeria), Israel. 0-150 m. Type Locality: ? Kóbe (Japan). Other localities: South Korea. 0-400 m. Type Locality: Golden Gate and southeast of the Farallon Light. Other localities: California. 10-470 m. Type Locality: West Africa (17%2'S, 11° 40’E). 54 m. Type Locality: ? Algiers (Western Mediterranean) Type Locality: Buffels Bay at Cape Point (South Africa). 10 m. Type Locality: ? Israel. Other localities: Eastern Mediterranean, off Turkey. 11-140 m. Type Locality: Coral Sea. Tropical Indo-Pacific Other localities: Indonesia, Philippines, Vanuato and the Marquesas. 352-760 m. 34 X 90-0-90 38 X 80.0.80 37 X 70-68.0.68-70 32 X 98.0.98 35 X 90.0.90 40-48 X 65.90.0.65.90 36 X 58.0.58 52.X%.52:0.52 35 X 65.0.65 32-40 X 55-68.0. 34 X 72.0.72 30 X 64.0.64 35 X 55.0.55 50-80.0.50-80 52 X 130-145.1.130-145 40 X 70.0.70 35 X 65.0.65 35 X 100.0.100 50 X 80.0.80 90 X 45.0.45 5 5-68 Body Length up to 100 up to 50 75-110 40-128 20 38-110 up to 60 12-100 up to 335 us ID 12-40 un m Cuticular stylet present No data present No data No data No data present No data present present No data present No data No data Metapodial References Spur present VAYSSIERE (1885, 1901, 1902) , BERGH (1897), MARCUS & GOSLINER (1984), CERVERA & GARCÍA-GÓMEZ (1988), CERVERA et al. (2006) present VAYSSIERE (1901), THOMPSON (1970), WILLAN (1983), MARCUS & GOSLINER, (1984), BURN (1989: 2006) present VAYSSIERE (1901), MARCUS & GOSLINER (1984) present BERGH (1897), VAYSSIERE (1901), MARCUS & GOSLINER (1984); GOSLINER (1985) No data BERGH (1892), MALAQUÍAS (2001), CERVERA et al (2006) present BERGH (1897), VAYSSIERE (1901), MARCUS & GOSLINER (1984) present BERGH (1897), VAYSSIERE (1901), MARCUS & GOSLINER (1984), JENSEN (1997) present BERGH (1897), VAYSSIERE (1901), ER. Marcus (1957, 1961), NIissen-MEYER, 1965), Er. MARCUS (1957), Ev. Marcus & ER. MARCUS (1966, 1967b), MARCUS & GOSLINER (1984), MUNIAIN (1997), ARDILA & RACHELLO (2004), VALDES et al. (2006), present study No data TSUBOKAWA et al. (1992) No data MACFARLAND (1966), MARCUS & GOSLINER (1984), WILLAN (1987) present MARCUS & GOSLINER (1984) present MARCUS & GOSLINER (1984) No data MARCUS & GOSLINER (1984) present MARCUS & GOSLINER (1984) present DAYRAT (2000) Bonner zoologische Beiträge Band 55 (2006) Heft 3/4 Seiten 301-310 Bonn, November 2007 Exploring Cerebral Features in Acochlidia (Gastropoda: Opisthobranchia)* Timea P. NEUSSER, Katharina JORGER & Michael SCHRODL!) DZoologische Staatssammlung München, Munich, Germany *Paper presented to the 2nd International Workshop on Opisthobranchia, ZFMK, Bonn, Germany, September 20th to 22nd, 2006 Abstract. Histological semithin sections of the marine acochlidian species Hedylopsis spiculifera (Kowalevsky, 1901), H. ballantinei Sommerfeldt & Schródl, 2005, Microhedyle remanei (Marcus, 1953) and Asperspina murmanica (Kudins- kaya & Minichev, 1978) and of the limnic Tantulum elegans Rankin, 1979 were (re)examined for different cerebral fea- tures: 1) the number of cerebro-rhinophoral connectives, 2) the presence of Hancock s organs, 3) the relative position and size of the eyes, the length and diameter of the optic nerve, and the presence of an optic ganglion, and 4) cellular ag- gregates attached to the cerebral ganglia. We describe novel structures such as double cerebro-rhinophoral connectives in T. elegans, and “lateral bodies” in A. spiculifera, H. ballantinei and A. murmanica. Cerebral features are discussed as a promising additional set of characters for phylogenetic analysis. However, (ultra)structural comparisons of acochlidians with basal opisthobranchs and pulmonates are overdue. Keywords. Cerebral nerves, “lateral bodies”, dorsal bodies, Hancock’s organ, optic ganglion. 1. INTRODUCTION Acochlidian opistobranch gastropods show high morpho- logical and biological diversity. However, the number of useful characters for phylogenetic analyses is still limit- ed by the paucity of comparative data available. The cen- tral nervous system (cns) of several euthyneurous taxa was described (e.g. HASZPRUNAR & HUBER 1990; HUBER 1993; MIKKELSEN 2002), comprising data about cerebral nerves and sensory organs. The value of these data in phyloge- netic studies is evident (DAYRAT 8 TILLIER 2002; MIKKELSEN 1996). In contrast, several of the species (re)descriptions in Acochlidia do not include any infor- mation on the cns (e.g. HAYNES & KENCHINGTON 1991; HuGHEs 1991; KIRSTEUER 1973; MARCUS & MARCUS 1955, 1959; SALVINI-PLAWEN 1973; WAWRA 1979, 1980, 1988). Other authors limited their descriptions of the ens to the main ganglia on the (pre)pharyngeal nerve ring and the visceral nerve cord (e.g. BERGH 1895; BUCKING 1933; CHALLIS 1968, 1970; Doe 1974; HERTLING 1930; KOWALEVSKY 1901; KUDINSKAYA & MINICHEV 1978; KUTHE 1935; Marcus 1953; MARcus & MARCUS 1954; Morse 1976; SWEDMARK 1968; WAWRA 1989; WESTHEI- DE & Wawra 1974). Unfortunately, the identification of the small and hardly separated ganglia on the visceral nerve cord is problematic. Even detailed histological de- scriptions, such as that of Tantulum elegans by RANKIN (1979), can be considerably misleading and thus cannot be trusted (see NEUSSER & SCHRODL 2007). Furthermore, very few studies give data about cerebral nerves and sen- sory organs reflecting the complexity of the acochlidian cns. HUBER (1993) gave a detailed overview of the cns in marine heterobranchs and determined the number of cere- bral nerves in Acochlidia to only two (the labiotentacular nerve and the proximally joint oral and rhinophoral nerve) plus the static nerve. SOMMERFELDT & SCHRODL (2005) confirmed these three nerves plus optic nerves for Hedy- lopsis spiculifera and H. ballantinei. The authors empha- sized the presence of large rhinophoral ganglia, from which the joint oral and rhinophoral nerve arise, and that was overlooked in H. spiculifera by HUBER (1993). The terminology and the homology of the different cerebral nerves in Acochlidia are still uncertain. Data about sensory organs are sparse, often consisting on- ly in the affirmation of presence or absence of easily iden- tified structures, such as eyes (e.g. CHALLIS 1970; MAR- cus 1953; Marcus & Marcus 1955; WESTHEIDE & WawRA 1974). Hancock’s organs, the primary chemosen- sory organs in architectibranchs and cephalaspideans (MIKKELSEN 1996, 2002), were thought to be absent in Acochlidia (e.g. NEUSSER et al. 2006; SOMMERFELDT & SCHRODL 2005; WAwRA 1987). However, Hancock’s or- gans like structures were reported from Microhedyle glan- 302 Table 1 . Comparison of cerebral features in different acochlidian species. +: present, — species feature Double cerebro- rhinophoral connective Hancock’s organ Eyes Eyes externally visible Eyes position Eye size in diameter Optic nerve Timea P. NEUSSER et al.: Cerebral features in Acochlidia Hedylopsis spiculifera 9 9 + pigmented dorsal and lateral well visible posterior to the rhinophores (in some distance) 25 um long, undulated Hedvlopsis ballantinei 9 9 + pigmented dorsal and lateral hardly visible slightly posterior to the rhinophores (at their base) 30 um long, undulated Asperspina murmanica absent, ?: not detected. Tantulum Microhedyle elegans remanei + 2 E ? + reduced unpigmented = not visible - slightly anterolateral - to the cerebral ganglion 20 um - short, not undulated = Optic nerve diameter 6-7 um 6-7 um - 3 um - Optic ganglion (diameter) — - + = (18 um) Lateral bodies + + + = x Cells above cerebral ? ? + ES 2 commissure dulifera (Kowalevsky, 1901) and Pontohedyle milasche- witchii (Kowalevsky, 1901) by EDLINGER (1980a, b), and recently confirmed for P milaschewitchii (JÖRGER et al. in press). Additionally, our re-examination of Tantulum el- egans revealed the presence of a small Hancock’s organ in this species too (NEUSSER & SCHRÖDL 2007). Among representatives of four traditional acochlidian fam- ilies (Hedylopsidae, Asperspinidae, Tantulidae and Micro- hedylidae), the present study (re)investigates a number of special cerebral nervous features using histological sec- tions. As far as information is available, these characters are compared with other acochlidian species and are eval- uated as a possible set of characters for future phyloge- netic analysis. 2. MATERIAL Serial semi-thin sections of five different acochlidian species were available for re-examination by light mi- croscopy: one series (section thickness: 1.5 um) of Hedy- lopsis spiculifera, Zoologische Staatssammlung Múnchen, ZSM N° 20070391 (Secche della Meloria, Livorno, Italy, September 2005) and one paratype series (section thick- ness: 2 um) of Hedylopsis suecica Odhner, 1937, Swedish Museum of Natural History, SMNH N° 27211; H. sueci- ca was considered as a synonym of A. spiculifera by WAWRA (1989) and confirmed by SOMMERFELDT & SCHRÖDL (2005). Five paratype series (section thickness: 2 um) of Hedvlopsis ballantinei, ZSM N° 20004766/1, 20004767, 20004768, 20004769 and N° 26X (Dahab, Gulf of Agaba, northern Red Sea, October 1999). Six series (section thickness: 1.5 um) of Microhedyle remanei, ZSM N° 20070079, 20070080, 20070081, 20070082, 20070083 and 20070084 (southwest of Castle Roads, Bermuda Is- lands, July 1999). Four series (section thickness: 1.5 um) of Asperspina murmanica, ZSM N° 20062163, 20062164, 20062165 and 20062167 (Yarnyshnaya Bay, Barents Sea, Russia, August 2005). Four original paratype series (sec- tion thickness: 3 um) and two recently prepared paratype series (section thickness: 1.5 um) of Tantulum elegans, Royal Ontario Museum, Canada, ROM N° SEI and 2F0 (Golden Grove, St. Vincent, West Indies, July 1972). All sections, except the original paratype series of 7. elegans, were stained with methylene blue-azure II according to RICHARDSON et al. (1960). Bonner zoologische Beiträge 55 (2006) 303 3. CEREBRAL FEATURES EXAMINED 3.1. Rhinophoral ganglia and cerebro-rhinophoral connectives A comparative overview of all examined features in the different species is given in Table 1. All species re-examined herein, except Microhedyle re- manei, have a pair of true rhinophoral ganglia, 1.e. large ganglia separated into a nuclei-free medulla and a cortex composed of cell bodies. The rhinophoral ganglıa of M. remanei are not subdivided into cortex and medulla; in- stead the nuclei are distributed homogeneously all over the ganglion (see NEUSSER et al. 2006, fig. 3d). Serial sec- tions of Hedylopsis spiculifera, H. ballantinei and M. re- manei show only a single nerve (approx. 5-10 um in di- ameter) that connects the cerebral ganglion to the rhinophoral one. In one specimen of Tantulum elegans ex- amined, we found two nerves connecting the cerebral gan- glion with the rhinophoral ganglion (Fig. 1). Both nerves are thin (approx. 7 um in diameter) and lie close togeth- er (distance between them approx. 31m). Nevertheless, the transition between the cerebral ganglion and the rhinophoral ganglion is well identifiable due to the pres- ence of dark stained fibres (Fig. 1A, D). 3.2. Sensory organs 3.2.1. Hancock’s organ and nerve Paired, small and ciliated invaginations posterior to the head appendages and innervated by cerebral nerves are present in Tantulum elegans (see NEUSSER & SCHRÓDL 2007, fig. 4b). Neither such organs of similar shape could be detected in Hedylopsis spiculifera, H. ballantinei and Microhedyle remanei, or cerebral nerves innervating the region where Hancock’s organs are present in other acochlidian species. 3.2.2. Eyes, optic nerves and optic ganglia Asperspina murmanica and Microhedyle remanei are eye- less and lack any optic nerve or optic ganglion. Both Hedv- lopsis species have pigmented lens eyes (Fig. 3A, B) that, however, differ in size and relative position. The eyes of H. spiculifera are clearly visible externally (Fig. 2A, B) from dorsal and lateral and reach up to 25 um in diame- ter (Fig. 3A). They are located on the rather lateral side of the head (Fig. 2B), and are in some distance posterior to the rhinophores (Fig. 2A, B) and anterior of the cere- bral ganglia. In contrast, the eyes of HA. ballantinei are hardly detectable by external view (Fig. 2C) even though they are slightly larger (approx. 30 um in diameter) (Fig. 3B). Furthermore, they are situated closer together and are just posterior to the rhinophores (Fig. 2C). The optic nerves show approx. 6-7 um in diameter in both species (Fig. 3A, B). They arise from the rhinophoral ganglia and are highly undulated. An optic ganglion is absent in A, spi- culifera as well as in H. ballantinei. In contrast, Tantu- lum elegans develops a very short and thin optic nerve (ap- prox. 3 um in diameter) leading to a reduced unpigment- ed eye of approx. 20 um in diameter (Figs. 1, 3C). The optic nerve arises from a small optic ganglion (approx. 18 um in diameter) that is subdivided into the outer cortex and the inner medulla (Fig. 3D). It is attached laterally to the cerebral ganglion, both of which are surrounded by a thin layer of connective tissue (Fig. 3D). No nerves can be detected by light microscope examination connecting the cerebral with the optic ganglion. 3.3. Aggregates attached to the cerebral ganglia 3.3.1. “Lateral bodies” A “lateral body” as defined herein consists of a more or less hemispherical cluster of cells that is lying laterally on the surface of each cerebral ganglion. Under a light mi- croscope, the cells of the “lateral bodies” cannot be dis- tinguished from the neuron bodies situated in the cortex of the cerebral ganglion. Each “lateral body” is surround- ed by a separate, relatively thin sheath of connective tis- sue and together with the cerebral ganglion by a second common and thick one. “Lateral bodies” are present in Hedylopsis spiculifera (Fig. 4A), H. ballantinei (Fig. 4B) and Asperspina murmanica (Fig. 4C). The “lateral body” lacks any subdivision. The nuclei are more or less uni- formly distributed over the entire “lateral body”. There are no nerves visible under the light microscope connecting the cerebral ganglion with the “lateral body”, and there are no nerves arising from the latter. None of the speci- mens examined of Microhedyle remanei and Tantulum el- egans had “lateral bodies”. 3.3.2. Cells near the cerebral commissure Additionally, we could find several cells of uncertain ori- gin and function dispersed in the connective tissue above the cerebral commissure in Asperspina murmanica (Fig. 4D). In contrast to the “lateral bodies”, these cells are not tightly attached to each other, and are not enclosed by an individual sheath of connective tissue. No data about the presence or absence of these cells can be given for Hedy- lopsis spiculifera, H. ballantinei and Tantulum elegans, due to very compressed tissue layers. 304 Timea P. NEUSSER et al.: Cerebral features in Acochlidia Fig. 1. Double cerebro-rhinophoral connective in Zantulum elegans. Four consecutive cross sections of series ROM N° SEI, 3.sli- de, 6. ribbon, section N° 17-20. A: section N° 17, first cerebro-rhinophoral connective. B and C: section N° 18 and 19, respecti- vely, without connective. D: section N° 20, second cerebro-rhinophoral connective. cg cerebral ganglion; ey eye: rhg rhinophoral ganglion; arrow, indicates fibres of the cerebro-rhinophoral connective. Scale bars A-D: 15 um. 4. DISCUSSION 4.1. Rhinophoral ganglia and number of cerebro-rhinophoral connectives The presence of rhinophoral ganglia were reported for Hedylopsis spiculifera and Tantulum elegans (see RANKIN 1979; WAwRA 1989), but both descriptions lack histolog- ical data of the rhinophoral ganglia. Recently, rhinophoral ganglia were described in detail for Hedylopsis ballanti- nei (see SOMMERFELDT & SCHRODL 2005), Microhedyle re- manel (see NEUSSER et al. 2006), 7. elegans (see NEUSS- ER & SCHRODL 2007) and Pontohedyle milaschewitchii (see JORGER et al. in press). Due to their position anterodor- sally of the cerebral ganglia and their similar innervation the homology of the rhinophoral ganglia can be assumed for all acochlidian species studied herein. In contrast to Hedylopsis species, Asperspina murmanica and T. elegans, rhinophoral ganglia of P milaschewitchii and M. remanei are not separated into medulla and cortex. The presence Bonner zoologische Beitráge 55 (2006) 305 Fig. 2. Position of eyes in different acochlidian species, external view. A: Hedylopsis spiculifera, dorsal view, length 3.5 mm. B: Hedylopsis spiculifera, lateral view, length 3.5 mm. C: Hedylopsis ballantinei, lateral view, length 5 mm. D: Pontohedyle mila- schewitchii, dorsal view, length 2.5 mm. ey eye; It labial tentacle; rh rhinophore. of rhinophoral ganglia within P milaschewitchii that is lacking any rhinophores might be explained by a modi- fied, e.g. neurosecretory function. Microhedyle remanei, however, possesses rhinophores and cell bodies evenly dis- tributed within the rhinophoral ganglia. Of all the specimens here studied, the double connection between the cerebral ganglia and rhinophoral ganglia could only be detected in one specimen of Tantulum ele- gans, and is only clearly visible on the right side of the nervous system. Unfortunately, the identification of these thin nerves depends critically upon preservation and stain- ing conditions as well as on the cutting plane. Tiny nerves can thus be overlooked and easily misinterpreted, or be invisible even on semi-thin serial sections. While “detect- ed” usually means “present”, “not detected” does not nec- essarily mean “absent”. The cerebro-rhinophoral connec- tive has been identified by the presence of dark stained fibres. HASZPRUNAR (1985, figs. 19, 20) described simi- lar fibres occurring at the transition between two differ- ent ganglia in Discotectonica discus Philippi, 1844. A dou- ble cerebro-rhinophoral connective has also been found in Pontohedyle milaschewitchii (see JORGER et al. in press); both nerves are even thinner than those in 7. elegans. There is no reliable data on further acochlidians. HASZPRUNAR & HUBER (1990) described a double cere- bro-rhinophoral connective for the enigmatic opistho- branchs Rhodope veranii Kólliker, 1847 and Rhodope transtrosa Salvini-Plawen, 1989, as well as a double con- nective attaching the cerebral ganglion with the procere- brum in the pulmonate Smeagol manneringi Climo, 1980. In fact, the double cerebro-rhinophoral connective of the acochlidian cns resembles the general pulmonate condi- 306 Timea P. NEUSSER et al.: Cerebral features in Acochlidia Fig. 3. Eyes and optic ganglion (cross sections). A: Pigmented eye in Hedylopsis spiculifera ZSM N° 20070391. B: Pigmented eye in Hedylopsis ballantinei ZSM N° 20004766/1. C: Unpigmented eye in Tantulum elegans ROM N° 8E1. D: Optic ganglion attached to the cerebral ganglion in Tantulum elegans ROM N° SEI. cg cerebral ganglion; ey eye; og optic ganglion: on optic ner- ve; rhg rhinophoral ganglion. Scale bars A-D: 15 um. tion (VAN MoL 1967). Therefore, the potential homology of acochlidian rhinophoral ganglia to the procerebrum of pulmonates should be investigated in detail. 4.2. Sensory organs 4.2.1. Hancock’s organ We were not able to detect any Hancock’s organ like struc- tures in the species examined herein except for Tantulum elegans which shows a pair of epidermal folds on the side of the head (NEUSSER & SCHRODL 2007). Such folds were reported for Pontohedyle milaschewitchii and Microhedvle glandulifera and regarded as Hancock’s organs by EDLINGER (1980a, b), 1.e. as true homologues of the pri- mary chemosensory organs in architectibranchs and cephalaspids (see MIKKELSEN 1996). According to their similar position, cerebral innervation, (although more tiny) structure, and probable sensory function, a general homol- ogy can be suspected. Some doubts persist, such as the Bonner zoologische Beiträge 55 (2006) 307 Fig. 4. Aggregates attached to the cerebral ganglia (cross sections). A: “Lateral body” in Hedylopsis spiculifera ZSM N* 20070391. B: “Lateral body” in Hedylopsis ballantinei ZSM N° 20004766/1. C: “Lateral body” in Asperspina murmanica ZSM N° 20062163. D: Cells above cerebral commissure in Asperspina murmanica ZSM N° 20062163. cc cerebral commissure; cg cerebral ganglion; Ib “lateral body”; arrow, cells near cerebral commissure. Scale bars A-D: 15 um. yet unclear homology of euthyneuran cerebral nerves, the unknown origin of the Acochlidia and reports of acochlid- ian “Hancock's organs” from only a few and supposedly derived microhedylid species, 1.e. P milaschewitchii and M. glandulifera, and the enigmatic 7. elegans. 4.2.2. Eyes, optic nerves and optic ganglia In the past, the description of acochlidian eyes often was limited to the affirmation of presence or absence of these sensory organs. Eyes are absent in all Asperspina species, Microhedyle remanei, Ganitus evelinae Marcus, 1953, Paraganitus ellynnae Challis, 1968 and Pontohedyle ver- rucosa Challis, 1970 (see CHALLIS 1968, 1970; KUDIN- SKAYA & MINICHEV 1978; MARCUS 1953; Morse 1976; SALVINI-PLAWEN 1973; SWEDMARK 1968). Our results show that the position, size and development of eyes in Acochlidia examined herein differ considerably. 308 Timea P. NEUSSER et al.: Cerebral features in Acochlidia The eyes of Hedylopsis spiculifera are clearly visible ex- ternally from a dorsal and lateral view. In the freshwater acochlidian species Strubellia paradoxa (Strubell, 1892) and Acochlidium fijiense Haynes & Kenchington, 1991 the eyes are clearly observable only in lateral view (unpubl. data of MS). In contrast, the eyes of the marine Micro- hedyle glandulifera (see KOWALEVSKY 1901; MARCUS & Marcus 1955; ODHNER 1952), Hedvlopsis ballantinei (Fig. 2C) and Pontohedyle milaschewitchii (Fig. 2D) are externally not that clearly visible through the head tissue. WESTHEIDE & WAwRA (1974) observed that eyes of Parhedvle cryptophthalma (Westheide & Wawra, 1974) were not visible externally in living specimens, and only as two small pigmented spots in preserved specimens. Eyes in Pseudunela cornuta (Challis, 1970) are poorly de- veloped and not visible externally (CHALLIS 1970, as Hedylopsis cornuta). The eyes of Hedylopsis spiculifera and H. ballantinei are both located dorsolaterally in the body cavity; while the eyes of H. ballantinei are situated at the base of the rhinophores, in HA. spiculifera they are somewhat more posteriorly. A similar dorsolateral eye position at or close to the base of the rhinophores is already known from the limnic acochlidian species Acochlidium amboinense Strubell, 1892, Palliohedvle weberi (Bergh, 1895) and Strubellia paradoxa (see BERGH 1895; BUCKING 1933; KUTHE 1935). In contrast, the eyes of Pontohedyle mi- laschewitchii are located more posteriorly and closer to- gether (Fig. 2D). WESTHEIDE & WAWRA (1974) described a similar eye position in the marine acochlidian Parhedyle cryptophthalma. The optic nerve 1s short in Strubellia paradoxa (see KUTHE 1935). The well-developed eyes of Acochlidium am- boinense, Palliohedvle weberi and $. paradoxa were de- scribed as attached anterodorsally to anterolaterally on the cerebral ganglia (BERGH 1895; BUCKING 1933; KÜTHE 1935), thus the optic nerves are probably short as well. The eyes of Pontohedyle milaschewitchii are directly at- tached to the cerebral ganglia (JÖRGER et al. in press), as are the eyes of Parhedvle cryptophthalma, Microhedyle nahantensis (Doe, 1974), M. elandulifera and M. odhneri (Marcus, 1955) (see DOE 1974; MARCUS & MARCUS 1955; WESTHEIDE & WAWRA 1974). The optic nerve is moder- ately long but thin in Zantulum elegans, while long and thick in both Hedvlopsis species. The long optic nerves observed herein may be phylogenetically informative in Acochlidia. All eyes described for Acochlidia are pigmented, except those of Tantulum elegans (present study) and of Micro- hedvle nahantensis (see DOE 1974). The “poorly devel- oped” eyes of Pseudunela cornuta described by CHALLIS (1970) should be reinvestigated. The eye sıze differs within the species: whereas eyes of Hedylopsis spiculifera and H. ballantinei measure approx. 25 and 30 um, respectively, eyes in Pontohedyle milasche- witchii reach approx. 20 um (JÖRGER et al. in press). The largest eye size known from an acochlidian species is 0.52 mm and was reported for the limnic Palliohedyle weberi (see BERGH 1895). The optic ganglion in Tantulum elegans was first described by NEUSSER & SCHRÖDL (2007) and is regarded to be a true ganglion with subdivision into cortex and medulla (see NEUSSER et al. 2006). More specifically, it is enclosed in a thin layer of connective tissue together with and at- tached to the cerebral ganglion. This feature should not be confused with the “lateral bodies” described in the pres- ent study, since the latter are lying inside the thick layer of connective tissue from the cerebral ganglion (see be- low). So far there are only two reports of ganglia being surrounded by a common layer of connective tissue with the cerebral ganglia: the rhinophoral ganglia of 7. elegans (see NEUSSER & SCHRODL 2007), and the rhinophoral gan- glia of Pontohedyle milaschewitchii (JORGER et al. in press). The presence of an optic ganglion only in 7. elegans is surprising, since eyes are unpigmented in this species, while for species possessing more well-developed eyes (e.g. both Hedylopsis species and Pontohedyle milasche- witchii) this character is lacking. Either there are some un- known sensory abilities involved in at least one ontoge- netic stage, or both eyes and optic ganglia are evolution- ary remnants of organs in the process of being reduced. The optic ganglia of Zantu/um do no more fuse with the rhinophoral ganglia, as may be the case in both Hedylop- sis species with large rhinophoral ganglia bearing optic nerves. We urgently need ontogenetic evidence for the de- velopment of acochlidian central nervous structures. The presence of optic ganglia, the origin and length of op- tic nerves, eye position in terms of situation and proxim- ity to the cerebral ganglion, as well as eye size and struc- ture should be reinvestigated in all acochlidian species, since these may be easily accessible and phylogenetical- ly informative characters (see MIKKELSEN 1996). 4.3. Aggregates attached to the cerebral ganglia 4.3.1. "Lateral bodies” SOMMERFELDT & SCHRÖDL (2005) described “dorsal bod- ies” attached to the cerebral ganglion in the acochlidian Hedylopsis ballantinei. We herein confirm the presence of such organs for both Hedvlopsis species and A. mur- manica. Their position is, however, more lateral than dor- sal. We thus propose to use the term “lateral bodies” for Bonner zoologische Beiträge 55 (2006) 309 such acochlidian structures until more detailed and com- parative data are available to assess their homology to pul- monate dorsal bodies. The “lateral bodies” of the re-examined acochlidian species are characterized by a group of neuronal cells that are enclosed within the thick connective tissue layer sur- rounding the cerebral ganglion. The dorsal bodies of ba- sommatophoran pulmonates consist of a pair of similar neuronal cell clusters that are, however, enclosed in a thin sheath of connective tissue, and are situated dorsally on the cerebral ganglia. Basommatophoran dorsal bodies can lie close together and appear as one group in Helisoma Swainson, 1840 and Planorbarius Duméril, 1806, or they can be distinguished as two separate tissue masses, as in Ancylus Mueller, 1774, Lymnaea Lamarck, 1801 and Siphonaria Sowerby, 1823 (SALEUDDIN 1999; SALEUDDIN etal. 1997; TAKEDA & OHTAKE 1994). SOMMERFELDT & SCHRÖDL (2005) described the “lateral bodies” of Hedylopsis spiculifera and H. ballantinei be- ing subdivided into an outer cortex and an inner medul- la. According to SALEUDDIN (1999), most of the dorsal bodies of basommatophoran pulmonates develop a cor- tex with nuclei and an inner medulla with cell processes that lie very close to the cerebral ganglia. In “lateral bod- 1es” of H. spiculifera, H. ballantinei and Asperspina mur- manica, no such clear subdivision into cortex and medul- la was found; instead all nuclei are distributed more or less uniformly. Similarly, the basommatophoran pulmonate Siphonaria pectinata Linnaeus, 1758 is described to pos- sess dorsal bodies without clear separation into cortex and medulla (SALEUDDIN et al. 1997). The function of the “lateral bodies” in Hedylopsis spiculif- era, H. ballantinei and Asperspina murmanica is unclear. Due to the absence of visible nerves arising from these aggregations, the “lateral bodies” are possibly not senso- ry but secretory organs. The role of dorsal bodies in pul- monates as an endocrine organ involved in female repro- duction is quite well known (SALEUDDIN 1999). Further- more a putative endocrine gland, called the juxtagan- glionar organ, has been described in several opisthobranch species (e.g. SwITZER-DUNLAP 1987). However, the ho- mology of these structures is still unclear. Future studies by means of transmission electron microscopy and (im- muno)histochemical studies are needed to understand ho- mologies and functions. Disregarding our deficient knowledge, within acochlidians the presence of “lateral bodies” in members of Hedylopsidae, Asperspinidae and Tantulidae versus their absence in two members of Mi- crohedylidae (Pontohedyle milaschewitchii, Microhedyle remanei) may represent characters with a phylogenetic sig- nal. 4.3.2. Cells near the cerebral commissure For the first time in an acochlidian species we describe several cells that are loosely dispersed within the connec- tive tissue above the cerebral commissure in Asperspina murmanica. Due to its position such a cell aggregation re- sembles the dorsal bodies of stylommatophoran pul- monates (e.g. Theba pisana Mueller, 1774, Helix asper- sa Mueller, 1774 and Achatina fulica Ferussac, 1821) which were described as diffusely scattered cells within the connective tissue sheath of the cerebral ganglion and located near the cerebral commissure (SALEUDDIN 1999; SALEUDDIN et al. 1997; TAKEDA & OHTAKE 1994). The presence, structure, origin and function of these cells in acochlidians cannot be revealed by light microscopy alone but requires ultrastructural studies. Acknowledgements. We thank the Royal Ontario Museum (Canada) and the Swedish Museum of Natural History (Sweden) for providing material for re-examination. Gerhard Haszprunar (ZSM) is thanked for helpful discussions. We are grateful to Liz Atwood (University of Washington) for improving the English. Two anonymous referees provided helpful comments on the manuscript. 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University Vienna: Dissertation N° 17335. Wawra, E. 1988. Sand-Opisthobranchia aus dem Golf von Ben- galen. Annalen des Naturhistorischen Museums in Wien B 90: 427-231. Wawra, E. 1989. Zur Kenntnis der interstitiellen Opisthobran- chierart Hedylopsis spiculifera (Kowalevsky) (Mollusca, Gastropoda). Zoologica Scripta 18(3): 397-403. WESTHEIDE, W. & Wawra, E. 1974. Organisation, Systematik und Biologie von Microhedvle cryptophthalma nov. spec. (Ga- stropoda, Opisthobranchia) aus dem Brandungsstrand des Mit- telmeeres. Helgolander Wissenschaftliche Meeresuntersuchun- gen 26: 27-41. Authors’ addresses: Timea P. NEUSSER (corresponding author), Zoologische Staatssammlung Munchen, Munch- hausenstrasse 21, 81247 Miinchen, Germany, E-Mail: ti- mea-neusser@gmx.de; Katharina JORGER, E-Mail: k_jo- erger(@hotmail.com; Michael SCHRÖDL, E-Mail: schro- edl(Wzi.biologie.uni-muenchen.de Bonner zoologische Beiträge | Band 55 (2006) Heft 3/4 Seiten 311-318 Bonn, November 2007 The cephalic sensory organs of Acteon tornatilis (Linnaeus, 1758) (Gastropoda Opisthobranchia) — cellular innervation patterns as a tool for homologisation* Sid STAUBACH & Annette KLUSSMANN-KOLB!) Institute for Ecology, Evolution and Diversity — Phylogeny and Systematics, J. W. Goethe-University, Frankfurt am Main, Germany *Paper presented to the 2nd International Workshop on Opisthobranchia, ZFMK, Bonn, Germany, September 20th to 22nd, 2006 Abstract. Gastropoda are guided by several sensory organs in the head region, referred to as cephalic sensory organs (CSOs). This study investigates the CSO structure in the opisthobranch, Acteon tornatilis whereby the innervation pat- terns of these organs are described using macroscopic preparations and axonal tracing techniques. A bipartite cephalic shield and a lateral groove along the ventral side of the cephalic shield was found in 4. tornatilis. Four cerebral nerves can be described innervating different CSOs: N1: lip, N2: anterior cephalic shield and lateral groove, N3 and Nele: posterior cephalic shield. Cellular innervation patterns of the cerebral nerves show characteristic and con- stant cell clusters in the CNS for each nerve. We compare these innervation patterns of A. tornatilis with those described earlier for Haminoea hydatis (STAUBACH et al. in press). Previously established homologisation criteria are used in order to homologise cerebral nerves as well as the organs innervated by these nerves. Evolutionary implications of this homologisation are discussed. Keywords. Haminoea hydatis, Cephalaspidea, axonal tracing, homology, innervation patterns, lip organ, Hancock’s organ. 1. INTRODUCTION Gastropoda are guided by several organs in the head re- gion which are assumed to have primarily chemo- and mechanosensory functions (AUDESIRK 1979; Davis & MATERA 1982; BICKER et al.1982; EMERY 1992; CHASE 2000; CROLL et al. 2003). In Opisthobranchia, these cephalic sensory organs (CSOs) present an assortment of forms including rhinophores, labial tentacles, oral veils, Hancock’s organs and cephalic shields. Recent investiga- tions of CSOs in Opisthobranchia have focussed prima- rily on functional aspects (CROLL 1983; BOUDKO et al. 1999; CROLL et al. 2003) while homology of the differ- ent types of CSOs in different taxa has never been inves- tigated in detail. We want to clarify their homology in sep- arate evolutionary lineages so as to elucidate key ques- tions regarding character evolution and phylogeny. Acteon tornatilis belongs to the subgroup Acteonoidea, formerly ascribed to the basal Cephalaspidea (ODHNER 1939, BURN & THOMPSON 1998). However, recent inves- tigations have either excluded the Acteonoidea from the Opisthobranchia (MIKKELSEN 1996) or proposed a sister group relationship of Acteonoidea and the highly derived Nudipleura (VONNEMANN et al. 2005) thus, rendering the phylogenetic position of Acteonoidea within Opistho- branchia unsettled. Acteonoidea are characterised by the presence of a prominent cephalic shield. This structure 1s also present in Cephalaspidea and has been considered to be an apomorphie of the Cephalaspidea (SCHMEKEL 1985). However, the structure of the cephalic shields differs con- siderably in Cephalaspidea and Acteonoidea with the lat- ter possessing two distinct hemispheres while the cephal- ic shield in the Cephalaspidea possesses uniform structure. Therefore, common origin of both types of cephalic shields and thus homology is questionable. Further CSOs have been described in Acteonoidea and Cephalaspidea such as lip organ and Hancock’s organ (RUDMAN 1971A; RUDMAN 1971B; RUDMAN 1972a, b; RUDMAN 1972c; EDLINGER 1980). Since the presence of these organs in members of the genus Acteon has been disputed by different authors (EDLINGER 1980; SCHMEKEL 1985), absolute clarification is certainly necessary . The intention of this study is to de- scribe the structure emphasizing the innervation of the CSOs in the acteonid 4. tornatilis. Our descriptions fo- cus on the cellular innervation patterns reconstructed for 312 Sid STAUBACH & Annette KLUSSMANN-KOLB: Cephalic sensory organ in Aceton the cerebral nerves using axonal tracing. In an earlier study (STAUBACH et al. in press) these cellular innervation pat- terns were shown to be more adequate in homologising cerebral nerves than ganglionic origins of nerves (HUBER 1993). By comparising the innervation patterns in A. tor- natilis to previously published data on A. hydatis (STAUBACH et al. in press), we want to survey whether the preliminary characteristic cell clusters in the central nerv- ous system (CNS) of both taxa can be identified by ho- mologising cerebral nerves across taxa. Based on the ho- mologisation of the nerves innervating the CSOs we want to clarify if A. tornatilis has homologous structures to the CSOs of Cephalaspideans. It is our intent interest that we shed light on the phylogenetic position and evolutionary history of Acteonoidea within the Opisthobranchia for fu- ture studies. 2. MATERIALS AND METHODS 2.1. Specimens A. tornatilis (Fig. 1A) were collected in the wild at St. Michel en Greve (Brittany, France). They were then stored alive at our lab in Frankfurt. Fourty specimens measur- ing a shell length between 15 and 20 mm were traced di- rectly (5 to 15 days after collecting) and five were fixed for SEM. 2.2. Tracing studies Animals were relaxed with an injection of 7 % magne- sium chloride. The central nervous system consisting of the cerebral, pleural and pedal ganglia was removed and placed in a small Petri dish containing filtered artificial seawater (ASW; Tropic Marin, Rebie-Bielefeld; GER- MANY). We then followed the procedures from CROLL & BAKER (1990) for Ni2*-Iysine (Ni-Lys) tracing of ax- ons. Briefly, the nerves of the right cerebral ganglion were dissected free from the connective tissue. The nerves were cut and the distal tip was gently drawn into a glass mi- cropipette using suction provided by an attached 2.5 ml syringe. Subsequently, the saline in the micropipette was replaced by a Ni-Lys solution (1.9g NiCI-6H)0, 3,5 g L- Lysine freebase in 20 ml double distilled H,O). The prepa- ration was then incubated for 12-24 hours at 8° C to al- low transport of the tracer. The micropipette was then re- moved and the ganglia were washed in ASW three times. The Ni-Lys was precipitated by the addition of five to ten drops of a saturated rubeanic acid solution in absolute Di- methylsulfoxide (DMSO). After 45 minutes the ganglia were transferred to 4 % paraformaldehyde (PFA) and fixed for 4-12 hours at 4° C. Thereafter the ganglia were dehy- drated in an increasing ethanol series (70/80/90/99/99% 10 minutes each), cleared in methylsalicylate and mount- ed on an objective slide dorsal side up in Entellan (VWR International) and covered with a cover slip. Ten replicates were prepared for each cerebral nerve of A. tornatilis. Samples with only a partial staining of the nerve were not used because of possible incomplete innervation patterns. Our criterion for a well-stained preparation was a dark blue stained nerve indicating intact axons (FREDMAN 1987). The Ni-Lys tracings were analysed by light microscopy (Le- ica TCS 4D). Camera lucida drawings were digitalised fol- lowing the method of COLEMAN (2003) adapted for Corel- DRAW 11. The somata in the innervation scemes occurs in all replicates. Somata only occurring in single samples are not considered part of the schematics. The axonal path- ways are estimated over all replicates. Additionally, we tested for asymmetries making axonal tracings (n = 2 to 3) for each cerebral nerve of the left cerebral ganglion. 2.3. Scanning electron microscopy studies The specimens were relaxed by an injection of 7 % Mg- Cl, in the foot. Thereafter, the entire head region was dis- sected from the rest of the animal. The CSOs were fixed in 2,5 % glutaraldehyde, 1 % paraformaldehyde in 0,1M phosphate buffer (pH 7,2) at room temperature. For the SEM, the fixed CSOs were dehydrated through a graded acetone series followed by critical point drying (CPD 030, BAL-TEC). Finally, they were spattered with gold (Sput- ter-Coater, Agar Scientific) and examined with a Hitachi S4500 SEM. All photographs were taken using DISS (Dig- ital Image Scanning System — Point Electronic) and sub- sequently adjusted for brightness and contrast with Corel PHOTO-PAINT 11. 3. RESULTS 3.1. Organisation and innervation of the cephalic sen- sory organs A. tornatilis possesses a prominent bipartite cephalic shield (cs) in which each hemisphere of this shield is divided in- to an anterior and a posterior lobe (Figs. 1A and B). Eyes are embedded deeply within the tissue of the shield. Along the lateral margin of the anterior lobe of the cephalic shield a groove is present (Fig. 1B, 2A). Hidden under the cs and above the foot, the mouth opening is situated at the me- dian frontal edge (Fig. 2B) surrounded by the lip (not vis- ible in Figure 2B). We found four nerves innervating the CSOs (Fig.1B). The NI (Nervus oralis) provides inner- vation to the lip and small median parts of the anterior cephalic shield. The bifurcated N2 (Nervus labialis/labio- tentacularis) innervates the complete anterior cephalic shield whereby the groove at the ventral anterior lobe of the cephalic shield is especially innervated. The small N3 (Nervus tentacularis/rhinophoralis) innervates a little re- Bonner zoologische Beiträge 55 (2006) 313 gion of the posterior cephalic shield. The Nele (Nervus clypei capitis) innervates the largest hind part of the pos- terior cephalic shield. We could not detect a lip organ (Fig. 2B), which according to EDLINGER (1980) should comprise two small lobes on the cephalic shield above the mouth. A Hancock 's organ described by EDLINGER (1980) for A. Tornatilis, here a folded structure separated from the cephalic shield was likewise not found in the present study. 3.2. Tracing studies By conducting the axonal tracing studies we were able to reconstruct cellular innervation patterns for the four cere- bral nerves of 4. tornatilis. Ten replicate tracings were per- formed each for the NI N2, N3 and Nelc using only the nerves of the right cerebral ganglion. The characteristic patterns of labelled somata for all nerves are shown in Fig- ure 3A-D, including the approximate pathways of the stained axons. The identified clusters were named with ab- breviations signifying the ganglion in which they are lo- cated, the nerve filled and a number indicating the order of their description (for example, Cnlc3: Cerebral Nervus labialis cluster 3). Nerve cells are grouped in clusters on the basis of their close positioning in the ganglia and the tight fasciculation of their axons projecting into the filled nerve. Asymmetries for tracings of the left nerves could not be detected. Fig. 1. bral nerves (excluding the optical nerve) and the cephalic sensory organs of Haminoea hydatis and Acteon tornatilis. Only the right cerebral nerves are shown. NI Nervus oralis, N2 Nervus labialis, N3 Nervus rhinophoralis, Ncle Nervus clypei capitis, ey eye, gr groove, al anterior lobe, pl posterior lobe, sh shell, cs cephalic shield, f foot. For the NI (n = 10) we identified six cerebral clusters (Cnocl-6) and one pedal cluster (Pdnocl) in each sam- ple (Fig. 3A). The variation between the samples was re- stricted to very few somata in some clusters. The cerebral clusters were distributed over the whole cerebral ganglion. The pedal cluster Pdnocl was located on the anterior mar- gin of the pedal ganglion above the pedal commissure. The innervation pattern of the N2 (n=10) consists of five cere- bral clusters (Cnlel-5) and three pedal clusters (Pdnlc1- 3) (Fig. 3B). The cerebral clusters show distinct spatial separations and are easy to identify. The third traced cere- bral nerve (n= 10) was the N3. Six cerebral (Cnrc1-6) and three pedal clusters (Pdnrc 1-3) were identified (Fig. 3C). We found an additional single cluster (Cc/nrc/) and a sin- gle soma in the left cerebral ganglion (see arrows in Fig. 3C). The contralateral cluster was located at the base of the N2 whereas the single soma was found at the root of the cerebral commissure. We observed slight intraspecif- ic variability between the ten samples which amounted on- ly to very few somata in some clusters. In the Nelc, the innervation (n = 10) pattern consisted of five cerebral clus- ters (Cncel-5) and a single soma at the lateral margin of the cerebral ganglion above the pedal connective (Fig. 3D). Additionally we found four pedal clusters (Pdnec 1 - 4). The Nele had the highest amount of pedal clusters in all investigated nerves. The number of pedal somata, how- ever, was comparable to the number of pedal somata for the N2 innervation pattern (Fig 3B). Nclc B A: Photograph of Acteon tornatilis with the cephalic shield visible. B: Schematic illustration of the CNS, the four cere- 314 Sid STAUBACH & Annette KLUSSMANN-KOLB: Cephalic sensory organ in Aceton anterior Fig. 2. A. Lateral SEM photography of the groove at the ventral surface of the cephalic shield of Acteon tornatilis. cs cephalic shield, gr groove. B. Frontal SEM photography of the mouth region of Acteon tornatilis. cs — cephalic shield, mo — mouth, f— foot. 4. DISCUSSION The present study demonstrates the constancy of nervous structures in the opisthobranch mollusc. Throughout our investigation of several individuals of the acteonid, Acteon tornatilis we found uniform innervation patterns of the head region via four cerebral nerves, which can be attrib- uted to characteristic neuronal cell clusters in the UNS. These cellular innervation patterns in A. tornatilis show an extremely high congruence with the cellular innerva- tion patterns described for the four cerebral nerves of Haminoea hydatis (STAUBACH et al. in press). In the N1, the number of cerebral clusters as well as the position of these clusters to each other 1s the same in 4. tornatilis and H. hydatis. However, we found some dif- ferences in the size and number of somata when compar- ing both species. Additionally, we could not detect a pleu- ral, a parietal and a pedal cluster in A. tornatilis, which were described for A. hydatis. This may be due to the dif- ferences in the peripheral innervation area of the NI. In A. tornatilis it only provides for the lip and very small parts of the median cephalic shield whereas in /7. hydatis, it in- nervates the lip and large parts of the anterior cephalic shield. For the second nerve, the N2 (Nervus labialis), we nearly found no differences between the presence and dis- tributions of the cell clusters for both species. The only ostentatious difference was the lack of a single pedal so- ma and its contra-lateral analogue in 4. tornatilis. In the Nelc (Nervus clypei capitis), the difference between the two species was also reduced to the presence of a single cerebral soma in 4. tornatilis. In contrast to the three nerves described above, we found a prominent difference in the structure of the N3 when comparing Acteon and Haminoea. On the other hand, in HA. hydatis the N3 ter- minates in a rhinophoral ganglion. Such a ganglion is missing in 4. tornatilis. Hence, we expected considerable differences in the cellular innervation patterns for the N3 of these species. However, these differences were margin- al and only amounted to the lack of one single cell soma in the cerebral ganglion of A. tornatilis. This implies that basic innervation patterns of the N3 are probably the same in both species. Additional functions of the N3 processed in the rhinophoral ganglion can be proposed for A. hydatis. These functions are probably related to the Hancock's or- gan, which is innervated by nerves originating in the rhinophoral ganglion (STAUBACH et al. in press). We were unable to locate such an organ in A. tornatilis ın contrast to earlier descriptions (EDLINGER 1980). Upon comparing the innervation patterns presented here for A. tornatilis with those for A. hydatis (STAUBACH et al. in press) we find constant features of these patterns across species. This 1s congruent with other findings that neuronal structures in the central nervous system of mol- luscs and other invertebrates seem to be highly conserved (CROLL 1987; ARBAS 1991; HAYMAN-PAUL 1991; KUTSCH & BREIDBACH 1994; NEWCOMB et al. 2006). Hence, we postulate the N1 of A. tornatilis to be homologous to the N1 (Nervus oralis) described by HUBER (1993) for Cepha- laspideans. Additionally, we postulate homologies of the N2 and the N3 of 4. tornatilis to the N2 (Nervus labialis) and N3 (Nervus rhinophoralis) of Chepalaspideans. This is congruent to the assumption of HOFFMANN (1939) that the c3 (after VAYSSIERE 1880) of A. hydatis represents the Nervus labialis and the c4 represents the Nervus tentac- ularis, here a synonym for the Nervus rhinophoralis (HU- BER 1993). Our data cannot support EDLINGER’S (1980) de- scription of independent nerves for the lip organ (N1 af- ter Edlinger 1980) and the anterior Hancock ’s organ (N2 after Edlinger 1980). The Nele of 4. tornatilis also seems to be homologous to the Nele of Cephalaspideans (Huber Bonner zoologische Beiträge 55 (2006) 315 Cc N1 N2 \ 7. Ceinre1 N3 aa N. Opt. Be. N2 DÁ 3 Nelc N. Opt. AS cnc 1) Panic 1 Pdnic3 Pancc Pdneca Fig. 3. Schematic outline of cell clusters providing the N1 (A), N2 (B), N3 (C) and Nele (D) of Acteon tornatilis. The size and position of the somata were digitalized from a camera lucida drawing, the distribution of the axons are averaged from all replica- tes. Nl Nervus oralis, N2 Nervus labialis, N3 Nervus rhinophoralis, Nelc Nervus clypei capitis, N. opt. Nervus opticus, CG cere- bral ganglia, RhG rhinophoral ganglia, PIG pleural ganglia, PdG pedal ganglia. 316 Sid STAUBACH & Annette KLUSSMANN-KOLB: Cephalic sensory organ in Aceton 1993). HOFFMANN (1939) described the same nerve as the Nervus proboscidis. We define this nerve however, as Nervus clypei capitis according to HUBER (1993). Considering the homologisation of the cerebral nerves in light of their neurological origin, neuro-anatomics and nervous innervation patterns, we postulate hypotheses of homologies respective of the organs innervated by these nerves. Thus, we consider the lip of 4. tornatilis to be ho- mologous to the lip of Cephalaspideans (HUBER 1993) since both organs are innervated by the Nl. The same holds true for the small median parts of the cephalic shield in Acteon and the anterior cephalic shield of Haminoea. We could not find a lip organ in A. tornatilis as described by EDLINGER (1980), but we detected a groove at the ven- tral side of the anterior cephalic shield. This groove 1s in- nervated by the N2 as is the lip organ of Cephalaspideans (Huber 1993). Therefore, we postulate this groove to be homologous to the lip organ. This hypothesis is also sup- ported by data on immunoreactivity against several neu- rotransmitters. In the groove of A. tornatilis as well as in the lip organ of A. hydatis, characteristic sub-epidermal sensory neurons containing catecholamines could be found in high density indicating that both organs are involved in contact chemoreception (S. FALLER, Frankfurt, pers. comm. 2007). The N2 of Haminoea is divided into two branches which are described as two single nerves by EDLINGER (1980). The first or inner branch provides the lip organ as de- scribed earlier. The second, outer branch ıs related to the anterior Hancock’s organ (EDLINGER 1980; HUBER 1993). In Acteon we also found two branches of the N2: the in- ner one providing the largest part of the groove whereas the outer branch is restricted to a small region between the anterior and posterior lobe of the cephalic shield. Therefore, this latter region may be homologous to the an- terior Hancock’s organ of A. hvdatis and not to the pos- terior Hancock’s organ as described by EDLINGER (1980). The N3 of 4. tornatilis provides a large part of the pos- terior cephalic shield but no identifiable posterior Han- cock’s organ. Additional immunohistochemical and ultra- structural investigations could also not detect a posterior Hancock’s organ in A. tornatilis (S. FALLER, Frankfurt, pers. comm. 2007; GOBBELER & KLUSSMANN-KOLB in press). The posterior parts of the cephalic shields in Acteon and Haminoea are probably equally homologous as both where innervated by the Nelc. The lack of a posterior Hancock’s organ in A. tornatilis might be due to three different reasons: 1. the ancestor of A. tornatilis never had a posterior Hancock's organ; 2. the posterior cephalic shield of 4. tornatilis may be a homol- ogous structure to the posterior Hancock's organ of HA. hy- datis; and 3. the posterior Hancock's organ has second- arily been reduced in A. tornatilis. The first hypothesis 1s rather implausible since we found a distinct N3 with conserved cellular innervation patterns in the central nervous system. If the ancestor of A. tor- natilis never had a posterior Hancock’s organ, this nerve and associated neural structures should be lacking. More- over, a Hancock’s organ has been described for other Acteonoidea (RUDMAN 1971a, b; RUDMAN 1972a, b; RuD- MAN 1972c). If we consider the second explanation for lack of a posterior Hancock’s organ in A. tornatilis, we imply that the posterior cephalic shield in this species, in- nervated by the N3, presents a sensory organ as the Han- cock ’s organ in Cephalaspidea. However, immunohisto- chemical and ultrastructural investigations of the respec- tive epithelia in A. tornatilis do not indicate a sensory func- tion at all (S. FALLER, Frankfurt, pers. comm. 2007; GÖBBELER & KLUSSMANN-KOLB in press). We reject this hypothesis of homology of the posterior cephalic shield in A. tornatilis and posterior Hancock's organ in A. hy- datis since we found no evidence for a function of the pos- terior cephalic shield as an olfactory sensory organ. More- over, the posterior cephalic shield is mostly innervated by the Nele and not by the N3. The third hypothesis regard- ing the reduction of a Hancock’s organ seems to be the most plausible when the habitat and the food sources of A. tornatilis in comparison to H. hydatis are considered. The posterior Hancock’s organ is believed to be an olfac- tory sensory organ (AUDESIRK 1979; EMERY 1992). H. hy- datis feeds on green algae which occur in patches in open water whereas A. tornatilis is a predator of soft inverte- brates living up to ten centimeters in solid sand (FRETTER 1939; Yonow 1989; own investigations). In such an en- vironment, an olfactory sensory organ is not plausible since olfaction or distance chemoreception is generally as- sociated with water currents, which are not substantial in a sandy substrate habitat. Here, a contact chemoreceptor, which is located near the edge of the cephalic shield is more plausible. This we witnessed in Acteon tornatilis via its display of a potentially chemoreceptive groove along the lateral margin of the anterior cephalic shield. This assumption of secondary reduction of the Hancock 's organ in the endobenthic A. tornatilis is also supported by the fact that a Hancock’s organ has been described for oth- er epibenthic Acteonoidea (e. g. Bullina, Micromelo, Hy- datina) (RUDMAN 1971a, b; RUDMAN 1972a, b; RUDMAN 1972a,b,c). Despite all discussion, homology of the described Han- cock’s organs to those in Cephalaspidea cannot undoubt- edly be proposed at this stage, particularly since data on innervation patterns in these acteonids are lacking to date. Moreover, current phylogenetic hypotheses (GRANDE et al. Bonner zoologische Beiträge 55 (2006) 317 2004; VONNEMANN et al. 2005) regardıng Opistho- branchia propose an independent origin of Acteonoidea and Cephalaspidea, indicating convergent development of these sensory organs in both evolutionary lineages. Fur- ther studies will have us utilizing cellular innervation pat- terns for CSOs in order to compare several taxa while ho- mologising the different types of CSOs in Opistho- branchia. This procedure will enable us to glean a better understanding of the evolution of these organs. Acknowledgements. Marc Hasenbank, Patrick Schultheiss and Christiane Weydig were constructive in locating the Acteon tor- natilis population at St. Michel en Greve. Thanks to Katrin Göbbeler and Simone Faller for their personal commitments. Roger Croll was always very helpful in discussing tracing pat- terns and CSOs. We are grateful to Angela Dinapoli, Adrienne Jochum and Alen Kristof for their comments on an earlier ver- sıon of this manuscript. This study was supported by the German Science Foundation, KL 1303/3-1 and by the Verein der Freunde und Förderer der Johann-Wolfgang-Goethe Universität. Also thanks to an unknown referee who provided valuable com- ments on the manuscript. REFERENCES ARBAS, E. 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STAUBACH, S.E., SCHÜTZNER, P., CROLL, R.P. & KLUSSMANN- Kors, A.Innervation patterns of the cerebral nerves in Hami- noea hydatis (Linnaeus 1758) (Gastropoda, Opisthobranchia) — A test for intraspecific variability. /n press, Zoomorphology 2007). VAYSSIERE, A. 1980. Recherches anatomiques sur les Mollusques de la famille des Bullidés Annales des Sciences Naturelle Zoo- logie 6: 9. VONNEMANN, V., SCHRODL, M., KLUSSMANN-KOLB, A. & WAGELE, H. 2005. Reconstruction of the phylogeny of the Opisthobranchia (Mollusca, Gastropoda) by means of 18S and 28S rDNA sequences. Journal of Molluscan Studies 71: 113-125. Yonow, N. 1989. Feeding observations on Acteon tornatilis (Lin- naeus) (Opisthobranchia, Acteonoidae). Journal of Molluscan Studies 55(1): 97-102. Author 's adresses: Sid STAUBACH (corresponding author) and Annette KLUSSMANN-KOLB, Institute for Ecology, Evo- lution and Diversity — Phylogeny and Systematics, J. W. Goethe-University, Siesmayerstraße 70, 60054, Frankfurt am Main, Germany. 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Inhalt SMITHSONIAN INSTITUTION LIBRARIES A 5543 3 9088 01431 Special Issue: Proceedings of the 2nd International Workshop on Opisthobranchia, | ZFMK, Bonn, Germany, September 20th to 22nd, 2006 AFFELD, Sven; WÄGELE, Heike; AVILA, Conxita; KEHRAUS, Stefan & KÖNIG, Gabriele M.: 181 Distribution of homarine in some Opisthobranchia (Gastropoda: Mollusca) DINAPOLI, Angela; TAMER, Ceyhun; FRANSSEN, Susanne; 191 NADUVILEZHATH, Lisha & KLUSSMANN-KOLB, Annette: Utility of H3-Genesequences for phylogenetic reconstruction — a case study of heterobranch Gastropoda Garcia, Francisco J.; DOMINGUEZ, Marta; & TRONCOSO, Jesús S.: 203 Biogeographic considerations of the Opisthobranchia (Mollusca: Gastropoda) fauna from the Brazilian littoral and nearby areas GOBBELER, Katrin & KLUSSMANN-KOLB, Annette: 228 Paddle cilia on the cephalic sensory organs (CSOs) of Opisthobranchia (Mollusca: Gastropoda) — genuine structures or artefacts? HÄNDELER, Katharina & WAGELE, Heike: 231 Preliminary study on molecular phylogeny of Sacoglossa and a compilation of their food organisms JENSEN, Kathe R.: 255 Biogeography of the Sacoglossa (Mollusca, Opisthobranchia) JORGER, Katharina M.; NEUSSER, Timea P & SCHRODL, Michael: 283 Re-description of a female Pontohedyle brasilensis (Rankin, 1979), a junior synonym of the Mediterranean P. wilaschemitchi (INowalevsky, 1901) (Acochlidia, Gastropoda) MUNIAIN, Claudia; ARDILA, Néstor E. & CERVERA, Juan Lucas: 291 Plenrobranchaea inconspicua Bergh, 1897 (Opisthobranchia: Pleurobranchidae): Redescription and distribution from Argentina and Colombia NEUSSER, Timea P.; JÖRGER, Katharina & SCHRODL, Michael: 301 Exploring Cerebral Features in Acochlidia (Gastropoda: Opisthobranchia) STAUBACH, Sid & KLUSSMANN-KOLB, Annette: 34 The cephalic sensory organs of Acteon tornatilis (Linnaeus, 1758) (Gastropoda Opisthobranchia) — cellular innervation patterns as a tool for homologisation Titelbild/Cover illustration: Flabellina affinis (Acolidoidea, Nudibranchia) from the Mediterranean Sea on Exdendrinm (Hydrozoa) | (see contribution of AFFELD et al., pp. 181-190) | Bonner zoologische Beiträge Editor-in-Chief Michael Schmitt / Fabian Herder Editors Renate van den Elzen Index Bernhard A. Huber Gustav Peters B d 5 6 Bradley Sinclair an Dieter Stüning | | | | Das Gupta, Sujit H.; MAZUMDAR, Abhijit & CHAUDHURI, Prasanta K.: 43-48 Biting Midges of the Genus Pa/pomyia Meigen (Diptera: Ceratopogonidae) in India Denys, Christiane; Missoup, Alain Didier; TCHIENGUE, Barthelemy; ACHOUNDONG, Gaston; 159-173 EKOBO, Atanga; BILONG BILONG, Charles Felix; LEMBE, Dieudonné Massoma & NICOLAS, Violaine: Altitudinal distribution of and anthropogenic influence on small mammal assemblages on Mount Kupe, SW Cameroon DENZER, Wolfgang & MANTHEY, Ulrich: 255-258 Remarks on the type specimen of Gonocephalus mjobergi Smith, 1925 (Sauria: Agamidae) DEUSCHLE, Jürgen & GLÜCK, Erich: 7-16 Colonisation and Steadyness of Carabid Beetles in Orchards DIETERLEN, Fritz: 185-200 Climbing mice of the genus Dendromns (Nesomyidae, Dendromurinae) in Sudan and Ethiopia, with the description of a new species ELZEN, Renate VAN DEN & NEMESCHKAL, Hans L.: 25-35 The Impact of Body Mass on Morphological Integration in Avian Skeletons (Aves, Fringillidae; Carduelinae, Fringillinae) GAEDIKE, Reinhard: 101-106 Some New and Interesting “Microlepidoptera “ from the Collection of the Zoologisches Forschungsmuseum Alexander Koenig (ZFMK), Bonn (Lepidoptera: Tineidae, Epermeniidae, Acrolepiidae, Douglasiidae) GOODMAN, Steven M.; RAHERIARISENA, Martin & JANSA, Sharon A.: 133-149 A new species of Eliuras Milne Edwards, 1885 (Rodentia: Nesomyinae) from the Reserve Spéciale d’Ankarana, northern Madagascar HALLERMANN, Jakob: 279-284 A new species of Bronchocela (Squamata: Agamidae) from Nicobar Island HUTTERER, Rainer « MONTERMANN, Christian: 201-208 A large new species of Sylvisorex (Mammalia: Soricidae) from Nigeria and the first record of Sy/visorex ollula from the country HUTTERER, Rainer; RIEGERT, Jan & SEDLACEK, Ondtej: 151-157 A tiny new species of Sylvisorex (Mammalia: Soricidae) from the Bamenda Highlands, Cameroon IKARUNARATHNA, D. M. S. Suranjan; AMARASINGHE, A. A. Thasun & STÖCKLI, Edi: 229-238 Taxonomic and biological study on Ca/otes ceylonensis Müller, 1887 (Reptilia: Agamidae) of Sri Lanka KERBIS PETERHANS, Julian; STANLEY, William T.; HUTTERER, Rainer; DEMOS, Terrence C. & AGWANDA, Bernard: 175-183 A new species of Surdisorex Thomas, 1906 (Mammalia, Soricidae) from western Kenya KOCH, André; ArıDA, Evy; RIYANTO, Awal & BOHME, Wolfgang: 107-129 Islands Between the Realms: A Revised Checklist of the Herpetofauna of the Talaud Archipelago, Indonesia, with a Discussion About its Biogeographic Affinities IKÖHLER, G; ROTH, S. & REINHARDT, K.: 17-24 Ten Instars in the Leprous Grasshoppers, Phywatheus leprosus (Fabricius, 1793) (Caelifera: Pyreomorphidae): Maximum Number Recorded in the Acrioidea LEACHE, Adam D.; CHONG, Rebecca A.; PAPENFUSS, Theodore J.; WAGNER, Philipp; BOHME, Wolfgang; 273-278 SCHMITZ, Andreas; RÖDEL, Mark-Oliver; LEBRETON, Matthew; INEICH, Ivan; CHIRIO, Laurent; BAUER, Aaron; ENIANG, Edem A. & BAHA EL Din, Sherif: Phylogeny of the genus Agama based on mitochondrial DNA sequence data Moravec, Jiri & BOHME, Wolfgang: 49-54 Second Find of the Recently Discovered Amazonian Giant Peccary, Pecary maximus (Mammalia: Tayassuidae) van Roosmalen et al., 2007: First Record from Bolivia SANDERA, Martin & STAROSTOVA, Zuzana: 225-228 A record of Asian agama of the genus Ca/otes Cuvier, 1817 (Squamata: Agamidae) in Kenya SCHMITT, Michael: 3-5 In memoriam Günther Osche, 7.8.1926-2.2.2009 TALAVERA, José A.; PÉREZ, Dolores I: 37-41 Occurrence of the Genus Microscolex (Oligochaeta, Acanthodtilidae) at Western Canary Islands WAGNER, Philipp; BARE], Michael FE & SCHMITZ, Andreas: 285-297 Studies on African Agama VII. A new species of the Agama agama- group (Linnaeus, 1758) (Sauria: Agamidae) from Cameroon & Gabon, with comments on Agama mehehi Tornier, 1902 WAGNER, Philipp; INEICH, Ivan; LEACHE, Adam D.; WiLMs, Thomas M.; TRAPE, Sébastien; 239-253 BÖHME, Wolfgang & SCHMITZ, Andreas: Studies on African Agama V1. Taxonomic status of the West African Agama (Sauria: Agamidae) with prominent tail crests: Agama bonlengeri Lataste 1886, Agama insularis Chabanaud, 1918 and Agama cristata Mocquard, 1905 WAGNER, Philipp; Wırms, Thomas M.; BAUER, Aaron & BÖHME, Wolfgang: 215-223 Studies on African Agama V. On the origin of Lacerta agama Linnaeus, 1758 (Squamata: Agamidae) WILMS, Thomas M.; SCHMITZ, Andreas; WAGNER, Philipp; LUTZMANN, Nicola & BÖHME, Wolfgang: 55-99 On the Phylogeny and Taxanomy of the Genus Uromastyx Merrem, 1820 (Reptilia: Squamata: Agamidae: Uromastycinae): Resurrection of the Genus Saara Gray, 1845 WILMS, Thomas M.; WAGNER, Philipp; SHOBRAK, Mohammed & BOHME, Wolfgang: 259-272 Activity profiles, habitat selection and seasonality of body weight in a population of Arabian Spiny-tailed Lizards (Uromastyx aegyptia microlepis Blanford, 1875; Sauria: Agamidae) in Saudi Arabia Buchbesprechungen / Book Reviews tw No) GRIMMBERGER, Eckhard & RUDLOFF, Klaus, unter Mitarbeit von KERN, Christian (2009): Atlas der Sáugetiere Europas, Nordafrikas und Vorderasiens (R. HUTTERER) KRAFT, R. (2008): 6 Mause und Spitzmáuse in Bayern: Verbreitung, Lebensraum, Bestandssituation (R. HUTTERER) MANTHEY, Ulrich (2008): 305 Terralog Vol. 7a — Agamid Lizards of Southern Asia, Draconinae 1 (P. WAGNER) TEMPLE, Helen J. & CUTTELOD, Annabelle (compilors) (2009): 210 The Status and Distribution of Mediterranean Mammals (R. HUTTERER) Publication dates: nos 1/2 31.03.2009, no. 3 30.09.2009, no. 4 30.11.2009 Author Index ACHOUNDONG, Gaston 159 AGWANDA, Bernard 175 AMARASINGHE, A. A. Thasun 229 ARIDA, Evy 107 BAHA EL DIN, Sherif 273 BAREJ, Michael E. 285 BAUER, Aaron 215, 273 BILONG BILONG, Charles Felix 159 BÖHME, Wolfgang 49, 55, 107, 215, 239, 259, 273 CHAUDHURI, Prasanta K. 43 CHIRIO, Laurent 273 CHONG, Rebecca A. 273 Das GUPTA, Sujit H. 43 Demos, Terrence C. 175 Denys, Christiane 159 DENZER, Wolfgang 255 DEUSCHLE, Jurgen 7 DIETERLEN, Fritz 185 EKOBO, Atanga 159 ELZEN, Renate VAN DEN 25 ENIANG, Edem A. 273 GAEDIKE, Reinhard 101 GLUCK, Erich 7 GOODMAN, Steven M. 133 HALLERMANN, Jakob 279 HUTTERER, Rainer 151, 175, 201 INEICH, Ivan 239, 273 JANSA, Sharon A. 133 IKARUNARATHNA, D. M. S. Suranjan 229 KERBIS PETERHANS, Julian 175 KOCH, André 107 KÖHLER, G. 17 LEACHE, Adam D. 239, 273 LEBRETON, Matthew 273 LEMBE, Dieudonné Massoma 159 LUTZMANN, Nicola 55 MANTHEY, Ulrich 255 MAZUMDAR, Abhijit 43 Missoup, Alain Didier 159 MONTERMANN, Christian 201 Moravec, Jifi 49 NEMESCHKAL, Hans L. 25 NICOLAS, Violaine 159 PAPENFUSS, Theodore J. 273 PEREZ, Dolores I. 37 RAHERIARISENA, Martin 133 REINHARDT, K. 17 RIEGERT, Jan 151 RIYANTO, Awal 107 RODEL, Mark-Oliver 273 ROTH, S. 17 SANDERA, Martin 225 SCHMITT, Michael 3 SCHMITZ, Andreas 55, 239, 273, 285 SEDLACEK, Ondfej 151 SHOBRAK, Mohammed 259 STANLEY, William T. 175 STAROSTOVÁ, Zuzana 225 STÓCKLI, Edi 229 TALAVERA, José A. 37 TCHIENGUE, Barthelemy 159 TRAPE, Sébastien 239 WAGNER, Philipp 55, 215 WILMS, Thomas M. 55, 2 av Herausgegeben vom Bonner Euleischen Forschungsmuseum z00logische |“... Beiträge ~ Heft 1/2 2009 (2007 ri! 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David Sissom, Dept. of Life, Earth and Environmental Sciences, W. Texas A. & M. University, WTAMU Box 60808, Canyon, Texas 79016, USA; E-Mail: dsissom@wtamu.edu. en | Dr. Miguel VENCES, Technische Universität Carolo-Wilhelmi- na, Zool. Inst., Abt. Evolutionsbiol., Mendelssohnstr. 4, D-38106 Braunschweig, Tel.: +49 531-391 3231, Fax: +49 531-391 3222; E-Mail: m.vences@tu-braunschweig.de. Prof. Dr. Heike WAGELE, Rheinische Friedrich-Wilhelms-Uni- versitat, Institut fiir Evolutionsbiologie und Okologie, D-53121 Bonn, Tel.: +49 228 73 5159, Fax: +49 234-322 4114; E-Mail: hwaegele@evolution.uni-bonn.de. Dr. Erich WEBER, Eberhard-Karls-Universitát, Zoologische Schausammlung, Sigwartstr. 3, D-72076 Túbingen, Germany; E-Mail: erich.weber@uni-tuebingen.de. Editorial Das vorliegende Doppelheft erscheint mit großer Verspá- tung. Eine erste Verzögerung war durch einen finanziel- len Engpass verursacht worden. In der Folge ließ der Ma- nuskripteingang so stark nach, dass wir keine Hefte fül- len konnten. Dies ist auch der Grund für die unvertretbar lange Wartezeit, die für einige Autoren entstanden ist. Wir bedauern dies sehr und entschuldigen uns dafür. Als Ausweg aus der Krise haben wir folgendes beschlos- sen: (1) Wir werden mehrere Themenhefte auflegen. Als erstes wird unter der Herausgeberschaft von R. Hutterer ei- ne Ausgabe mit lauter Art-Neubeschreibungen erschei- nen. Ein zweites Themenheft wird die Verhandlungen eines Symposiums über Agamen enthalten, herausge- geben von P. Wagner. Weitere Themenhefte sind in Vorbereitung, und wir laden ausdrücklich Kolleginnen und Kollegen ein, ein solches Heft als Gastherausge- ber zu betreuen. (2) Vom jetzigen Heft an werden Abbildungen in Farbe für die Autorinnen und Autoren kostenfrei publiziert. (3) Sämtliche in den BzB erschienene Artikel — von Band l an — werden nach und nach digitalisiert und online gestellt. Sie sind unter www.bzb.zfmk.de von der web- site des ZFMK abrufbar. (4) Autorinnen und Autoren dürfen die PDF-Dateien ih- rer Beiträge legal im Internet verfügbar machen (so- lange das Original-Layout nicht verändert wird). (5) Von Band 57 an werden zum Druck angenommene Beiträge unverzüglich elektronisch veröffentlicht. Mindestens solange die Internationalen Regeln für zoologische Nomenklatur eine Publikation von wissen- schaftlichen Namen und nomenklatorischen Handlun- gen „unter Verwendung eines Verfahrens, welches zahlreiche identische und dauerhafte Exemplare ge- währleistet“ (Artikel 8.1.3.) vorschreibt, werden wir auch eine auf Papier gedruckte Version produzieren. Wir hoffen, auf diese Weise eine höhere Zahl guter Ma- nuskripte zu erhalten und die Bonner zoologischen Bei- träge zu einem attraktiven Publikationsort zu machen. Michael SCHMITT, Schriftleiter The present double issue have been greatly delayed. An initial delay was caused by a financial bottleneck, which was later confounded by the submission of fewer and few- er manuscripts, and consequently we could not complete an ıssue. This ıs apparently also the reason for the drama- tic setback of contributions from some authors. We deeply regret this inconvenience and apologise for it. As a possible loophole we decided the following: (1) We shall publish several Special Issues. First, R. Hut- terer will act as guest editor for an upcoming issue con- taining exclusively numerous species descriptions. A second Special Issue will be devoted to the proceed- ings of a symposium on Agamidae, edited by P. Wag- ner. Additional Special Issues are in preparation, and we explicitly invite interested colleagues to publish such issues as guest editors. (2) From this issue onwards, all colour illustrations will be published free of charge. (3) All articles ever published in BzB will — step by step — be put online. They can be downloaded from www.bzb.zfmk.de. (4) Authors are entitled to distribute their contributions as PDFs freely over the internet, as long as they do not alter the original layout. (5) From Volume 57 onwards, contributions accepted for publication will immediately be published electroni- cally. At least as long as the International Code of Zoo- logical Nomenclature requires publication of new names and nomenclatural acts by means of “a method that assures numerous identical and durable copies” we shall continue to produce a version printed on paper. With this anticipated increase in exposure and turn around time resulting from the above upgrades, we hope to re- ceive a higher number of high quality manuscripts and to make the Bonner zoologische Beitráge a more attractive publication option. Michael SCHMITT, editor-in-chief Bonner zoologische Beitráge | Band 56 co | Heft 1/2 Seiten 3—5 Bonn, Marz 2009 In memoriam Günther Osche 7.8.1926—2.2.2009 Nach einer kurzen, aber schweren Krebserkrankung starb am Montag, 2. Februar 2009, Prof.Dr.Dr.h.c. Giin- ther Osche in Freiburg im Breisgau in seiner Wohnung im Kreis seiner Familie. Er wurde am 7. August 1926 als Sohn eines Bankangestellten in Haardt in der Nahe von Neustadt an der Weinstraße geboren. Seine Eltern zogen aber schon ein Jahr nach seiner Geburt nach Nürnberg, wo er aufwuchs. Dieser Stadt blieb er bis zu seinem Tod treu. Fig. 1. (Photo: M. Schmitt). Prof.Dr.Dr.h.c. Günther Osche, September 30, 2006 Schon als Kind und Jugendlicher war Gúnther Osche be- geistert für die Natur, im besonderen für die Tiere, und ganz besonders für die Vögel. Mit 16 Jahren musste er Flakhelfer werden, mit siebzehn wurde er, nach einem No- tabitur, zum Kriegsdienst eingezogen. Im November 1944, er war gerade einmal achtzehn Jahre alt, wurde er schwer an beiden Beinen verwundet, wovon eine lebenslange Gehbehinderung blieb. Günther Osche studierte vom Wintersemester 1946 an Biologie an der Universität Erlangen und wurde dort mit einer unter der Betreuung von Prof.Dr. Hans-Jürgen Stam- mer angefertigten Dissertation über die Systematik, Phy- logenie und Ökologie der Gattung Rhabditis (Nematoda) am 10. Juni 1951 promoviert. Im selben Jahr heirateten er und Elisabeth Riedel. Nach Assistenten- und Dozentenzeit in Erlangen wurde er 1967 auf einen Lehrstuhl für Zoologie an die Univer- sıtät Freiburg im Breisgau berufen. Bis zu seinem vorzei- tigen Rückzug in den Ruhestand 1988 las er dort 22 mal die umfang- und gehaltreiche Vorlesung über spezielle Zoologie der Wirbellosen. Aus der Reihe seiner 84 wissenschaftlichen Publikationen ragen besonders heraus seine Habilitationsschrift über die systematische Stellung und Phylogenie der Pentastomida (1963) und der Handbuchbeitrag über die „Grundzüge der allgemeinen Phylogenetik“ (1966a). Er selbst hielt seine „Beiträge zur Morphologie, Ökologie und Phylogenie der Ascarıdoidea (Nematoda)“ (1958) für dıe am besten ge- lungene (siehe SCHMITT 1996). Fraglos haben seine po- pulären Bücher über „Die Welt der Parasiten“ (1966b), „Evolution“ (1972) und „Ökologie“ (1973) die größte Breitenwirkung entfaltet. In den letzten Jahren seiner wissenschaftlichen Betätigung faszinierte ihn zunehmend die Wechselbeziehung zwi- schen Blüten und ihren Bestäubern, ganz besonders die evolutive Entstehung von Signalen und Täuschungen auf Seiten der Blüten (z. B. 1983). Günther Osche wurde 2001 die Ehrendoktorwürde der Rheinischen Friedrich-Wilhelms-Universität Bonn verlie- hen, 2006 wurde er zum Ehrenmitglied der Deutschen Zoologischen Gesellschaft ernannt, deren Präsident er 1973 und 1974 war (s. SAUER 2007). Wer ihn erlebt hat, wei, dass noch viel eindrucksvoller als der Autor der Redner Giinther Osche war (Fig. 1). In Vorlesungen, Vortragen und persónlichen Gesprachen hat er seine eigene Begeisterung für die Vielfalt der Organis- men und ihrer Leistungen immer wieder auf seine Zuhö- rer übertragen. Auch wurde er nicht müde seine Bewun- derung für August Johann Rösel von Rosenhof, Johann Wolfgang von Goethe und Charles Darwin farbenreich auszudrücken. Er vertrat pointierte Ansichten zu Politik, Erziehung und Ästhetik, an denen sich ohne weiteres leb- hafte Diskussionen entzünden konnten. Dabei wurde er nie verletzend, es lag ihm stets an respektvollem Ausgleich. Für diejenigen, die ihm persönlich nahe standen, war er 4 In memoriam Gúnther Osche, 7.8.1926-2.2.2009 eine wohlwollende, Vertrauen und Autorität ausstrahlen- de Vaterfigur. Während der 70er und 80er Jahre des 20. Jahrhunderts prägte er die deutschsprachige Evolutions- biologie durch seine publizierten Werke ebenso wie durch sein persönliches Auftreten. Eine kraftvolle Stimme ist verstummt. Wer sie je gehört hat, wird sie immer in sich hören. Danksagung. Ich danke Gabriele Uhl (Bonn) für die hilfreiche Durchsicht des Manuskripts. After a short but severe bout of cancer, Prof.Dr.Dr.h.c. Günther Osche passed away on 294 of February, 2009, in Freiburg im Breisgau, Germany, in the circle of his fam- ily. He was born on 7 August 1926, the son of a bank clerk in Haardt near Neustadt an der Weinstrasse, a small town in southwestern Germany. A year after he was born, his parents moved to Nürnberg, where he grew up. He re- mained true to this town for all his life. Already as a child and youngster, Günther Osche was an ardent nature lover. All animals interested him, especial- ly birds. At the age of 16 he served in a flak battery, and when he was 17 he was, after an abridged final school ex- amination (“Notabitur”), called into the regular army. In November 1944 he was severely wounded in both legs which remained a lifelong impairment. Gúnther Osche studied biology at the University of Er- langen from winter semester, 1946, receiving his doctor- al degree in 1951. His dissertation, supervised by Profes- sor Hans-Júrgen Stammer, treated the systematics, phy- logeny, and ecology of the genus Rhabditis (Nematoda). The same year, he and Elisabeth Riedel married. After his degree, he served as an assistant, earned his habilitation in 1963, and obtained a position as a university docent in Erlangen. In 1967 Günther Osche was appointed full pro- fessor of zoology at the Albert Ludwigs-Universitát Freiburg im Breisgau. Until his voluntary retirement in 1988 he presented his famous, extensive and rich lecture series on special zoology of the invertebrates for a total of 22 times. Of his 84 scientific publications, his habilitation thesis on systematic position and phylogeny of the Pentastomida (1963) and his handbook chapter on basics of general phy- logenetics (1966a) are especially outstanding. He regar- ded his contribution on morphology, ecology and phyloge- ny of the Ascaridoidea (Nematoda) (1958) as his best pa- per (see SCHMITT 1996). Unquestionably his popular books on the world of parasites (1966b), evolution (1972), and ecology (1973) reached the broadest readership. During the last years of his professional activity he turned increasingly fascinated by the interaction between flow- ers and their pollinating insects, especially the evolution of signals and deceptions by plants (e. g. 1983). In 2001, he was awarded the degree of an honorary doc- tor by the Rheinische Friedrich Wilhelms-Universitát, Bonn, and in 2006 he was elected as an honorary mem- ber of the German Zoological Society, of which he was president in 1973 and 1974 (see SAUER 2007). Whoever met him in lectures and discussions knows that even more impressive than Osche the author was Osche the speaker (Fig. 1). Through lectures, talks and perso- nal conversation, he transferred his own enthusiasm of the multifariousness of organisms and their attributes to his listeners. Tirelessly he expressed in vivid colours his ad- miration of August Johann Rósel von Rosenhof, Johann Wolfgang von Goethe, and Charles Darwin. He advanced definite views on politics, education, and aesthetics, which easily led to lively discussions. However, he never offen- ded, but always aimed at a fair and respectful balance. To all who were close to him, he was a benevolent fatherly mentor who gave off an air of trustfulness and natural au- thority. During the seventies and the eighties of the 20th century, he guided evolutionary biology in the German speaking community by his publications, but likewise by his personal performance. A powerful voice has fallen silent. Whoever has been for- tunate to hear it, will hear it forever. Acknowledgement. | thank Walter J. Bock (New York) for care- fully improving the English manuscript. REFERENCES Oscue, G. 1958. Beiträge zur Morphologie, Ökologie und Phy- logenie der Ascaridoidea (Nematoda). Parallelen in der Evo- lution von Parasit und Wirt. Zeitschrift für Parasitenkunde 18: 479-572. OscHE, G.1963. Die systematische Stellung und Phylogenie der Pentastomida — embryologische und vergleichend-anatomi- sche Studien an Reighardia sternae. Zeitschrift für Morpho- logie und Ökologie der Tiere 52: 487-596. OscHE, G. 1966. Grundzüge der allgemeinen Phylogenetik. Pp. 817-906 in: Handbuch der Biologie 3/2. Akademische Ver- lagsgesellschaft Athenaion, Frankfurt am Main. ÖSCHE, G. 1966. Die Welt der Parasiten (Verstándliche Wissen- schaft 87). VIII+160 pp., Springer, Berlin etc. Bonner zoologische Beiträge 56 (2007) 5 OscHE, G. 1972. Evolution. Grundlagen, Erkenntnisse, Entwick- lungen der Abstammungslehre (Studio visuell). 116 pp., Her- der, Freiburg. OSCHE, G. 1973. Ökologie. Grundlagen, Erkenntnisse, Entwick- lungen der Umweltforschung (Studio visuell). 142 pp., Her- der, Freiburg. OscHE, G. 1983. Optische Signale in der Coevolution von Pflan- ze und Tier. Berichte der Deutschen Botanischen Gesellschaft 96: 1-27. SAUER, K.-P. 2007. Laudatio zur Verleihung der Ehrenmitglied- schaft in der Deutschen Zoologischen Gesellschaft an Dr.Dr.h.c.mult. Günther Osche, Professor Emeritus und vor- mals Direktor am Zoologischen Institut der Universität Frei- burg. Pp. 13-16 in: Zoologie 2007 — Mitteilungen der Deut- schen Zoologischen Gesellschaft. Basilisken-Presse: Marburg an der Lahn. SCHMITT, M. 1996. Günther Osche — a man of the spoken word. Zoologischer Anzeiger 235: 1-9. Michael SCHMITT, Zoologisches Forschungsmuseum Alexander Koenig, Bonn, Germany (m.schmit@uni- bonn.de). 6 Buchbesprechung KRAFT, R. (2008). Mäuse und Spitzmáuse in Bayern: Verbreitung, Lebensraum, Bestandssituation. 111 S., 63 Farbfotos, 51 Karten, 25 Diagramme, 4 Tabellen. Eugen Ulmer, Stuttgart. ISBN 978-3-8001-5609-2. Preis 39€, Dieser schmale Band ergänzt das 2004 im gleichen Ver- lag erschienene Buch „Fledermäuse in Bayern“ (von A. MESCHEDE und B.-U. RUDOLPH). Anders als bei Fleder- mäusen konnte sich der Autor des vorliegenden Buches weniger auf die Beobachtungen ehrenamtlicher Naturschützer stützen (für Mäuse und Spitzmäuse gibt es keine Lobby), sondern erarbeitete die Daten für die Ver- breitungskarten selbst durch langwierige Analyse von Eu- lengewöllen, die zahlreiche Ornithologen in 332 bay- erischen Ortschaften gesammelt hatten. Insgesamt wurden über 40.000 Kleinsäugerbelege durchgesehen und be- stimmt. Ergänzend wurde die im Bayerischen Landesamt für Umwelt vorliegenden Daten und Belege aus Muse- umssammlungen herangezogen. Viel Zeit und Mühe ver- wendete der Autor auch auf die Anfertigung von Farbfo- tos lebender Kleinsäuger. Das Resultat ist eine sorgfältig kompilierte Übersicht der für Bayern vorliegenden Ken- ntnisse von 27 Kleinsäugerarten. Die beigefügten Lebend- fotos wird von allem derjenige zu schätzen wissen, der selbst einmal versucht hat, eine Spitzmaus oder eine Wald- maus zu fotografieren. Schade ist, dass nicht alle Arten von Nagern (Biber, Bisam und Nutria fehlen) und Insek- tenfressern (Igel, Maulwurf) behandelt wurden. Die hier vorgelegten Daten ergänzen unsere Kenntnisse über die Kleinsäugetiere in Deutschland und sind relevant für den Naturschutz in Bayern. Rainer HUTTERER Zoologisches Forschungsmuseum Alexander Koenig, Bonn Seiten 7-16 Bonn, Márz 2009 Bonner zoologische Beitráge Band 56 (2007) | Heft 1/2 Colonisation and Steadyness of Carabid Beetles in Orchards Jürgen DEUSCHLE & Erich GLUCK Abstract. From April 1995 to November 1997 investigations were carried out in the nature reserve “Limburg” (48.36 N/9.23E): data were collected on the type and frequency of grassland use and data on the carabid fauna. 17 sample plots were selected. Part of the plots had 25 years of unchanged management regimes: three-cutting meadows (3), two-cutting meadows (3), mulched meadows (4), abandoned meadows (3), a horse pasture, a sheep pasture with rotational grazing, a continuously grazed sheep pasture, and a sheep pasture abandoned in 1994. 5229 beetles representing 68 carabid species were caught in pitfall traps during the three years of investigation on the 17 study plots (Tab. 2). On nearly all studied areas/plots changed the relative frequency of dominant species. 3-cutting mead- ows frequently show a one-sided activity dominance structure. Within the yearly spectrum of activity dominance on all areas it is obvious that species occured and are leaving continously. Rare species in the area (steadyness group I: present on 1-3 plots) reached percentages between 10 % and 42 %, the cor- responding values for steadyness group II (species present on 4-6 plots) are 0 % up to 67 % und steadyness group III (present on 7-10 plots) 20 up to 64 % (Fig. 2, Tab. 3). The management of the grass vegetation in orchards and the in- fluence on the carabid community is briefly discussed. Keywords. Ground beetle, grassland management, species colonisation, species steadyness. 1. INTRODUCTION Vegetation and fauna of extensively managed orchards are mainly determined by the site and its maintenance, the type of grassland management and its land use intensity (DEUSCHLE et al. 2002, GLUCK et al. 2004). Until now animal communities of extensively managed orchards have not been analysed based on the direct com- parison of different forms of grassland management There are few investigations dealing with the influence of man- agement systems on arthropod communities in extensive- ly managed grassland, whereas the effects of grazing have been documented more frequently (HANSSEN & HINGST 1995, MAELFAIT et al. 1988, RUSHTON et al. 1989, SCHNIT- TER 1994, DEUSCHLE & GLUCK 2001, GLUCK & DEUSCHLE 2003). Carabids showed a higher activity on cut and mulched meadows than on plots in state of succession during aban- donment trials (SOUTHWOOD & VAN EMDEN 1967). But the abandonment of two meadows, one fertilised and the oth- er one unfertilised, lead to an increase of the activity den- sity during the course of succession (SCHNITTER 1994). Low food resources — determined by differences in a de- crease of the population — caused a lower activity densi- ty in woodlands (GUILLEMAIN et al. 1997). Beside describing represent field studies in the last years was tried to explain and scale the differential character of carabid communities. Abiotic and management specific characters and gradients have been worked out. (BAGUETTE 1993, BAUER 1989, BUTTERFIELD & COULSON 1983, DENNIS ef al. 1997, DUFRENE & LEGENDRE 1997, EYRE ef al. 1990, HuHTA 1979, Lurr et al. 1992, LUFF 1996, Mc FERRAN et al. 1994, Morris & RISPIN 1987, PREISZNER 1996, RODE 1993, VOWINKEL 1996, GLÜCK & DEUSCHLE 2003). The aim of this paper is to ask wether carabids colonize relatively homogenous orchards constantly in nearly the same species and individual densities or in a more stochas- tic manner, or is there a concentration on some plots/ habi- tats of specieal species. The species therefore were divid- ed into three classes of steadyness (see 2.2). The constan- cy in time of the composition of carabid community is in- spected and how it is linked up management specific at the background of different management regimes, differ- ent area, the distribution with altitude and the shading of the plots. Analysis of similarity of the carabid communi- ties and the activity dominances on the plots show a pat- tern in time and space of the species compsition and dom- inance idendities of different management regimes. 8 Jürgen DEUSCHLE & Erich GLUCK: Colonisation and Steadyness of Carabid Beetles in Orchards 2. METHODS 2.1. Census and registration of land use From the beginning of April until the middle of October 1997 the following parameters were recorded weekly or every two weeks in the study area orchards in the nature reserve “Limburg” (48.38N/9.23E), south west Germany: type of use, time of mowing, “mowing device”, “where- abouts” of mowed grass, and number of grazing animals. The amount of ground shaded by trees under perpendicu- lar solar radiation was estimated and assigned to six cat- egories. All areas, which are mowed completely on a regular ba- sis every year and where the cut grass is removed, will be called “typical meadows”. On “mulched meadows” the cut grass remains regardless of the mowing device and mowing frequency. “Continuous grazing” defines the management type of pastures, where the grazing animals can be found permanently on the same area, while “rota- tional grazing” 1s the type, where a particular area is be- ing grazed for a period of only a few days, but several times a year. Combinations of the three types of manage- ment can also be found in the extensively managed or- chards of the nature reserve area. Landuse 1997 Mixed landuse SJ for grazing and mowing (sheep) BEER Mulching and pasturing === mukching and mown crop with clear up "77, mwaiching, mown crop and pasturing Succession recently abandoned meadow old abandoned meadow MH Woodland 50 0 50 100 Meter Fig. 1. Land use intensity of grassland in the study area during the vegetation period of 1997 and the sample plots. Bonner zoologische Beitráge 56 (2007) 9 2.2. Selection of sample plots and Population census In 1995 the selection of sample plots took into consider- ation areas which had been under the same management system for more than 25 years (see Fig.1, Tab. 1). Six pitfall traps were placed in a single line 10 m apart as a transect through the centre of each sampling plot. Eth- ylene glycol (50 %) was used as preservative solution, with detergent added to reduce surface tension. A cover made of Perspex (120 by 120 mm) was installed 30-50 mm above ground level. In-1995 (5. April to 1. Novem- ber) traps were controlled and emptied regularly weekly. In 1996 (5. May to 4. November) and 1997 (8. April to 4. November) the control interval was two weeks (DEUSCHLE & GLUCK 2001). The division in classes of steadyness helps to classify the rareness in the study area, independent from their plot spe- cific frequency, where ever this scarcity 1s only valuable for the individual study plots. The species were divided into three classes of steadyness: steadyness l are all species occuring on 1-3 plots, steadyness II species occuring on 4-6 plots, steadyness III species occuring on 7-9 plots. 2.3. Analysis of data and statistical methods All registered data concerning management practices, veg- etation, soil condition, and carabid population were inte- grated into a database. After examining the necessary con- ditions the following statistical tests were applied: U-test by Mann & Whitney (MWU-test), and X?-test. The rank correlation was calculated according to Spearman and the coefficients were tested on their significance. 3. RESULTS 3.1. Acitivity dominance The activity density of carabids on the various plots is quite different (Tab 2). The yearly activity density on the three fold mowed meadow (3CM1) showed differences in the distribution of the second up to the fifth dominant species. The percentages of these species (Amara aenea, Pseudoophonus rufipes, and Nebria brevicollis) decrease continually within the three year study. In 1995 P. rufipes and N. brevicollis were the third respective fifth dominant species an 1997 both were totally absent. In contrast to this the percentages of Clivina fossor, Poecilus cupreus, and Carabus ullrichi steadily increase throughput the three year period. Only Anisodactylus binotatus in all three years remained as the cominant species despite of its changing perdentages in the area. Six species occured 1995 in a low density, they were absent in 1996 and occured again in low density 1997. In 1996 and 1997 seven species are regis- tered which were not collected in 1995 in the pitfall traps, two of the species were captured only in 1996, and three species out of them in 1997. On the 2-cutting meadow (2CM1) the dominant species changed during the three years of investigation: Harpalus latus, the activity of which decreased countinously is removed by €. ullrichi the percentage of which increased. The next following species hold nearly the same dominance ranks with the years. Three species are only in 1995 and 1997, seven species only in 1995 registered. Eight more species are on- ly present in 1996 and 1997, two out of them only in 1997. On the horse pasture (HP) all years the dominance of species changes from C. fossor to Pterostichus vernalis on to Pterostichus melanarius, the percentage of the lat- ter increased from 1% in 1995 to 8.5% up to 29.1% in 1997. The percentages of the other dominant species de- crease with the years. A. plebeja is registered only in 1995; Amara familiaris and €. ullrichi restricted their occurance in 1995 and 1997 also on this plot. In contrast in 1996 new species occured in the community, six more species in 1997 by most of them only one individual was collected. On the anbandoned sheep pasture (SPA) P. melanarius re- mains as the most dominant speacies and reached eudom- inant status, with 39,4 % in 1995 and 58,4 % in 1997. N. brevicollis disappeared on this plot in 1997. This species ranked in 1995 still on the second dominance position. Three species were registered exclusive on this plot in 1995, five only in 1996 and six species only in 1997. On the sheep pasture rotational grazing (SPR) C. ullrichi was the most dominant species througout the three years. Pterostichus ovoideus strongly increased and was collect- ed in the same frequency as C. ullrichi. On this plot four species occured only in 1995 or 1996, five more occured only in 1997. On the mulched meadow (MMI) Leistus ferrugineus re- moved the so far most frequent C. fossor 1997. L. ferrug- ineus occured the first time in this community, reached a percentage of 6,1 % and is ordered in status to the sub- dominant carabids. The percentages of species belonging to the dominant ones in 1995 changed strongly and are displaced by the so far subdominant species as C. ullrichi. Seven species are trapped only in 1995, ten more species in 1996 and 1997 four out of these only in one of both years. The dominance spectrum is wellbalanced on the mulched meadow (MME2). A. latus reaches all the years’ eudom- inant status. Three species are registered only in 1995, two more in 1996 and no more in 1997. 10 Jürgen DEUSCHLE & Erich GLUck: Colonisation and Steadyness of Carabid Beetles in Orchards Table 1. Morphology and land use of sampled plots. Plot Land use Type of Area Circumference Altitude Shading Sampling land use (Ar) (m) (mNN) (%) period 3CM2 3-cutting meadow Meadow 26 ~ 209 405 20 “97 3CM3 3-cutting meadow Meadow 31 287 425 20 “OF: 3CM1 3-cutting meadow Meadow 16 204 392 20 95,96, 97 2CM3 2-cutting meadow Meadow 40 446 415 80 ‘97 2CMI 2-cutting meadow Meadow 10 218 445 60 95596; 97 2CM2 2-cutting meadow Meadow 15 250 410 60 "97 SPA abandoned sheep Pasture 34 234 405 60 195,96, 97 pasture (continuous grazing), see text SPC Sheep pasture Pasture 15 213 415 80 >97 (continuous grazing) HP Horse pasture Pasture 124 580 400 20 "95,96,'97 SPR Sheep pasture Pasture 11 259 445 20 "95,96, 97 (rotational grazing) MMI Mulched meadow Mulched 12 162 455 20 "95, 96, 97 (4 - 6 cuttings) meadow MMEI Mulched meadow Mulched 47 380 440 100 >97 (3 cuttings) meadow MME2 Mulched meadow Mulched 29 217 420 60 "95,96, 97 (2 — 3 cuttings) meadow MME3 Mulched meadow Mulched 25 206 517 80 "97 with | cutting meadow AMR Recently abandoned Succession 13 153 445 0 95596, 97 meadow (4 years) AMO Old abandoned Succession 13 202 420 100 95,°96, 97 meadow (10 years) WDL Woodland Succession 13 222 520 100 "97 On the recently abandoned meadow (AMR) A. latus dis- placed 1996 the so far most frequent A. binotatus, the lat- ter belonged in 1996 and 1997 just to the group of sub- dominant species. The percentages the persuiting dominant species on the plot P. ovoideus, C. fossor and Bembidion obtusum changed strongly during the years. Four species are reg- istered only 1995, six more only in 1996 and 1997. In the dominance spectrum of the old abandoned mead- ow (AMO) set apart from the percentages of the most fre- quent and dominant species from year to year consider- ably and lead to a yearly change in the ranking of the com- munity, during the course1996 and 1997 only few species added new into the community. Spectrum and the rankıng of dominant and subdominant of species change strongly within the study plots and years. Regarding comparative management forms there are great differences regarding the main species in the spectrum (Tab. 2). The percentage of the most frequent species compared to the number of all species together did not show any con- nection with the number of species or individuals nor the management of the plots. The maximal and minimal val- ues are given on three year investigated plots on the aban- doned sheep pasture (SPA, 58,4 %) in 1997 and on the horse pasture (HP, 13,9 %) 1996). On the one year investigated plots owns the 3-cutting meadow (3CM3) with 16,4 % the lowest value , the like- Bonner zoologische Beitráge 56 (2007) 11 > 3 8 o5848683 (%) MME2 (%) AMR ©1995 26.3 21. 52.9 m1995 29.4 17.6 50.0 m199 13.3 66.7 20.0 2199 17.6 47.1 35.3 =1997 18.8 18.8 62.5 “1997 37.5 0.0 56.3 81995 333 16.7 50.0 81996 28.6 42.9 ! 28.6 21997. 18.2 18.2 63.6 Fig. 2. Steadyness of carabid species from 1995-1997 on plots of different management 3CM1, 2CM1, SPA, HP, MME2 and MMI (left) / MME2, AMR and AMO (right) (species percentages). wise 3-cutting meadow (3CM3) with 49,4 the highest val- ue. Eleven different species built up the most frequent carabid on the three year investigated plots; H. latus was for seven times the most frequent on the nine investigat- ed plots. Only on the areas 3CM1, MME2 and SPA in all three years the same species reach the highest frequency (4. binotatus, H. latus and C. ullrichi). From the other species are C. ullrichi, C. fossor, A. binotatus, P. vernalis und P ovoideus high steady and occur on nearly all oth- er plots. P. melanarius und A. parallelepipedus are regis- tered only on a few investigated plots. The arrangement of the most frequent and second most frequent species does not show any affinity to compatible or to near by standing management forms. 3.2. Steadyness of species on the plots The determined steadyness allows on the three year stud- ied areas/plots a differentiation between three groups. The first group consists of species which colonize regularily two-third of the study plots (Steadyness III, Fig. 2). This group comprises 1995: 11 (24 %) of altogether 46 species, 1996: 6 (15 %) from 39 species and 1997: 10 from 38 (26 %) species. The second group (steadyness II) comprises 1995: 8 (17 %), 1996: 14 (36 %) und 1997: 7 (18 %) of the species. The third group comprises the large part of all detectable species (Steadyness I). 27 (59 %) of the species 1995 contained in the pitfall traps were only pres- ent on one to three study areas/plots. In 1996 19 (49 %) species belonged to this group, 1997: 21 (55 %). With this the percentages of the classes of steadyness in 1995 und 1997 are comparable. 1996 evidence of some rare species in the study area/plots are absent (probably dependent on the low activity). In 1995 more frequent species are de- tected on less study plots, so in 1996 the steadyness class II was comparatively high occupied (Fig. 2). The highest proportions of steadyness I were registered on the 3-cutting meadow (3CM1) and on the abandoned sheep pasture (SPA) (39 %), the lowest percentages of this class were in the carabid community on the plot 2CM1 (1997: 10 %, Fig. 2). The percentages on the plots are be- tween 7 % (MME2) and 23 % (2CM2), beside the horse pasture (HP) the values are in 1996 negligible lower com- pared to 1995 oder 1997. Some of this rare species in the study area are able to es- tablish in seperate years or continously on distinct plots larger populations. To this species belong Abax paral- lelepipedus, Amara aenea, Bembidion properans or Pterostichus melanarius. 19 Jürgen DEUSCHLE Erich GLÚCK: Colonisation and Steadyness of Carabid Beetles in Orchards Table 2. Density of species, activity abundance, dominance index, area specific proportions of main and accessory species and the most frequent species each in the perennial sample plots (1995-1997). MME2 Area: SPA HP SPR MMI AMR AMO Number of 1995 23 24 18 22 18 19 19 ily 12 species 1996 17 20 20 20 18 16 15 17 7 1997 21 20 20 24 16 16 16 16 11 Number of 1995 226 144 226 200 76 95 86 84 81 individuals: 1996 64 58 81 130 69 49 4] 82 56 1997 125 75 267 234 87 106 113 60 58 Percentage of 1995 26,1 31,8 27,8 36,4 55,6 42,1 52,6 41,2 66,7 Main species: 1996 58,8 65,0 35,0 45,0 50,0 56,3 66,7 41,2 85,7 (%) 1997 42,9 50,0 25,0 37,5 62,5 62,5 31,3 50,0 1257) Percentage of 1995 73,9 68,2 72,2 63,6 44,4 57,9 47,4 58,8 33,3 Accessory sp.: 1996 41,2 35,0 65,0 55,0 50,0 43,8 33,3 58,3 14,3 (%) 1997 57,1 50 75,0 62,5 37,5 37,5 68,8 50,0 2153 dominance 1995 25,5 31,3 39,4 18,0 31,6 33,7 29,1 16,7 37 index 1996 15,6 22,4 25,9 13,9 18,8 18,4 29,3 24,4 48,2 (%): 1997 31,2 21,3 58,4 29,1 18,4 19,8 28,3 30,0 24,1 1995 Anisod. Harpalus Pt. mela- Clivina Carabus Clivina Harpalus Anisod. A.paralle- Most frequent binotatus latus narius fossor ullrichi fossor latus binotat. lepipedus species: 1996 Anisod. Harpalus Pt. mela- Pt. Carabus Clivina Harpalus Harpalus. Molops binotatus latus narius vernalis ullrichi fossor latus latus elatus 1997 Anisod. Carabus Pt. mela- Pt. mela- C. ullrichi/ L.ferru- Harpalus Harpalus. A.parallelus binotatus ullrichi narius narius Pt. ovoideus gineus latus latus /Harpalus latus The area specific percentages in class II stagger between 0% (AMR 1997) und 50 % (SPR, MMI 1996). The scat- tering on the plots within the 3 year study 1s essentially higher in this class of steadyness I, the values are beween 14 % (HP) und 47 % (AMR). Species of steadyness III colonized all study plots/areas. The highest percentages in this class are on AMO (64 %) und MME2 (63 %) in1997. On the same areas are the low- est percentages in 1996 (29 % bzw. 20 %). The area spe- cific percentages of the community stagger considerably (13-43 %). The unequivocal distribution of the steadynes shows clear, that within the relatively homogeous and enclosed or- chards in the study area exist numerous small-patterned differences which can lead to inhomogenous distributions on neughbouring areas. A smaller part of species colonizes the orchards in changing frequencies nearly overall. The majority of species concentrate abviously only on small- patterned and local distribution centres, Their discontin- ually and recedental occurance in other areas allows the interpretation that ther behaviour is a navigational explo- ratıon. 1996 and 1997 the percentage of rare species (steadyness I) increased with increasing area of the plots, 1997 the cor- relation was statistically significant (Tab. 3). In the three years of investigation the percentage descreased with in- creasing altitude, the correlation in 1997 was statistical- ly significant. The steadyness seems not to be correlated with the param- eters of the plots like management, circumference and shading; the correlation analysis did not show any tenden- cies. Only 1997 the percentage of frequent species (stea- dyness III) increased with decreasing management inten- sitymit, 1996 the percentage decreased statistically signif- Bonner zoologische Beiträge 56 (2007) 13 Table 3. Rank correlation coefficient of species percentages of different steadyness classes of the carabid community with area parameters (n = 9; level of significance: * =p < 0.05, ** =p < 0.01, *** = p < 0.001). Species Management percentages 5 1995 SCI -0.56 SCH 0.12 SC HI 0.38 1996 SCI -0.35 SCH 0.31 SCM -0.28 1997 SCI -0.11 SCH -0.55 Area circumference altitude shading 0.28 -0.01 -0.59 0.33 0.19 0.36 -0.18 0.03 0.13 0.27 0.11 -0.34 0.52 0.32 -0.60 0.22 -0.63 -0.27 0.70* 0.09 -0.32 -0.26 0.13 0.84** 0.81*7% 0.08 -0.76* -0.37 -0.34 0.16 0.12 0.01 SC HI 077 -0.40 -0.43 0.57 0.34 icant with increasing shading. So that seems between the comparatively rich species mowing meadows and pastures and the scarce species areas like abandoned and mulching meadows are no differences in the percentages of rarely or frequent species in the sudy area (Tab. 3). In contrast the percentages of rare species (steadyness I) increase with increasing area 1996 and 1997, 1997 the cor- relation was statistically significant. In all three years the percentage of these species decreased with increasing al- titude of the study plots. 4. DISCUSSION Three cutting meadows are colonized by numerous species, which occur either in a high population density and/or are not present on other areas. The typical speceis on three cutting meadows are predominantly spreeding and euritopic forms, which colonize a broad spectrum of different habitats. Some of this species, as P. melanarius or €. fossor indicate with high population densities inten- sive management forms (DESENDER ef al. 1994, TIETZE 1985, 1987). The species colonizing 3-cutting meadows belong most- ly to steadyness III or I, but seldom to I. They are there- fore either high frequent or very rare. In four of these species the individual density was significantly higher than on other study areas, on which they occured regularily in scarce populations (Tab. 2). These species belong most- ly to steadyness III (A. binotatus, P. cupreus, C. fossor, exception: Pt. melanarius). A. binotatus, P. cupreus and C. fossor colonized all 3-cutting meadows continously and therefore in time and space on all areas of this manage- ment form present. All four species are capable to fly, two species are macropteric and two are alae dimorphic and therefore comparable mobil. However from P. melanarius and C. fossor no completely macropteric individuals were registered. Especially P. melanarius colonized the area 3CM2 and 3CM3 as well as the neighboured pastures in high population densities but was absent on 3CM1 (DE- SENDER ef al. 1994). Obviously in its spreading pattern seems to be content an area dependent gradient, which had not been jet registered: in the sout-western part of the study area the species occured 1997 on nearly all studied plots therby also on pessimissm areas (MME3). In the north- eastern part of the area there are only some lonely indi- viduals captured (1996: SPR, 1997: AMR) and in our opinion optimal areas (3CMI) have not been colonized. Large 3-cutting meadows and pastures are common in the south-eastern part of the study area, wheras they are on the remaining area more rare and smaller and mostly sur- rounded by mulching meadows. (Fig.1). The unability to fly and therefore beeing scarcely mobile in the investigat- ed populations seems potentially more favorable coloniz- ing, but prevents colonizing of secluded mowing mead- ows in other parts of the study area. Therefore the species overall distribution is not only based on suitable manage- ment forms or habitats but also on the attainment of ex- ploring individuals of local populations, which depends on the species immanent mobility. With more typical species of 3-cutting meadows the density of individuals are clear lower, they colonize the plots with other man- agement forms essentially more rare and fall mostly in steadyness class I. (Amara aenea, Bembidion properans, Agonum mülleri, Poecilus versicolor, Carabus granula- tus, Diachromus germanus, Amara montivaga). Except C. granulatus (brachypteric), all individuals of the species are macropteric. They are therfore capable to fly and more mobil in contrast to the differential species of stedyness group III. In contrast to the differentiating species from steadyness group I on the studied 3-cutting meadows they are less continously present in time and space. This dis- 14 Jürgen DEUSCHLE & Erich GLUCK: Colonisation and Steadyness of Carabid Beetles in Orchards continous presence and the lower population density in- dicates a lower plasticity and a higher risk of extinction in local populations of this species, whereas eurytopic species with high plasticity even are able to colonize pes- simal areas. For the latter therefore seems the ability to fly of reduced importance. Some of the differential species on 3-cutting meadows (es- pecially of steadyness group III) colonize pastures more frequently. These are P melanarius and P. vernalis which colonize both management forms in higher populations densities than 2-cutting meadows, mulching or abandoned meadows. They mark the similarity of the carbid commu- nity of both management forms which obviously repre- sent comparable living conditions. Species colonizing on- ly pastures in high densely populations were only pres- ent by Pt. macer und P. rufipes. Their rarely population densities did not allow to mark off statistically signifcance to other management forms. (Fig. 2). N. brevicollis and C. ullrichi have been trapped signifi- cantly more frequent on 2-cutting meadows compared to other management forms. Both species are counted to the more frequent ones (steadyness II / steadyness III) and col- onize all other management forms at least in low densi- ties (Fig. 2). But both species dave been very frequent on the area 2CM3 out of the direct Limburg area, and hint the first time an influence independent from the manage- ment form on the composition of specific carbid commu- nities in orchards. The eurytopic N. brevicollis is a typi- cal species of the woodland edge (THIELE 1964). Their densely population on 2-cutting meadows and scarce ac- tivity on abandoned areas is different from this findings, but a overall high presence on the study plots confirms the morphological relationship from orchards and thin woodlands. Both habitats offer obviously for N. brevicol- lis comparable living conditions (see GRUSCHWITZ 1983). After the scarce specifications, the rarely found species C. ullrichi (RL 3) inhabits a broad spectrum of various habitats with ist centre of distribution in Southern Europe. The statements are from loamy fields, meadows, and ru- ral areas up to thin woodlands (HURKA 1996).This plas- ticity seems to correspond to the high steady presence on all studied plots. Leistus ferrugineus, the only species with its centre of dis- tribution in mulching meadows increased rapidly over the three year study (1995: steadyness I, 1996: II, 1997: III). In contrast euritopic species of abandoned meadows (Carabus auratus, Abax parallelus, Abax paral- lelepipedus) are more often present on meadows and pas- tures. A. parallelus (1995 — 1997: steadyness II) is more common than 4. parallelepipedus (1995, 1997: steadyness I, 1996: II) and reaches moreover a higher population den- sity on mulching meadows. A. parallelus therefore 1s the species with higher plasticity and a lower bonding to cul- tivating or succession phases. So called stenotopic wood- land species however are in extensive orchards restricted on succession areas or border on woodlands (M. elatus, C. coriaceus, C. nemoralis). They indicate thereby a strong change of the carabid community on a complete abandon- ment. Beside the differential species there are obviously some highly steady species (steadyness III), which colonize all studied plots in nearly the same density, whenever it is re- marquable that they also reflect a specific tendency of cul- tivation. H. latus (macropteric) colonizes on less nutrient cutting meadows and pastures in a slightly higher densi- ty than on mulching meadows and abandoned meadows. Its centre of distribution in the grassland remains more likely on ologotrophic areas (TIETZE 1985, Fig. 2). Pt. ovoideus (brachypteric), colonises 2-cutting meadows, mulching meadows und abandoned meadows more fre- quently. The area specific percentages of the steadyness groups I- III are obviously not influenced by the management form of the study plots. On the species rich cutting meadows and pastures especially rare species are not found more frequent compared to low species areas as mulching mea- dows and abandoned meadows. But the area specific per- centages of steadyness group I decrease tendentious with decreasing size and increasing altitude of the plots. For larger plots are more frequent at the bottom of the ”Lim- burg“ area, and therefore at the border of the study area, the higher percentage of rare species on this plots can be true to influences of the surrounding areas. This influen- ce became scarcer — regarding overall influences — with increasing distance of the studied plots from the border in the centre of the ”Limburg‘ area. 5. ZUSAMMENFASSUNG Von April 1995 bis November 1997 wurden in den Streu- obstwiesen des Naturschutzgebietes Limburg bei Weil- heim/Teck (48.38 N/9.23 E) auf einer Fläche von etwa 62,7 ha (857 Flurstücke, Abb. 1) Kartierungen, Erfassun- gen und Messungen der Grünlandnutzung und -häufig- keit, Vegetation und Carabidenfauna durchgeführt. Dabei wurden 17 Probeflächen mit einem teilweise seit über 25 Jahren unveränderten spezifischen Management ausge- wählt. Nutzungsformen waren: dreischürige Mähwiesen (3), zweischürige Mähwiesen (3), Mulchwiesen (4), Suk- zessionsflächen (3), eine Pferdeweide, eine Schaf-Stand- weide, eine Schaf-Umtriebsweide und eine 1994 aufge- lassene Schaf-Standweide. Bonner zoologische Beitráge 56 (2007) 15 Im Gebiet der 17 Probefláchen wurden insgesamt 68 Spe- cies registriert (Tab. 2). Auf annáhernd allen Fláchen wechselte in den einzelenen Jahren die relative Háufig- keit der dominanten Spezies. Dreischúrige Máhwiesen be- sitzen häufig eine einseitige Dominanzstruktur. Innerhalb der järlichen Spektren der Aktivitätsdominanz wurden auf allen Flächen kontinuierlich Neuzugänge und Abgänge re- gistriert. Im Areal seltene Species (Stetigkeitsklasse I: auf 1-3 Flä- chen präsent) besaßen Anteile zwischen 10 % und 42 %, die entsprechenden Werte für Stetigkeitsklasse II (auf 4— 6 Flächen präsent) betrugen 0 % und 67 % und für Ste- tigkeitsklasse III (auf 7-10 Flächen präsent) 20 und 64 % (Abb. 2, Tab. 3). Die Bewirtschaftung des Unterwuchses von Streuobstwiesen hinsichtlich der Beeinflussung der Zönosen wird diskutiert. Acknowlegments. We are grateful Jutta Deuschle for revising the English text. REFERENCES BAGUETTE, M. 1993. Habitat selection of Carabid beetle in de- ciduous woodlands of southern Belgium. Pedobiologia 37: 365-378. BAUER, L. J. 1989. Moorland beetle communities on ,,limestone habitat islands”: I. 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The arthro- pod community of the edge of an intensivly grazed pasture. In: SCHREIBER, K.-F. (1988): Connectivity in Landscape Ecol- ogy. Proc. ond Intern. Seminar of the International Associa- tion for Landscape ecology. Múnstersche Geographische Ar- beiten 29: 115-117. Mc FERRAN, D. M., MEHARG, M. J., MONTGOMERY, W. I. & MCADAM, J. H. 1994. The impact of grazing on communities of ground-dwelling beetles (Coleoptera, Carabidae) in upland vegetation in north-east Ireland. Pp 325-330 in: DESENDER, K. et al. (eds.) Carabid beetles: Ecology and Evolution. Kluw- er Academic Publishers, Dordrecht etc. Morris, M. G. & Rispin, W. E. 1987. Abundance and diversity of coleopterous fauna of a carcous grassland under different cutting regimes. Journal of Applied Ecolology 24: 451-465. PREISZNER, J. 1996. Effect of habitat heterogenity on Carabid Populations on a sandy grassland. Acta Phytopathologica En- tomologica Hungaricae 22: 433-438. Rope, M. 1993. 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Ökologische Untersuchungen an bodenbe- wohnenden Coleopteren einer Heckenlandschaft. Zeitschrift Morphologie und Ökologie der Tiere 53: 537-586. TIETZE, F. 1985. Veränderungen in der Arten- und Dominanz- struktur der Laufkäferzöosen (Coleoptera — Carabidae) bewirt- schafteter Graslandökosysteme durch Intensivierungsfaktoren. Zoologische Jahrbücher Systematik 112: 367-382. TIETZE, F. 1987. Changes in the structure of carabid beetle Taxo- coenoses in grassland affected by intensified management and 16 Jürgen DEUSCHLE & Erich GLUCK: Colonisation and Steadyness of Carabid Beetles in Orchards industrial air pollution. Acta Phytopathologica Entomologia Hungaricae 22: 305-319. VOWINKEL, K. 1996. Eignen sich Carabiden als Indikatoren fúr Nutzungsintensitätsunterschiede im Grünland”? Artenschutz- report 6: 57-60. Authors’ addresses: Prof. Dr. Erich GLÜCK (correspon- ding author), Gingener Weg 61/1, 73072 Donzdorf, 0049- 7162/25360; E-Mail: glueck.donzdorf@t-online.de; Dr. Jürgen DEUSCHLE, Käthe-Kollwitz-Str. 14, 73257 Köngen, 0049-7024/805326; E-Mail: deuschle@tloe-deuschle.de. Received: 23.04.2007 Accepted: 31.07.2007 Corresponding Editor: M. Schmitt Bonner zoologische Beitráge Band 56 (2007) Heft 1/2 Seiten 17-24 Bonn, März 2008 Ten Instars in the Leprous Grasshopper, Phymateus leprosus (Fabricius, 1793) (Caelifera: Pyrgomorphidae): Maximum Number Recorded in the Acridoidea G. KOHLER!), S. ROTH?) & K. REINHARDT?) Institute of Ecology, Friedrich-Schiller-University, Jena, Germany 2)University of Bergen, Department of Biology, Bergen, Norway 3)Department of Animal and Plant Sciences, The University of Sheffield, Sheffield, UK Abstract. First instars of the pyrgomorphid Phymateus leprosus (Fabricius, 1793) were collected in the Great Karoo (South Africa). In the field about 190 poppers were distributed in small groups over 1.5 m? during the day, but densely aggregated during the night. The hoppers reared in ca pare were fed with Vincetoxi- cum, Euphorbia and Rubus. The mortality of the first instar hoppers was 97 %, but few of those that survi- ved reached adult moult. Ten juvenile instars of variable duration (2-8 weeks each) occurred in both sexes, which is the highest known stage number in the Acridoidea. Postfemur and body length measurements (exu- viae) as well as short descriptions are presented for each instar. The juvenile development lasted about one year, and the phenology corresponded with ce field observations by BisHop (1940). One male and three female grasshoppers completed the biennial life cycle (April 2000 to January 2002). The adult lifespan amounted to 2-8 months, with mating attempts after three months, but no egg pods were produced. The ovariole numbers varied from 116 to more than 187. Fully developed cegs were found in females that E died in July/August, but not in a female that died the following January. The meters of the genus Phymateus are summarized. w known reproduction para- Keywords. Phymateus, juvenile stages, biennial life cycle, reproduction, South Africa. 1. INTRODUCTION The genus Phymateus belongs to the tribe Phymateini within the family Pyrgomorphidae (KEVAN & AKBAR 1964). It is distributed with eight species in continental Africa south of the Sahara and two species in Madagas- car (distribution maps in ANONYMOUS 1982). These giant and colourful Orthoptera are popularly known as bush grasshoppers or bushhoppers, sometimes (and erroneous- ly) as bush locusts or milkweed locusts. They inhabit semi- deserts with Karoo vegetation, tree savannas, woodland, as well as gardens and other cultivated areas (ANONYMOUS 1982, JOHNSEN 1990, PICKER et al. 2002). They can cause damage in several crops (summarized by Uvarov 1977, ANONYMOUS 1982, SITHOLE 1986). This initiated studies on their biology, dealing with life cycles (BisHop 1940, DE Lotto 1951), food habits (KEVAN 1949, CHAPMAN 1962), and hopper behaviour (ROFFEY 1964, ROWELL 1967). The chemical defence of the pyrgomorphids includ- ing Phymateus was summarized by WHITMAN (1990). Re- cently, the genus became a study object in biochemical and endocrinological research (e.g. GADE & KELLNER 1995, GADE 2002), pharmacophagy (SEIBT et al. 2000), and flight capability (KUTSCH et al. 2002). Ephemeral studies dealt with local dispersion, thermoregulation, and short-term movement of adult bush grasshoppers (KOHLER et al. 1999, 2001a). Despite of these research activities, few data ex- ist about the complete life cycle in either the field or the laboratory. Information on hopper stages are available from only two Phymateus species (BisHop 1940, DE LOT- TO 1951, KAUFMANN 2000). Therefore, the present article focuses on the instars of Phymateus leprosus (Fabricius, 1793), particularly their number, morphometry and dura- tion. The species, here popularly named as leprous grasshopper (according to SAMWAYS 2005), was reared over its whole life cycle in the laboratory, enabling us for the first time to observe an exact number and differences of the instars. 2. MATERIAL AND METHODS In the afternoon of the 24th April 2000 a bulk of densely aggregated hatchlings of P /eprosus was found in the Eas- tern Great Karoo, South Africa, about 100 km N Port Eli- zabeth near a camp with typical Karoo vegetation (S.R.). With respect to an ovariole number >200 per female (see Table 2) and the same size and colour of hoppers within the bulk, it was very likely the complete offspring of one egg pod deposited nearby in the soil, but which was not found. Most of the hoppers were collected next morning in a plastic contain and, together with some not specified native plant species, transported to Jena/Germany. Later the adults were identified as Phymateus leprosus (Fabri- cius, 1793). 18 Günter KÖHLER et al.: Ten Instars in the Leprous Grasshopper About one week after collection the hoppers were reared in a terrarium (1 34 x w 18 x h 46 cm) in the Institute of Ecology (G.K.), placed on the sunny window directed to SSE. During field-work in the Great Karoo the adults of P. leprosus were mostly found on Asclepias bushes (KOH- LER 1999, 2001a). And BisHop (1940) mentioned a fre- quently attacked Euphorbia species. Therefore we offered food plants of the same families, Vincetoxicum hirundi- naria (Asclepiadaceae) and Euphorbia cyparissias (Eu- phorbiaceae), from the surroundings of Jena from May to October 2000. Because these plants were not available in winter of 2000/2001, leaves of Rubus sp. (Rosaceae), a generally accepted food plant for herbivorous insects, were provided during further captivity. According to ANONY- MOUS (1982) and our field observations grass is not eat- en by the species. In regular intervals the exuviae of the hoppers were col- lected and preserved dry. Later the length of the left hind femur and the body length from these exuviae were meas- ured with an Electronic Vernier Calliper (Helios-digit, ac- curacy 0.02 mm). The formation of wing pads was not- ed. For measuring the early instars a stereoscope (x10) was used. The exuviae of all instars are deposited in the Insti- tute of Ecology at the University of Jena. The culture was maintained until the death of the last adult. Immediately after death, the females were dissected for classifying the ovarian development according to the terminology (I-IV) of PHIPPS (1949) and for counting the ovarioles resp. eggs. Later the preserved and mounted adults were also meas- ured (body length, pronotum, postfemur, forewing). For not stressing the few individuals, no photos or line draw- ings were made. The sex ratio was determined from the dead first instar hoppers, and from those that survived. To do this, the hardened hoppers were softened with water vapour. Despite the black colouration the sex of most in- dividuals was identified by the shape of the external geni- talia of the terminal abdominal segments (subgenital plate in male, upper and lower ovipositor valves in female — as described in UVAROV 1966). 3. RESULTS 3.1. Offspring, mortality, and hopper aggregation From the aggregated bulk in the field (Fig. 1), altogether 172 first instar hoppers had been sampled, whereas about 5-10 % had escaped. Thus, about 180-190 hoppers should have hatched from this obviously one egg pod. The sex ratio of the bulk could be confirmed as nearly equal, with altogether 84 male and 80 female first instar hoppers, and 8 of unknown sex. These first instar hoppers suffered from a very high mortality, perhaps to a lower extent by the transport stress and after that mainly by the habituation to new food plants in the culture (Vincetoxicum, Euphor- bia). Otherwise, from the potential native food plants giv- en into the plastic contain only few were eaten during the transport. After one week of transport, 30 dead hoppers were found in the plastic contain, resulting in a mortali- ty of 17 %. During the following two weeks in the terra- rium nearly all individuals of the surviving bulk also died, so the total first instar mortality amounted to 97 %. On- ly five hoppers moulted in early May to the second instar, but these five survived to adult moult, with Vincetoxicum, Euphorbia and Rubus as food plants. Fig. 1. Bulk of first instar hoppers of Phymateus leprosus (bo- dy size about 6 mm) showing aggregation behaviour in the la- te afternoon. Eastern Great Karoo, South Africa, late April 2000. Photo: S. Roth. The field observations in the Karoo showed that in the af- ternoon (24.04.2000, about 20°C, some clouds) the indi- viduals were distributed over an area of about 1.5 m? most- ly in small groups of five to ten hatchlings sitting at a height of 30-40 cm on high shrubs and grasses. With de- creasing evening temperature and during the relatively chilly night (8-10 *C) all individuals gathered into a sin- gle dense bulk occupying about 0.15 m in diameter (Fig. 1). The aggregation behaviour was also observed in the following days within the plastic contain: dense aggrega- tions during the night and looser ones during the day. 3.2. Number, colouration and duration of instars Both hind femur and body length of exuviae originating from successive moults showed ten size classes with al- Bonner zoologische Beitráge 56 (2007) 19 Table 1. Morphometrical parameters of all developmental stages of Phymateus leprosus (Fabricius). Measurements: in juveniles — exuviae, in adults — dry mounted specimens. One male and four (juv.) resp. three females (ad.). In brackets — length influenced by shrinkage. Parameter Postfemur length (mm) Body length (mm) instar / sex male female male female 1 2.9 2.93.0 6.8 6.2-6.7 2 3.9 3.9-4.3 8.4 7.1-8.0 3 5.0 4.7-5.0 10.1 8.8-9.6 4 5.8 5.8-6.6 10.7 10.2-11.8 5 7.4 7.3-8.3 12.7 11.8-13.7 6 9.0 8.5-10.7 16.0 13.8-16.1 7 ES 10.9-13.2 [15.5] 16.8-17.8 8 14.9 13.1-16.4 19.1 20.1-25.1 9 177: 18.1-19.7 24.9 27.1-34.7 10 23.3 23.1-25.8 42.8 42.5-42.8 adult 30.0 30.0-33.0 [50.0] 58.0-62.0 Fig. 2. Last instar nymph of Phymateus leprosus (body size about 40 mm). Great Karoo (S.A.), Tierberg Research Centre. Photo: J. Samietz. most no overlap in the male and females of P. leprosus (Table 1). According to the location of the wing pads, there are eight hopper (L1-L8) and two nymphal stages (L9-L10; Fig. 2). There is an isometric growth between body and postfemur length (Fig. 3). The body size of both sexes was largely identical over the entire juvenile devel- opment, but showed considerable differences in the adult stage (Table 1, Fig. 3). The instars one to three were mostly black with several small yellowish or orange markings (Fig. 1). Two light spots were on the frons. On the upper and lateral prono- tum as well as along the margins there were yellowish markings. On each of the first eight abdominal segments there was a lateral drop-shaped to trapezoid spot, where- as dorsally a fine creamy stripe occurred. Ventrally the tho- Phymateus leprosus postfemur (mm) 0 10 20 30 40 50 60 70 body length (mm) Fig. 3. Isometric growth in juvenile development of both se- xes in Phymateus leprosus. Male (n = 1), females (n = 3). See also Table 1. rax had four lateral spots near the coxae of the first and third pair of legs, while the antenna and legs were com- pletely black. In the female hoppers the markings appeared slightly larger than in the male hoppers. Beginning with instar four the lighter portions (more or less of a bright green) extended successively, beginning on the tibiae of the hind and median legs, followed by the pronotum and other parts of the body (Fig. 2). In both sexes, the anten- nal segments increased from eight (instar 1-4) over eleven (instar 5-6) and 15 (instar 7) to 18-20 (instar 8-9) and 21 (instar 10 — Fig. 2), and in the females they varied 20 Günter KÖHLER et al.: Ten Instars in the Leprous Grasshopper AM JJASONDJYJFFMA MJ JAS ON D J Fig. 4. Biennial life cycle of laboratory-reared Phymateus leprosus, starting with hatchlings, collected in late April in the Eastern Great Karoo (S.A.) and brought to the laboratory at Jena University. September December February September September November December February Eggs March April May June July Hopper rt Period Hoppers Adults W TI 4 SE) Js as: 5 Il Fig. 5. Biennial life cycle of Phymateus leprosus in the field in the Eastern Cape Province, South Africa (detail from BisHop 1940). slightly within an instar (exuviae). According to our ob- servations, the two prominent tubercles on the pronotum separating P. leprosus from other Phymateus species are distinctly visible from the 9% instar. There is a major change in colouration after adult moult to a mainly green or red morph, briefly described in KOH- LER et al. (1999). In the green morph the head, pronotum, tegmina and legs are olive-green or greyish-green, where- as in the red morph these parts are dark red (except some individuals with a green pronotum). As far as we know, the red morph of P. /eprosus seems to occur mainly in the Southern Great Karoo, whereas the green morph is dis- tributed from the Eastern Cape (and the coast) to Trans- vaal. The juvenile development extended over one year from 24. April 2000 (hatchlings collected in the field) to the be- ginning of May 2001 (adult moult in the laboratory) (Fig. 4). Despite the sunny place in the window and warm tem- peratures caused by a radiator (15-25 *C) development of the instars 8 to 10 retarded somewhat in the European winter months (Fig. 4). The duration of the instars varied from 4-6 weeks in instar 1-2 (May—June), 2-3 weeks in Bonner zoologische Beiträge 56 (2007) Table 2. Reproduction parameters in Phymateus species. Mean (min—max). Species Origin Ovarioles Eggs in a female Eggs per pod Source P. leprosus S. Africa 340, 367 BisHop (1940) S. Africa 360, 480 KOHLER et al. (1999) S. Africa 116, >187 157 present ms P. viridipes Eritrea 287 (233-335) [n=12] 135 (108-151) [n=6] De Lotto (1951) Ghana 282 CHAPMAN (1962) Kenia 200710 [n=?15] 162 (60-231) [n=?15] KAUFMANN (2000) Malawı >146 present ms Phymateus sp. 393 UVAROV (1966; cit. Phipps) instar 4-7 (July-September), 4-6 weeks again in instar 8-9 (October to January) and finally about eight weeks in instar ten (February-April). This phenology nearly per- fectly fits with field observations over five years (1933-37) in the Eastern Cape Province by BisHor (1940) and with observations of adults in the Karoo (KÖHLER et al. 2001b) (Figs 4 and 5). 3.3. Adult development One male and four females survived to adult moult, but one female died during the adult moult. After final moult, two females developed with somewhat scrambled tegmi- na and alae, only the male and one female had more or less normally folded wings. The adults eclosed in early Fig. 6. Mating Phymateus leprosus in the field. Blyde River Canyon, N-Transvaal, South Africa, January 1995. Photo: K. Reinhardt. 22 Günter KÖHLER et al.: Ten Instars in the Leprous Grasshopper May 2001, the last female died on 28. January 2002, re- sulting in a maximum adult lifespan of about eight months (Fig. 4). Two other females died on 9. July and 31. Au- gust 2001, reaching only two resp. four months of life- span, which was not related to wing folding failure. The adults obtained were all of the green morph and in the four body parameters measured (length, pronotum, postfemur, forewing) smaller than individuals from the wild in the Southern Karoo near Prince Albert (P.A., KÓHLER et al. 1999). The only reared male reached about 50 mm body length (P.A. 53-62 mm) and 30 mm postfemur length (P.A. 27-34 mm), and the three reared females reached body lengths of 58-62 mm (P. A. 67-83 mm) and post- femur lengths of 30-33 (P.A. 32-40 mm). Between 6. August and 10. September 2001 on six days it was observed that the male settled on the back of the female with the normal developed wings, although with- out actual coupling of the genitalia. This behaviour was seen mostly in the morning between 8:45 and 9:30 hours, on one occasion extending to 14:00 hours, and one case was seen only around 15:00 hours (Fig. 6). On 3. August 2001 a plastic cup with a mixture of garden soil and fine sand was put into the terrarium, but no egg pod found on 11. December 2001. During these months no egg-depositing behaviour was observed nor were egg pods found on another medium. The dead female from 9. July 2001 was in a reproductive stage (III) with 116 fully developed terminal oocytes (6.0 mm long, 1.5 mm in diameter). The female, which died on 31. August 2001, was around the ovulation (IV) and had 157 light brown eggs. The third and oldest female, which died on 28. January 2002, was in the pre-reproduc- tion stage (II) and had still poorly developed ovarioles with more than 100 on the left and more than 87 on the right (not clearly separated). Similarly, a female P. viridipes col- lected on 14. September 2001 (Mulanje Mountains, Malawi, K.R.), also had very thin, undeveloped ovarioles (stage II) with at least 75 and 71 on either side. Summa- rizing the few available exact reproduction data from the genus, a broad variability has been detected reaching in reared P. leprosus from 116 to >187 ovarioles resp. eggs per pod and in wild females from 340-480. In P. viridipes these numbers vary mainly between 100 and 300 (Tab. 2). 4. DISCUSSION Because of their occasional economic importance, the dis- tribution and biology of the ten Phymateus species are summarized in the “Locust and Grasshopper Agricultur- al Manual” (ANONYMOUS 1982). In addition to its consid- erable body size, members of the genus Phymateus exhib- it at least five common characteristics: polyphagous on trees and shrubs, hopper aggregation behaviour, slow ju- venile development, late adult maturation, and complex defence strategy of the adults. In P. leprosus, four of these characteristics are treated in the present paper, whereas the complex defence behaviour of this species was described as early as 1940 in BisHop (1940) and recently in KÖH- LER et al. (1999), but it is well-known in Phymateus species in general (DE VILLIERS 1985, WHITMAN 1990). 4.1. Food plants and rearing The biology of P leprosus has been studied only once nearly 70 years ago by BisHop (1940) in the Eastern Cape Province, South Africa, where this species became very abundant in 1933 near Coega, Port Elizabeth district. It invaded farmlands, causing considerable damage by feed- ing on pumpkins and other crops, and in a citrus nursery. In general, pyrgomorphid grasshoppers are non-gramini- vorous (GANDAR 1982). However, all Phymateus species use a wide range of food plants, including toxic families like Euphorbiaceae, Asclepiadaceae, and Apocynaceae, as well as many crop species and garden plants with prefe- rences for trees and shrubs (ANONYMOUS 1982, SITHOLE 1986). In the milkweed (genus Asclepias) the cardiac gly- cosides proved to be phagostimulants in P. leprosus (SEIBT et al. 2000). Despite this, there are few rearing attempts and therefore the juvenile and adult development is rarely documen- ted, possibly because of the high early mortality and be- cause it is time-consuming to follow the very extended life cycle. There are two studies in P. viridipes (DE LOTTO 1951, KAUFMANN 2000), one in P. morbillosus (REYNEKE 1941) and one in P. leprosus (BIsHoP 1940) that report suc- cessful rearing of laboratory populations over their com- plete life cycle. For P. leprosus BISHOP (1940) used mass and single rearing, but failed to ascertain the exact num- ber of and the differences between the instars due to con- tinuous high hopper mortality. Our hoppers suffered from a very high initial mortality, possibly caused by the un- usual food plants, despite them being from the same families, Asclepiadaceae and Euphorbiaceae, as those eat- en in the field (Bishop 1940, KOHLER et al. 1999). Final- ly, our rearing showed that after a period of habituation a complete development of the leprous grasshopper up to the adult stage is also possible with non-native food plants. As secondary plant compounds, Vincetoxicum arundina- ria contains bitter glycosides, whereas in the milk of Eu- phorbia cyparissias some phenolic substances were found (HEGNAUER 1962-2001). Our successful rearing with Rubus spec. as food plant from instar 8 to the death of adults suggests that leaves of Rubus alone are sufficient for Phymateus in captivity. Bonner zoologische Beitráge 56 (2007) 23 4.2. Hopper aggregation Several species of Pyrgomorphidae (and of Romaleinae) are typically gregarious as hoppers, but not as adults, a phenomenon named as juvenile gregariousness by Uvarov (1977). In the Phymateus species hitherto stu- died at most the young hopper stages remain gregarious forming dense aggregations on the food plants and bands of at least hundred individuals moving on the ground (e.g. P. morbillosus — REYNEKE 1941, P. aegrotus — KEVAN 1949, P. viridipes — DE Lotto 1951, CHAPMAN 1962). In later instars the tendency to aggregate becomes weaker (Uvarov 1977), and immature adults tend to disperse, but reaggregate to some extent for mating and oviposition (ROFFEY 1964, ROWELL 1967). In P. leprosus this aggre- gation behaviour, as here observed for a bulk of first in- star hoppers, was already described by BisHop (1940) oc- curring immediately after hatching in the field and also in captivity. He observed that after any disturbance (even the shadow of the observer) the hoppers scattered in all directions in search of dense vegetation, and later they reaggregated again. This suggests that dispersion rather than aggregation is an anti-predator behaviour. In the same manner the hoppers behaved when collected in the field and brought into a plastic contain (S.R.). According to ob- servations of DE LOTTO (1951) on P. viridipes and the few observations presented here on P. /eprosus, aggregation is also affected by the circadian light-dark-regime. This suggests that hopper aggregation seems to have a ther- moregulatory function. 4.3. Instar number and phenology Apparently, the hoppers of all Phymateus species are very similar in colouration and morphological details and hard to distinguish, something that already KARNy (1910) had to confess. So BisHop (1940) described juvenile stages of P. leprosus, which resembie considerably those of P viridipes from Eritrea, described in detail and with fine drawings by DE Lotto (1951). In P. leprosus BısHoP (1940) estimated from mass rea- ring at least six juvenile instars in both sexes. In P viridipes seven instars (DE Lotto 1951), or six (males) and seven instars (females) were exactly observed (KAUF- MANN 2000). Differing from both studies, we found alto- gether ten instars in males and females, which is the high- est number presently known in the Acridoidea. Previous maxima reached up to seven to eight also in the Pyrgo- morphidae, and nine instars in both sexes of one species, the Bombay locust, Patanga succincta (RAMSAY 1964, Uvarov 1966). The instar number in Caelifera is relative- ly stable (despite unfavourable rearing conditions), so in several acridoid species variability by one instar has been shown with geographic origin or food quality (RAMSAY 1964, Uvarov 1966), but never three or four additional instars as observed here. Because of their long juvenile and adult development Phy- mateus species have a biennial life cycle with reproduc- ing adults occurring every two years. The onset of the cy- cle may vary geographically (Uvarov 1977, ANONYMOUS 1982). According to BisHop (1940), P. leprosus from the same region as our individuals, occurs ın the wild from June to March as eggs in the soil with a dormancy peri- od of at most six months (September to February). From March to next March the hoppers and nymphs develop, and from March to November the adults are present. Be- cause of an extended egg-laying period over three months a considerable overlap of all stages from two to three months occurs in the field (compare Fig. 5). Such long duration of juvenile and adult period in the field cor- responds closely to our rearing results, although the cy- cle is slightly delayed compared to that presented by BIsH- oP (1940). When the hoppers were brought to Central Eu- rope, the altered temperature and photoperiod may have had some influence on the developmental period. This is apparent both in the variable duration of the instars and in the development of the adults. Our first adults moult- ed in May, whereas BisHop (1940) mentioned April, and in 1995 a Karoo population of immature adults occurred even in mid-March with still poorly developed ovaries in the females (KOHLER et al. 1999, 2001b). The ovarian mat- uration should last about two months, so we found in cap- tivity females with fully developed eggs in July/August, but not in January. Apparently, no egg pod of P. /eprosus has, as yet, been found in the field. In addition, females in captivity never laid eggs, despite a large number of reared adults (BisHop 1940) and a few observed mating attempts in our culture. So the information about egg num- bers was reached by dissecting gravid females (BISHOP 1940, KOHLER et al. 1999). The few known reproduction parameters in the genus Phymateus show for P. leprosus and P. viridipes a considerable variation in ovariole num- ber, eggs in a female and eggs per pod (Table 2). 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Pp. 357-391 in: CHAPMAN, R. F. & JOERN, A. (eds.) Biology of Grasshoppers. John Wiley & Sons, New York. Authors’ addresses: Gúnter KOHLER (corresponding au- thor), Institute of Ecology, Friedrich-Schiller-University, Dornburger Str. 159, D-07743 Jena, Germany; E-Mail: Guenter.Koehler@uni-jena.de; Steffen ROTH, Universi- ty of Bergen, Department of Biology, Postbox 7803, N- 5020 Bergen, Norway; E-Mail: steffen.roth@macnews.de; K. REINHARDT, Department of Animal and Plant Sciences, The University of Sheffield, Sheffield S10 2TN, UK; E- Mail: K.Reinhardt@sheffield.ac.uk. Received: 8.12.2005 Revised: 14.06.2006 Accepted: 20.06.2006 Corresponding editor: D. Stiining Comment from the Editor-in-Chief: Because we had some special issues to publish to which an external manuscript could not be added, the publication of the present paper was considerably delayed. We apologise to the authors and to the readers and hope that data and results presented here will find interested reception anyway. Bonner zoologische Beitráge Band 56 (2007) Heft 1/2 Seiten 25-35 Bonn, März 2008 The Impact of Body Mass on Morphological Integration in Avian Skeletons (Aves, Fringillidae; Carduelinae, Fringillinae) Renate VAN DEN ELZEN!) & Hans L. NEMESCHKAL?) DZoological Researchmuseum Alexander Koenig, Bonn, Germany 2)Department of Theoretical Biology, University of Vienna, Austria Abstract. The dependence of the individual and of correlated skeletal variables of Carduelinae and Fringillinae upon bo- dy mass (=size) was tested by means of multiple regression and single linkage R-cluster analyses. The most obvious re- sult is the significantly high degree of body mass dependences of all individual variables within the three functional com- plexes feeding, hind limb locomotion and flying. Elimination of body mass as a measure of size decoupled several units of correlated variables. Nevertheless the general robustness of units was strong but the removal of “size” had an impact on the extent of morphological integration, measurable as agglomeration level of variable units. Body mass (size) in our sample thus influenced the quantity rather than the quality of morphological integration. The key feature in fringilline/car- dueline skeletal morphology — coevolution of skull and hind limb traits — thus appears to be relatively unconstrained by size. Keywords. Size, ecomorphology, functional complexes, biological roles, skeletal morphology, developmental gene- tics. 1. INTRODUCTION 1.1. The role of body size in avian morphology Body size has been identified as an important factor driv- ing the diversification and evolution of organisms. OLSON & MILLER (1958, 1999), e. g. quantified size as a strong integrative force of morphology, FAIRBAIRN (1997) iden- tified its role in sexual dimorphism, KLINGENBERG & SPENCE (1997) analyzed its function in life-history evo- lution. BONNER (2004) proposed a third size dependent rule besides the already well known “weight-strength-rule” and “weight-surface-area-rule”. His “weight-complexity- rule” explains that increase and decrease in size of organ- isms are strongly related to increase and decrease in com- plexity in general, from differentiation of cell types to the social organization of organisms. These examples implement size as a morphological key feature in the organization of life, at all hierarchical lev- els from the individual to species (BARBOSA et al. 2000; BONNER 2004; CHERNOFF & MAGWENE 1999; PETERS 1993). Size, as the physical magnitude of objects, can be quan- tified using either measures or weights. Linear length measures are used to seize both the two dimensional area as well as the three dimensional volume, whereas weight quantifies the proportionality to volume alone. Measures widely accepted to represent overall size of organisms in- clude also factors like the first principal component (BOOKSTEIN et al. 1985; ZELDITCH & FINK 1995; LEISLER & WINKLER 2006), landmarks or ratios. BIORKLUND & MERILÄ (1993), e.g. used orthogonal rotation to remove size from species mean vectors, LEISLER (1980) calculat- ed the cube root of a compound measurement of the body core skeleton (sternum length plus pelvis length plus cora- coid length) times (sternum width plus pelvis width) times height of crista sterni as measure for “body size”, follow- ing HOERSCHELMANN (1966). For the characterization of volume, body mass is the ad- equate measure. Several rules explaining the construction- al constraints on morphology are related to body mass: The “weight-strength-rule” already mentioned above, ex- plains why e.g. longer arms, wings and legs have to be proportionally thicker than shorter ones, the “weight-sur- face-area-rule” describes physiological relations of e.g. gas diffusion in large versus small lung surfaces. These are geometric growth laws, that explain the change of propor- tions — so-called allometric relations — within a biologi- cal organism. The underlying process is differential growth during development brought about by differences in the growth rates of the various parts (WADDINGTON 1966). In small bodied flying vertebrates, flight imposes identi- cal physical constraints on the flying organism that lead to a functional symmetry in body sizes (MAURER et al., 26 Renate VAN DEN ELZEN & Hans L. NEMESCHKAL: Impact of Body Mass on Morphological Integration in Avian Skeletons 2004). The body plan of birds is primarily constrained by the weight-strength-rule, that regulates flying ability. In extant species 22 kg is the limit for take off; the world- wide tiniest bird 1s the Bee Hummingbird, Mellisuga he- lenae, found in Cuba and weighing from 1,6-1,9 g (CHAI 1999). The most successful clade in modern birds, the passerines (or perching birds, Passeriformes) have minia- turized to an average body mass minimum of 4,5 g in some tropical species (Black-faced Flycatcher Warbler, 4bro- scopus schisticeps, Broad-billed Flycatcher Warbler, Tickellia hodgsoni, Cuban Gnatcatcher, Polioptila lem- beyei; DUNNING 1993). The heaviest and thus largest song bird, the Raven, Corvus corax, from Alaska weighs 1.240 g. Perching birds successfully evolved and radiated with- in any terrestrial habitat -with exception of the polar re- gions where their occurrence is physiologically con- strained by the weight-surface-area-rule. 1.2. Key features of the carduelid skeleton Our paper focuses on the impact of body mass on the ar- chitecture of the avian skeleton, taking the song bird sub- families Fringillinae and Carduelinae as a case study. Mor- phological diversity in skeletal elements in this phyloge- netic entity (clade, monophylum) is mostly confined to changes in overall size (BJORKLUND 1991, 1994; VAN DEN ELZEN & NEMESCHKAL 1986, 2006; NEMESCHKAL & VAN DEN ELZEN 1990, VAN DEN ELZEN & NEMESCHKAL 2006). Differentiations in shape are (despite their unquestioned ecological importance) only expressed as an insignificant portion of the total variation. Whereas three shape axes explained only 9,3 % of variance, the size axis account- ed for 85,5 % of variance (VAN DEN ELZEN & NEMESCHKAL 2006). The observed pattern has been identified as a gen- eral rule in passerine birds by BIÖRKLUND (1994, 2006). “*... making species to be larger or smaller copies of each other, all moving up and down a common line of allo- metry”. Mapping of skeletal variables along the first prin- cipal components (NEMESCHKAL & VAN DEN ELZEN 1990) revealed, that morphological diversification in Fringilli- nae and Carduelinae focuses on variables of the beak. Thus, morphological change in these subfamilies is main- ly reflected in feeding mechanisms. In terms of ecomorphology, a discipline taking the bio- logical role of morphological structures under consider- ation (BOCK & VON WAHLERT 1965), elements of the avian skeleton belong to three different functional complexes: skeletal elements of the skull and beak are allied to the feeding complex, wing bones, pectoral elements and bones of the shoulder girdle represent the flying apparatus and skeletal elements of pelvis and legs the hindlimb locomo- tion complex. In the carduelid skeleton, nevertheless, units of covarying variables (UCCs) contain combinations oth- er than proposed by the functional complexes described above. These UCCs were detected to reflect units carry- ing out common biological roles. Besides a more or less undisturbed module of the “flying complex”, formed sole- ly by wing bones plus elements of the shoulder girdle, the variables of the feeding and hindlimb locomotion com- plexes were united in one single common cluster, and demonstrate the superimposure of feeding on hindlimb lo- comotion. In search of additional modern approaches in the study of morphological integration, CHERNOFF & MAGWENE (1999) propose a hierarchical framework for integrative hypothe- ses: Morphological variables are judged as integrated at the broadest, i. e. the uppermost inclusive level due to their covariation with size. At a less inclusive level variables covary due to developmental and/or functional associa- tions and at still lower levels due to anatomical (spatial) associations. Starting out from physics we may assume that changes of singular parts from a macroscopical phys- ical (e. g., mechanical) system will be ruled by various law-dependencies when the whole system changes size (e.g., mass). As a consequence, we test in the current study the dependence of individual and correlated skeletal vari- ables upon body mass taken as a measure of size. 1.3. Presumptions Based upon our knowledge from preceding analyses (VAN DEN ELZEN, NEMESCHKAL & CLASSEN 1987, NEMESCHKAL & VAN DEN ELZEN 1990, NEMESCHKAL, VAN DEN ELZEN & BRIESCHKE 1992, VAN DEN ELZEN & NEMESCHKAL 2006) our hypotheses are that: l.a significant dependence of skeletal variables on body mass exists. It is expected to differ between variables and to be highest in beak elements; 2.thus the three theoretical functional complexes -feed- ing, hindlimb locomotion, flying- differ in their size de- pendence; a. variables of the feeding apparatus — showing highest morphological differentiation between species — should exhibit the largest amount of size dependence, b. elements of the hindlimb locomotion complex and of the flying apparatus are thought to show lesser dependence, as of lesser interspecific variability. 3.units of variables (UCCs) are also expected to be size dependent, elimination of “size” is thought to decouple variables within units; decoupling is expected to occur particularly within “mixed” variable units of beak and hindlimb osteometrics. Bonner zoologische Beiträge 56 (2007) 27 2. MATERIAL AND METHODS In 313 specimens, representing 43 species (Table 1), 42 equivalent skeletal variables (measurements of bone lengths, widths and depths) were taken, as described in detail in NEMESCHKAL (1999). Twenty measurements stem from the feeding apparatus, 13 represent the flying appa- ratus and 9 hindlimb locomotion (Table 2). Firstly, centroids of 43 species over 42 variables were built from the log scaled original data matrix consisting of 313 specimens. This procedure was essential, because body mass data were available as species means only. Morpho- logical variation between species was then quantified by the total variance of species centroids. Body mass varia- tion in the actual fringilline-cardueline sample ranges from a maximum of 54 g in the Hawfinch Coccothraustes coc- cothraustes to a minimum of 9 g in neotropical siskins (Table 1). To test hypotheses about size impact, linear re- gression analysis was applied. Log transformed species means of body mass were taken as predictor variable and each of the 42 skeletal measurements as criterion variable. The 42 resulting residuals are used as the variables under study — the variables corrected for body mass (= variables of which size was partialized out; BCM). The coefficients of determination between body mass and original variables (Table 2) were tested for significance using random boot- strap (1000 replicates each; for computer programme package see NEMESCHKAL (1999)). They are figured as profiles of variance (Figs 1, 2). Single linkage R-cluster analyses were chosen to figure correlations of original da- ta (Figure 3- BO; based upon the variable intercorrelation matrix between species means for original data) and cor- relations data corrected for body mass (Fig. 3- BCM; vari- able intercorrelation matrix between species means for body mass adjusted data). Units of correlated variables were taken into account, when they agglomerated at the uppermost quartile (r? > 0.924) in the original dataset and reappeared in the size corrected dataset again. E-units are stable clusters with topological identity of variable posi- tions in both the R-cluster analyses of original data and the body mass corrected data set. S-units are less stable units built of identical variables but with topological dif- ferences between analyses of original and size corrected variables. 3. RESULTS The most obvious findings are on the one hand the sig- nificantly high degree of body mass correlation of all in- dividual variables (Fig. 1) and on the other hand the ro- bustness of several covarying variable units found in both the original and the “residual” data set, adjusted for body mass (Fig. 2). 0,8 1 /— 0,7 4 -- 0,6 + 0,5 coefficient of determination 0,4 en a BERN BEER BERN BER BER FEN BEE BER BERGE oe TE TR 0,3 Ll 4 1 3 5 7 9 11 13 45. 17-19 21, 23, 25: 27.29 31 33 238 37 skeletal characters Fig. 1. Dependence of 42 skeletal characters on body mass as shown by coefficients of determination. 28 Renate VAN DEN ELZEN & Hans L. NEMESCHKAL: Impact of Body Mass on Morphological Integration in Avian Skeletons variances and residuals 13 5 7 9 1% 13 45 47 19 21 23 25 27 29 31 23-35 37 ac skeletal characters —— BO --- BCM Fig. 2. Comparison of variance profiles. Variance of original characters (BO) occur in the upper line and residuals for characters corrected for body mass (BCM) at the bottom. 0.696 0.772 0.848 0.924 1 1.0 0.797 0.594 0.391 0.188 pl eee Figs 3. Single linkage R-cluster analysis of original data (BO; left) and of body mass corrected data (BCM; right). Black squares indicate measures of the pectoral girdle and forelimbs, black triangles mark skull measurements and open circles measures of the pelvic girdle and hindlimbs. Clusters are explained in the text. Bonner zoologische Beiträge 56 (2007) 29 3.1. Influence of body mass on individual variables All 42 variables are correlated with body mass at a high significance level (p< 0.001) as revealed by random boot- strap procedures. The degree of correlation with “size”, measured by coefficients of determination (COD) between (log transformed) original data of species means and body mass, is highest in coracoid length (variable number 35; COD=0,91), post orbital length (12; 0,90) and equal at COD= 0,89 in tibiotarsus width (27), total skull length (16) and synsacrum length (21). The largest amount of body mass independence exhibit carpometacarpus (41) and ul- na lengths (39; COD= 0,34 and 0,47) as well as tar- sometatarsus length (No 28) and width (29; COD= 0,70, resp. 0,67) and proximal end width of the humerus (38; COD= 0,71). Skull variables in general reveal the high- est correlation with body mass, hindlimb and forelimb variables indicate a larger variation in their correlation with body mass. A comparison of variance profiles of original and size cor- rected skeletal variables (Fig. 2) confirms and strength- ens these findings: The relatively higher variances in the original variables appear smoothened in the size correct- ed residuals (BCM; bottom line in fig. 2), indicating that the largest amount of total variance between species is due to a “size factor”, measurable by body mass. This espe- cially holds true for skull variables (skeletal variables 1-20) and less for hindlimb and forelimb measures (skele- tal variables 24-29, 37-42). 3.2. Influence of body mass on united variables The single linkage cluster analysis of original data (Fig. 3) extracted two main units fused at a high level, several disintegrated variables and one isolated variable set (E1). The module BOC 1 is composed of the variables of the flying apparatus, mainly its “engine-supporting” part, in- cluding measures of sternum, coracoid and scapula. Wing measures that represent “flight performance” are highly disintegrated, only humerus length (37) and carpometacar- pus width (42) are included. The second main module BOC 2 combines skull and hindlimb measures. The two main modules BOC 1 and BOC 2 are composed of two different types of subsets: Robustness 1s observed in nine smaller units (E1-E9), where only two to three variables are coupled. Similar structure is maintained af- ter correction for size in four units (S1—S4), including three to 17 variables as well as including several of the stable units (E4-E9). Whereas, for example, femur length (24) and tibiotarsus length (26) are next neighbours in the S4 cluster in BO, in BCM tibiotarus length (26) and tar- sometatarsus length (28) are agglomerated at the highest level. The larger SI and S2 units also include some of the smaller, topologically stable E units: E4-E6 always oc- cur within SI, E8-E9 within S3, S4 is always part of Sl. BOCs identified in the R-cluster analysis of original da- ta are to a great extent also found in the analysis of body mass corrected data (Fig. 3). Again in the body mass cor- rected data analysis, a main cluster composed of engine- supporting variables from the flight complex (S2 in BOC1, fig. 3) opposes a main cluster structured by a mixture of skeletal elements of the feeding and hind limb locomo- tion apparatus characters. The main differences to the orig- inal data analysis are the rather low agglomerative level (coefficients of determination in the “size-corrected” dataset being two to three times lower than in the origi- nal dataset), reduction of the involved variable units and some changes in variable topology. BOC2, that was com- posed of SI (including S4), S3, E2, E3 and E7 in the orig- inal dataset, comprises in the size corrected data set only S1 (including $4) and E3, the remaining components — units $3, E2 and E7 — are clustered at lower agglomera- tion levels. Subcluster SI differs in its hierarchical structure of ag- glomeration levels of E units and arrangement of variables within S clusters. In S2 sternum length (30), keel length (32) and keel depth (31) are differently arranged, in S3 post orbital length (12), and within S4 femur-, tibiotarsus and tarsometatarsus lengths (24, 26, 28) changed their po- sition along branches. In both illustrations of single linkage R-cluster analyses (Fig. 3), El shows up as a more or less disintegrated unit. It consists of measures of ulna (39) and carpometacarpus lengths (41), which are tied together at a relatively high agglomeration level, especially in the analysis of “size- corrected” data, in which both measures are combined with carpometacarpus width (42). 4. DISCUSSION 4.1. Size and morphological integration Do these results confirm or contradict our predictions and expectations of hypotheses on morphological integration? Corroborating our first assumption, a general body mass dependence was identified at a high significance level for all individual characters within all three classical function- al complexes. “Size” could thus be confirmed as a strong integrative force in cardueline and fringilline skeletal mor- phology. Variability was, as assumed in our second hy- pothesis, relying to a greater extent on body mass in skull and beak measurements than in characters allied to the oth- er functional complexes. Elimination of “size”, decoupled, 30 Renate VAN DEN ELZEN & Hans L. NEMESCHKAL: Impact of Body Mass on Morphological Integration in Avian Skeletons Table 1. List of species under study, their body masses and information sources. Sources: DU= Dunning 1993; RE= labels of specimens at ZFMK, Bonn; RO= McLean 1988; WA= Ward 2003. Species body mass in g sources Fringilla montifringilla 24 DU Fringilla coelebs 21,4 DU Coccothraustes coccothraustes 54 DU Pyrrhula pyrrhula 21,8 DU Pinicola enucleator 56,4 DU Loxia curvirostra 40,38 DU Chloris chloris 21,8 DU Chloris sinica 31,3 DU Acanthis flammea 13 DU Carduelis carduelis 15,6 DU Linaria cannabina 153 DU Agriospiza flavirostris 15,4 DU Rhodospiza obsoleta 23,9 DU Spinus spinus 14,5 DU Spinus cucullatus 3 RE Spinus barbatus 16,6 DU Spinus xanthogaster 12,7 DU Spinus yarrellii 35 RE Spinus psaltria 93 DU Serinus pusillus LG DU Serinus serinus 11,2 DU Serinus syriacus 12,1 BE Serinus canicollis 13,8 DU Serinus citrinella 12 DU Alario alario 11,8 RO Serinus canaria 12,97 DU, RE Ochrospiza reichenowi 11,5 RE Ochrospiza atrogularis 11,4 DU Ochrospiza leucopygia 9 RE Ochrospiza mozambica 10,6 DU Ochrospiza dorsostriata 14,4 DU Ochrospiza xanthopygia 10 RE Crithagra gularis 15,5 DU Crithagra striolata 22,4 DU Crithagra albogularis 25.9 DU Crithagra donaldsoni buchanani 24 DU Crithagra sulphurata 19,2 DU Serinops flaviventris 16,3 DU Crithagra mennelli 153 RO Crithagra burtoni 299 DU Dendrospiza hyposticta 1 DU Dendrospiza scotops 15,4 RO Pseudochloroptila totta 13,4 WA Bonner zoologische Beitráge 56 (2007) 31 as predicted in the third assumption, several units of cor- related variables. Nevertheless, contradicting our expec- tations, displacement of variables from units or units from larger modules was low and robustness of units strong. Body mass (size) itself in our sample thus influenced the quantity rather than the quality of morphological integra- tion and did not affect the key feature in fringilline/car- dueline skeletal morphology — coevolution of skull and hindlimb variables. According to classical hypotheses on morphological in- tegration, “size” 1s the uppermost integrative level, at deep- er agglomerative levels, traits covary due to developmen- tal or functional associations and at still deeper levels due to spatial adjacency (CHERNOFF & MAGWENE 1999) re- spectively morphological neighborhood (ALPATOV & BOSCHKO-STEPANENKO 1928). Indeed, in our study, re- moval of the first hierarchical layer “size”, mostly effect- ed the degree of morphological integration, measurable as agglomeration level of character units. Structure and re- lation of character compositions however remained to a great extent constant and robust. For variable units at the body mass reduced level, three types of trait correlations are identifiable: S-modules are subunits agglomerated at higher integrative levels within BOC1 and BOC2, the orig- inal, not body mass reduced, modules. E-modules that are not composed of characters from immediate morpholog- ical neighborhood or partial overlap (E4, ES, E9) can be assigned to two categories: Classical functional units (E2, E3, E6 and E7) are with one exception (E7) fused at low- er agglomerative levels (below 0.6 COD) than the units of biological roles El and ES (above 0.8 COD). Our find- ings thus corroborate the integrative hypotheses quoted above, that anatomical association is a primary integra- tive level in morphology. 4.2. Interpretation of the observed character mod- ules EBLE (2005) classifies four different kinds of modules in morphology: 1. modules due to structural relations, 2. modules of pleiotropic genotype-phenotype mappings, 3. developmental units (gene expressions, domains of epi- genetic dynamics, regions with localized allometric growth) and 4. functional units. We would like to extend the definition of the last catego- ry by modules accomplishing common biological roles (NEMESCHKAL et al. 1992) and thus differentiate between 4.a. mechanical functioning (like kinetics) and 4.b. biological functioning (like morphological traits used in e. g. feeding). Units E4 (postorbital width and internasal width, 9, 10), ES (skull width and interorbital maximal width, 6, 7) and E9 (premaxilla length and total skull length, 11, 16) may be explained by their immediate morphological neighbour- hood and /or partial overlap (“rule of neighbourhood”, AL- PATOV & BOSCHKO-STEPANENKO 1928; structural relations EBLE 2005). Most of the other results allow for mechan- ical functional explanations, few for explanations of their biological roles. E2 is composed of two skull characters (caudal length of the mandible and quadratojugal length, 3, 13) and a pelvis measure (synsacrum distal length, 23). Whereas the first two skull variables are functional coun- terparts and act in seed husking, their —also body mass in- dependent- correlation with a synsacrum length measure comes unawares. E3 combines two mandible depths (4, 5) with quadratum length (19), characters functioning in the kinetism of the avian skull, especially in lateral bill movements enhancing seed husking (NUIJENS & ZWEERS 1997). Also units E6 (skull height and premaxilla depth, 15, 20) and E7 (femur and tibiotarsus width, 25, 27) are easier to explain by constructional constraints than bio- logical roles. E6 depicts the interdependence of skull and beak height, E7 that of leg diameters, in ES.Only units El and ES where measures of the flying apparatus (car- pometacarpus (41) and ulna lengths (39)) respectively bill tip are tied together (length of pars symphysialis (1) with dentary length of the mandible (2)) allow for an interpre- tation of biological roles: The first unit comprises the feather-carrying bones of the wing, responsible for ma- noevrability in flight per se. Unit ES represents “the bill tip grasp”, a feeding tool well developed in carduelid finches (NEMESCHKAL et al. 1992). S4 with the lengths of femur (24), tibiotarsus (26) and tar- sometatarsus (28) obviously reflects a character pattern of concerting limb dimensions as guided by developmental processes during ontogeny (NEMESCHKAL 1999). Comparison of interspecific correlation patterns (macroevolutionary pattern, this study Fig. 3) with infra- specific analyses of variable correlations (microevolution- ary level, NEMESCHKAL & VAN DEN ELZEN 1994, fig. 7) re- veal that six modules (E3, E4, ES, E7, E8 and E9 of this study) correspond to units also found within carduelid finches and three modules (ES, E7 and E9) even to units occurring within the phylogenetically distant pigeons. At the microevolutionary level NEMESCHKAL et al. (1992) pro- vided evidence for clade specificity of variable units (mod- ules) and their correspondence to expressions of develop- mental control genes (NEMESCHKAL 1999). Morphologi- cal modularity might generally be seen as product of de- velopmental modularity, because morphological patterns of organization emerge in ontogeny (EBLE 2005). Conse- quently, each of the extracted variable complexes might additionally also correspond to developmental or body plan modules. 32 Renate VAN DEN ELZEN & Hans L. NEMESCHKAL: Impact of Body Mass on Morphological Integration in Avian Skeletons Table 2. Skeletal variables under study and their coefficients of determination. BO represent original data, BCM resi- duals for the body mass reduced variables. Skeletal variables Coeff. of determination Skull measures 01 mandible, pars symphysialis length 0,7014 02 mandible, dentary length 0,7641 03 mandible, pars caudalis length 0,7642 04 mandible, dentary depth 0,6924 05 mandible, pars caudalis depth 0,7883 06 skull width 0,8239 07 interorbital, maximal width 0,8353 08 interorbital, minimal width 0,6704 09 postorbital width 0,8048 10 internasal width 0,7856 11 premaxilla length 0,8240 12 postorbital length 0,86 13 quadratojugal length 0:7953 14 orbital width 0,7888 15 skull height 0,7280 16 skull length 0,8610 17 palatinum width 0,6176 18 pterygoid length 0,6932 19 processus orbitalis quadrati length 0,7988 20 premaxilla depth 0,7911 Pelvic girdle and hindlimbs 21 synsacrum length 0,7748 22 synsacrum width 0,7076 23 synsacrum distal length 0,7464 24 femur length 0,8355 25 femur width 0,7781 26 tibiotarsus length 0,7675 27 tibiotarsus width 0,7562 28 tarsometatarsus length 0,6180 29 tarsometatarsus width 0,3034 Pectoral girdle and forelimbs 30 sternum length 0,8413 31 keel depth 0,6897 32 keel length 0,7632 33 sternum, distal width 0,6617 34 sternum depth 0,7393 35 coracoid length 0,8401 36 scapula length 0,8114 37 humerus length 0,7644 38 humerus, proximal end width 0,6246 39 ulna length 0.3233 40 ulna width 0,6235 41 carpometacarpus length 052333 42 carpometacarpus width 0,6454 Bonner zoologische Beiträge 56 (2007) 33 How genes and developmental pathways co-operate is best documented for the avian beak. It is constructed by mul- tiple facial prominences (HELMS et al. 2005, HELMS & BRUGMANN 2007), where the frontal nasal mass (FNM), lateral nasal prominences (LNP), and maxillary promi- nences (MXP) comprise the upper beak and the mandibu- lar prominence (MDP) forms the lower beak. The identi- ties of facial prominences are specified early in the neu- ral crest stage and may involve homeobox genes such as Hox (CouLy et al. 1998) or MSX (BROWN et al. 1997, Wu et al. 2006). In metazoa, at least 17 signal transduction pathways op- erate to activate or repress different genes at distinct times and places in the embryo. Five predominate in early em- bryonic development: the Wnt, TGF-beta, Hedgehog, RTK, and Notch pathways. Five more are used in late de- velopment, and seven more in the functions of differen- tiated cells (GERHART 1999). Diversity in beak shapes of Galapagos finches, e. g., is known to originate from the activity of a morphogenetic bone protein (BMP4 of the TGF-beta family; ABZHANOV et al. 2004) and calmodulin, a protein that binds and trans- ports calcium ions (CaM; ABZHANOV et al. 2006). Where- as BMP4 stimulated growth of beaks along two dimen- sions, — it produced deeper and wider beaks and thus ex- plained the linkage in the variation of these traits, — the authors found CaM as regulator of beak length. Thus in Galapagos finches, beak length develops independently from width and depth due to two different factors that lead to changes in growth along different dimensions (ABZHANOV et al. 2006). In ducks, chickens and cockatiels, BMP4 is also differently expressed (WU et al. 2004, Wu et al. 2006). At late stages of development, chicken and duck embryos had two localized growth zones in the FNM, which melted in the chicken, but stayed separated in the duck. Ducks, moreover exhibited a wider FNM and more activity in another growth factor (fibroblast growth factor 8). In cockatiels a thicker FNM increased in a dif- ferent direction and the mandibular prominence (MDP) was suppressed. BMP4 was involved in all species in me- diating activity in all localized growth zones. Experimen- tal overexpression in BMP4 altered beak shapes among all species and beak curvation was induced by asymmet- ric growth activity in a facial prominence. BMP4 was al- so found to be responsible for regulating a homeobox gene: As it increased the expression of MSX1, the authors suggested that MSX1 activity is regulated by the BMP pathway. 4.3. Conclusions Under the assumption that BMP4 is a main regulator for the expression of developmental control genes in the avian skeleton, this protein, for instance, can be seen as a key growth factor accounting for size and shape variation in our skeletal variables. A developmental growth factor can also explain the strong size dependence of single variables, but cannot elucidate the interspecific correlation patterns of variable units. Thus in the light of current knowledge on gene expression variability and research in develop- mental genetics the classical view on morphological in- tegration and genesis of modularity might be too one- sided. WAGNER et al. (2007) offer a broader approach to modu- larity. Summarizing empirical evidence, they discern be- tween two levels: The elements the modules consist of and their kinds of interaction. Elements vary from molecular level (nucleotides) to morphological traits and their con- nections from physical to dynamical and statistical. They define three kinds of modules: Variational, functional and developmental modules, but stress a duality, as modules may belong to several categories at the same time. Units observed in the present study on cardueline/fringilline skeletal variables fall in the category of variational mod- ules, as their measure is statistical. For the interpretation of variational modules the authors offer several answers: identical developmental origin, similarity of covariation and gene expression territories, and pleiotropic effects. So our variable units (or character complexes; NEMESCHKAL et al. 1992) can be defined not only as a duality but a plu- rality: according to their operability they are functional modules, according to their mode of exploration they are variational modules, and according to their genesis devel- opmental modules. The open problem is, whether modules arise through the action of natural selection or because of biased mutation- al mechanisms (WAGNER 2007). Both neutral models and models based on natural selection are offered: e. g. LYNCH (2007) favors the neutral model and stated that “... emer- gent biological features such as complexity, modularity, and evolvability, all of which are current targets of con- siderable speculation, may be nothing more than indirect by-products of processes operating at lower levels of or- ganization.” We do not follow the author in refusing nat- ural selection and external evolutionary forces, but as- sume, that selection via fitness promotes certain pheno- types derived from their genetic basis and processes in on- togeny. Under the viewpoint of developmental constraints as internal evolutionary forces, we like to generalize the findings of Wu et al. (2004, 2006) on the origin of beak shape in birds. The authors offer a more reconciling con- clusion, that morphological diversity may be achieved by modulating prototypical molecular and cellular modules. 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Morphoregulation of avian beaks: comparative mapping of growth zone activities and morphological evolu- tion. Developmental Dynamics 235: 1400-1412. ZELDITCH, M. L. & Fink, W. L. 1995: Allometry and develop- mental integration of body growth in a piranha, Pygocentrus nattereri (Teleostei: Ostariophysi). Journal of Morphology 223: 341-355. Authors” addresses: Renate VAN DEN ELZEN (correspon- ding author), Section Ornithology, Zoologisches Forschungsmuseum Alexander Koenig, Adenauerallee 160, D-53113 Bonn, Germany; E-Mail: r.elzen.zfmk@uni- bonn.de; Hans L. NEMESCHKAL, Department of Theoret- ical Biology, University of Vienna, Althanstrasse 14, A- 1090 Wien, Austria; E-Mail: Hans.Leo.Nemeschkal@uni- vie.ac.at Received: 21.02.2008 Revised: 25. 06. 2008 Accepted: 09. 09. 2008 Corresponding editor: M. Schmitt Bonner zoologische Beitráge Band 56 (2007) Heft 1/2 Seiten 37-41 | Bonn, Marz 2009 Occurrence of the Genus Microscolex (Oligochaeta, Acanthodrilidae) at Western Canary Islands José A. TALAVERA & Dolores I. PEREZ Faculty of Education (Module A), University of La Laguna, Tenerife, Canary Islands, Spain Abstract. Two species of Microscolex are present in the western Canary Islands: M. dubius and M. phos- phoreus (new record for Hierro). The latter belongs to the modern fauna of the Canaries and was recently introduced by man. Both species prefer to live at the basal altitude level, mainly in the highly anthropized band between 50 and 300 m a.s.l. The habitat range of M. phosphoreus is rather narrow and its principal nu- cleus of dispersion are tropical fruit plantations (papaya, avocados and bananas); however, M. dubius is mo- re diversified, its range is wide, even reaching into the native Pinus canariensis and laurel forests at the mon- tane level. Other data provide evidence that the greatest number of specimens and locations are on the nort- heast slopes of the Canaries, as reflected in the first distribution maps of each of the islands. The biogeogra- phical affinities with Macaronesia are discussed, along with the probable links with Latin America as a long- standing (500 year-old) source for importing agricultural and ornamental plants into the Canaries. Keywords. Earthworms, Acanthodrilidae, Canaries, habitats, chorology. 1. INTRODUCTION The taxonomy and distribution of the Acanthodrilidae have been under study for a long time, BEDDARD (1895) and MICHAELSEN (1900) having compiled comprehensive information on these annelids in their magnificent scien- tific works. They are well known in Australasia, the Neoarctic and Neotropical regions, where they are spread over a wide range of both anthropized deforested habitats and those with abundant vegetation (e.g. tropical forests). The family is increasing in numbers and distribution area, as revealed by successive biogeographical records and de- scriptions of new taxa (CsuzDI & SZLÁVECZ 2002; FRAGOSOS 1988; FRAGOSOS & REC FERNANDEZ 1994; REYNOLDS & RIGHI 1994; Zicsi & Csuzoı 1991). In the Canary Islands, few acanthodrilids have been recorded in the literature to date. KRAEPELIN (1895) cites Microscolex poulteni (BEDDARD, 1892) as presumably from Tenerife but with no details. MICHAELSEN (1900) re- ports Microscolex dubius (Fletcher, 1887) and Mi- croscolex phosphoreus (Duges, 1837). Later, other authors merely copied them literally (COGNETTI DE MARTUS 1906; BALDASSERONI 1912). More recent data were obtained dur- ing direct field surveys performed by BOUCHE (1973) and TALAVERA (1992a), in particular those concerning the pres- ence of M. dubius in banana plantations and laurel forest in Tenerife. In any case, the taxonomy, ecology and choro- logy of the genus Microscolex Rosa, 1887 in the Canary Islands have been insufficiently studied. Therefore, in or- der to update and increase the data we have sampled a to- tal of 22 districts and reviewed more than 1550 specimens, producing original iconography and the first species dis- tribution maps for each island (Tenerife, La Palma, Gomera and Hierro). 2. MATERIAL AND METHODS Specimens were collected mainly by qualitative sampling along routes of up to one Kilometre in length, so as to in- clude a wide range of habitats with the greatest possible species diversity. A geologist’s hammer was used for dig- ging and specimens were picked out manually. Less fre- quently, 0.04 % formalin was poured over surface areas of 100 x 50 cm, providing complementary data on spec- imen abundance. The collected earthworms were usually transported to the laboratory inside plastic bags along with one Kilogramm of earth (winter time). After washing to remove the remaining earth they were submerged in 70 % alcohol, introduced into glass test-tubes containing 10 % formalin and left for four days, then finally con- served in 75 % alcohol. On very warm days and to avoid possible over-drying and desiccation, the worms were pre- pared and conserved “in situ” following the same steps. 38 José A TALAVERA & Dolores I. PÉREZ: Occurrence of the Genus Microscolex at Western Canary Islands In TALAVERA (1990a) keys were used to identify species. To prepare the original iconography the specimens were placed on a sheet of cork at the bottom of a beaker half- full of 70 % alcohol and the dissection was carried out un- der a binocular lens using an unsheathed razor blade and entomological pins. The specimens examined came from my own collections made during the Doctoral Thesis (1980-1986), and from surveys carried out personally be- tween 2000-2004, as part of research projects financed by TENERIFE NATIONAL PAR" FIDA HIERRO the General Directorate of Universities of the Canary Gov- ernment. A minimum of five station were sampled from each municipal district and montane localities (total 35), distributed as follows: 14 in Tenerife, ten La Palma, four Gomera and seven Hierro. These districts prospected (Fig. 1) are as follows: 1 San- ta Cruz de Tenerife, 2 La Laguna, 3 Anaga, 4 El Rosario (Monte La Esperanza), 5 Acentejo, 6 Icod, 7 Daute, 8 Los Fig. 1. Maps western Canary Islands showing of municipal district. Green, distribution of Microcolex dubius; Mallow, distribu- tion of Microscolex phosphoreus; Orange, distribution of together species. EPIA Bonner zoologische Beitráge 56 (2007) 39 Silos, 9 Candelaria, 10 Gúimar, 11 Chasna, 12 Abona, 13 Arona 14 Adeje, 15 Santa Cruz de La Palma, 16 Las Bre- ñas, 17 Mazo, 18 Fuencaliente, 19 Valle de Aridane, 20 Taburiente, 21 Puntagorda, 22 Garafía (Cueva del Agua), 23 Barlovento (Las Cancelitas), 24 Puntallana, 25 San Se- bastián, 26 Valle Gran Rey, 27 Hermigua (Cedro-Garajo- nay 28 Vallehermoso (Chorros de Epina), 29 Valverde, 30 Meseta de Nizdafe, 31 Frontera, 32 Dehesa 33 Julán, 34 El Pinar, 35 Tajace. 3. RESULTS 1550 earthworms assignable to Microscolex dubius and Microscolex phosphoreus were studied. Figs 1-3 show the main anatomical-morphological characters and the pres- ent range of distribution for each species. Microscolex dubius (Fletcher, 1887) Eudrilus dubius Fletcher, 1887 Microscolex dubius Rosa, 1890 Remarks. The morphological variability between the ex- amined specimens is minor, except in the pigmentation (different tones of pinkish grey), and body length vary- ing between 30-75 cm. Male pore and prostates tubular on 17. Genital papillae and spermathecae are generally ab- Fig. 2. Microscolex dubius. External morphology (lateral view); General dissection of the anterior region. sent, whereas small aracemose seminal vesicles nearly al- ways appear in segments 11-12. A hydrophilic species tol- erating high Na concentrations, it was found at humid sites rich in natural or artificial organic matter, and occasion- ally in soils with as little as 15.5 % relative humidity and 1.1 % organic matter (Buenavista, Tenerife). Habitat spectrum. It is common in anthropized biotopes in the basal belt below 300 m asl (gardens, parks, green- houses, tropical fruit plantations), and also often found in the medium altitude zones (400-600m) occupied by tra- ditional agriculture, and even in the native forests (Canary pine, and laurel forest). Nevertheless, it was not found in the high mountain zones where the extensive lava fields present a practically insurmountable natural barrier (Tei- de National Park, at altitudes over 2000 m). M. dubius has been found throughout the northwest of Tenerife, La Pal- ma and Hierro, being less common on south-facing slopes, mainly in areas lying leeward of the trade-winds (Fig. 1). The presence of this species in Tenerife is note-worthy over deforested or tree-covered biotopes on northern slopes, whereas 1t diminishes gradually towards the south, which explains the smaller number of specimens found there, due to the dry gritty soil, as well as to the tremen- dous impact of urban expansion associated with tourism. This is repeated on the south slopes of La Palma, and Hi- erro, where the habitat range of M. dubius is restricted to valleys and ravines, gardens, and avocado, tomato and ba- nana plantations. It also inhabits the deep gorges running through Garajonay National Park (Gomera), and remote areas of Taburiente National Park (La Palma), where it found underneath stones and rotting trunks together with Dendrodrilus rubidus and Eisenia eiseni. Chorology. TENERIFE: Santa Cruz—La Laguna metro- politan area; above Las Mercedes laurel forest and La Es- peranza pine forest; north-northwest slopes from the towns of Acentejo to Icod, Daute and Santiago del Teide; south to southwest slopes from the Malpais of Cande- laría-Valle de Gúimar, to Chasna, Abona and Adeje. LA PALMA: Metropolitan area of Santa Cruz, rural center of Mazo, and Las Breñas; northwest districts between the dis- perse evergreen laurel forest of Cueva del Agua and conifers around Puntagorda; northeast districts from the antropized zones of Puntallana to the undisturbed laurel forest of Cancelitas; villages of Valle Aridane, and dis- persed pine forest in the southern district of Fuencaliente. GOMERA: Woodlands on the central plateau of El Ce- dro; agricultural areas on the northwest slopes (Valle- hermoso and Valle Gran Rey. HIERRO: Metropolitan area of Valverde; deforested soils on north slopes between Nizdafe; El Pinar (Barranco de la Vieja, 28RBR0667)); wide forested bands in Las Dehesas, Julán, and Tajace. 40 José A TALAVERA & Dolores I. PÉREZ: Occurrence of the Genus Microscolex at Western Canary Islands nr NS Ll = Fig. 3. Microscolex phosphoreus. External morphology (late- ral view); General dissection of the anterior region. Microscolex phosphoreus (Duges, 1837) Lumbricus phosphoreus Duges, 1837 Microscolex phosphoreus Bouché, 1972 Remarks. The body length between two and 5.5 cm (av- erage 3.4) is evidence that the Canary island specimens are smaller than those described from continental areas. In addition, it differs from M. dubius in its smaller size, genital papillae in segment 11, penial chaetae present in 17, and a pair of diverticulate pyriform (pear-shaped) sper- mathecae in segment 9. M. phosphoreus is recorded for the first time from Hierro. Habitat spectrum. We consider M. phosphoreus a species indicative of agricultural activity, whose pattern of habitats is tropical fruit crops by drip-feed irrigation and cultivated areas subject to the use of fertilizers. It was oc- casionally found in gardens with exotic plants and defor- ested sites up to 1500 m, sharing ecological niches with M. dubius. This species has a fragmentary distribution clearly related to human action; it is present on all the western Canary Islands, Gomera contributes the least num- ber of specimens whereas Tenerife the greatest number, mainly in the northern zone in banana and avocado plan- tations. It is quite scarce in the South, where it arrived accidentally in soil transported from the northern slope to improve the ground used for tropical fruit and tomatoes. In La Palma, M. phosphoreus is noteworthy in some hor- ticultural areas near the port and a few rural towns and villages in the basal belt, where produce for export is grown by irrigation. On the other hand, this species prefers the pineapple, banana and avocado plantations at the northeastern end of Hierro, whereas its presence on Gomera is restricted to the bay of San Sebastian and its suburbs, which suggests recent colonization and incipient populations of M. phosphoreus with little capacity to ex- pand towards other sites. Chorology. TENERIFE: Parks in Santa Cruz; Botanical Garden above Puerto de la Cruz; anthropized biotopes on the north slopes (from La Victoria of Acentejo to Los Si- los), and south to southeast (Arona - Adeje). LA PALMA: Around the port of Santa Cruz de la Palma; vegetable gar- dens in Las Brenas districts; and fragmentedly in the Valle Aridane area; Barlovento (northeast); Garafia, (northwest). GOMERA: Metropolitan area of San Sebastian. HIERRO: cultivated plots with non-native crops in Valverde; adja- cent hamlets with farming tradition; Dehesa (northwest); southeast of El Pinar (Barranco de la Vieja, 23RBR0667). 4. DISCUSSION AND CONCLUSIONS This new contribution, as did the earlier study (TALAVERA 1990b, 1992b), adds to the knowledge of these earthworms widely distributed over tropical America and Africa. The family Acanthodrilidae Claus, 1880, is made up of more than twenty genera (Sims 1980), of which Microscolex has been able to reach the western Canary Islands but with un- equal success, the seaports being the main entry point of M. dubius and M. phosphoreus. This latter species was in- troduced by man in recent times, perhaps from the 18! century on, as a result of the increase in foreign trade and imports of exotic plants from the Caribbean colonies and Latin America. However the colonizing process of M. du- bius may have begun in more remote times, as shown by its presence deep within the native forests in the Canaries. In any case, both peregrine species initially used empty niches free of predators and native parasites, which favoured the installation of pioneer individuals in a short period of time. This hypothesis is in agreement with GATES (1972) for European lumbricids transported to America by man. Fig. 1 reveals some differences at the chorological level, for example that M. dubius is more abundant than M. phosphoreus. In fact, the latter is restricted to tropical fruit plantation parks and gardens with exotic plants im- ported from America. The habitat spectrum of M. dubius is wider, it was even found in the native Pinus canariensis and evergreen lau- rel forests, where it may constitute independent popula- tions, or otherwise occasionally share its ecological niche with epigeous lumbricids or the endogeous species Al- Bonner zoologische Beiträge 56 (2007) 4] lolobophora chlorotica Savigny, 1826 belonging to the an- cient fauna of the Canaries. Indeed, no indications of in- terspecific competition among them were detected, which concords with what ABBOT (1982) reported for Western Australia. We suggest three reasons why M. dubius has prospered more than M. phosphoreus: ecological plastic- ity, mode of transportation, and the colonization period; since M. dubius has been able to adapt to a greater diver- sity of habitats, use other dispersion routes besides the merely human one, and its colonization process began long ago, as testified by finding it in montane zones with dif- ficult access. All this is compatible with the possible ad- vantage that would be provided by parthenogenesis in re- cently colonized sites, where the probability of finding a mate is less. KRAEPELIN (1895) in his naturalist study cites Mi- croscolex poulteni Beddard, without mentioning number of specimens, locality, or which island. However, all the collected indications and data lead us to remove it from the present Canary Islands catalogue of valid species, as well as from any list of species “to be confirmed’ or con- sidered as extinct, simply because it did not exist nor has it been introduced into the Canary Islands. We suspect that the researcher K. Kraepelin, inexpert in earthworm tax- onomy performed an imprecise identification, perhaps due to the similarity between M. poulteni and M. dubius. Be- sides this, it is unlikely that this latter peregrine species could have been passed by unnoticed, since it is the only acanthodrilid with a wide distribution in the Canary Is- lands. In general the Microscolex genus 1s scarcely rep- resented in the Atlantic islands, which contrasts with its wide continental presence. In particular, when comparing our results with those obtained in Madeira (TALAVERA 1996), the same is confirmed (only two species: M. du- bius and M. phosphoreus), showing similarities between the habitat preferences and distribution range of both. The colonization process in the Canaries and Madeira proba- bly followed the same pattern, mainly as regarding the role of human activity and the routes along which these acan- thodrilids became dispersed over the basal and montane belts, where we consider that the easy altitudinal move- ments of bird flight were one of the possible modes of transport of cocoons-larvae (adhered to legs or plumage). This hypothesis is supported by other authors, among them TERHIVUO (1988) suggested it as a pathway in the colo- nization of oceanic islands. REFERENCES ABBOT, I. 1982. The distribution of earthworms in the Perth Met- ropolitan Area. records Western Australian Museum 10 (1): 11-34. BALDASSERONI, V. 1912. Nuovo contributo alla conoscenza dei lombrichi italiani. Monitore Zoologico Italiano 23 (6-7): 141-148. BEDDARD, F. E. 1895. A Monograph of the Order of Oligochaeta, Oxford, Clarendon Press 769 pp. BouchHÉ, M. B. 1973. Observation sur les lombriciens (4éme se- rie: X,XI). Prospection de l’ile de Tenerife: Lumbricidae et Acanthodrilidae. Revue d’ Ecologie et de Biologie du Solio 10 : 327-336. COGNETTI DE MARTIIS, L. 1906. Contributo alla conoscenza de- lla drilofauna delle Isole Canarie. Bollettino dei Musei di Zoo- logía ed Anatomia comparata della Universitá di Torino 21 (521): 14. Csuzpl, Cs. & SZLÁVECZ, K. 2002. Diplocardia patuxentis sp.n., a new earthworm species from Maryland, North America (Oli- gochaeta: Acanthodrilidae). Annales Historico-Naturales Mu- seo Nationalis Hungarici 94: 45-51. FRAGOSO, C. 1988. Sistemática y ecología de un género nuevo de lombriz de tierra (Acanthodrilini, Oligochaeta) de la sel- va Lacandona, Chiapas, Mexico. Acta Zoologica Mexicana 25:1-41. FRAGOSO, C. & Rojas Fernández, P. 1994. Earthworms from southeastern Mexico. New Acanthodrilidae genera and species (Megascolecidae, Oligochaeta). Megadrilogica 6 (1): 1-12. GATES, G. E. 1972. Contributions toward a revision of the earth- worm familyLumbricidae. IV. The trapezoides species group. Bulletin Tall Timbers of Research Station 12: 1-146. KRAEPLEIN, K. 1895. Zoologische Ergebnisse einer Frühjahrs- Exkursion nach Madeira und den Canarischen Inseln. Ver- handlungen des naturwissenschaftlichen Vereins Hamburg Hamburg 2: 6-17 MICHAELSEN, W. 1900. Oligochaeta. Das Tierreich 10: 1-575. REYNOLDS, J. & RIGHI, G. 1994. On some earthworms from Be- lize, C.A. with the description of a new species (Oligochaeta, Acanthodrilidae, Glossoscolecidae and Octochaetidae). Megadrilogica 5 (9): 97-106. Sims, R. W. 1980. A classifications and the distribution of earth- worms, suborder Lumbricina (Haplotaxida: Oligochaeta). Bul- letin of the British Museum Natural History (Zoology) 39 (2): 103-124. TALAVERA, J. A. 1990a. Claves de identificación de las lombrices de tierra (Annelida: Oligochaeta) de Canarias. Vieraea 18: 113-119. TALAVERA, J. A. 1990b. Considerations about Ocnerodrilus oc- cidentalis (Oligochaeta: Ocnerodrilidae) in the Canary Is- lands). Bonner zoologische Beitráge 41 (1): 81-87. TALAVERA, J. A. 1992a. Earthworms of the banana groves from Tenerife (Canary Islands). Soil Biol.ogy & Biochemistry 24: 1369-1375. TALAVERA, J. A. 1992b. Octochaetid earthworms of the Canary Islands. Bonner zoologische Beitráge 43 (2): 339-348. TALAVERA, J. A. 1996. Madeira earthworm fauna. Italian Jour- nal of Zoologia 63:8 1-86 TERHIVUO, J. 1988. The finnish Lumbricidae fauna and its for- mation. Annales Zoologici Fennici 25: 229-247. Zıcsı, A. & Csuzoı, Cs. 1991. Der erste Wiederfund von Zapo- tecia amecamecae Eisein, 1900 aus Mexiko (Oligochaeta: Acanthodrilidae). Miscellanea zoologica Hungarica 6: 31-34. Received: 21.05.2007 Revised: 15.11.2007 Accepted 22.11.2007 Corresponding Editor: B. Huber Bonner zoologische Beitráge | Band 56 (2007) Heft 1/2 Seiten 43-48 Bonn, März 2008 Biting Midges of the Genus Palpomyia Meigen (Diptera: Ceratopogonidae) in India Sujit K. Das Gupta!), Abhijit MAZUMDAR?) & Prasanta K. CHAUDHURI) Department of Zoology, Presidency College, Calcutta, India 2)Department of Zoology, University of Burdwan, Burdwan, India Abstract. The Indian species of Palpomyia Meigen are revised with the addition of two new species, P. pseudorivularis and P. simplitheca and two new records of P magna Tokunaga and P. stella Tokunaga previously described from New Guinea. The species P. flexidigita Sinha et al. and P. novita Saha & Das Gupta are proposed as new synonyms of P cras- sipalpis Sinha et al., and P. barbara Saha & Das Gupta is considered here as a new synonym of P deminutipalpis Sin- ha et al. Keywords. Diptera, Ceratopogonidae, Palpomyia, new species, synonyms, India. 1. INTRODUCTION Palpomyia was established as a genus by MEIGEN (1818) (as a synonym of Ceratopogon Meigen) with the single type species, the European Ceratopogon flavipes Meigen. EDWARDS (1926) divided the genus into three groups on the basis of the number of spines and the degree of swelling of the fore femora. GROGAN & WIRTH (1975) rec- ognized four distinct groups based on a combination of morphological features. The larvae of Palpomyia are an important source of food for fishes preying on zoo benthos (REMM 1976) and are utilized as indicators of water quality on account of their response to certain types of pollution (GROGAN & WIRTH 1979). According to BORKENT & WIRTH (1997) and up- dated by BORKENT (2008), there are 264 extant species, including nine Indian species: Palpomyia albiditarsis Ki- effer 1910, P himalayae (Kieffer, 1911), P. leucopogon Kieffer 1911, P rivularis Kieffer 1911, P crassipalpis Sin- ha, Das Gupta & Chaudhuri 2003, P deminutipalpis Sin- ha, Das Gupta & Chaudhuri 2003, P flexidigita Sinha, P. barbara Saha & Das Gupta 2005, and P. novita Saha $ Das Gupta 2005. This paper presents a revision of the Indian species of Palpomyia including two new species and new records of two species previously known only from New Guinea. Palpomyia flexidigita and P. novita are proposed as new synonyms of Palpomyia crassipalpis and P. barbara as a new synonym of Palpomyia deminutipalpis. Thus, the to- tal number of valid species of Palpomyia in India adds up to ten. 2. MATERIAL AND METHODS The insects were collected with light traps at different lo- calities in West Bengal, India. They were cleared and mounted in a mixture of phenol and Canada Balsam on microscope slides after appropriate orientation. Morpho- logical terminology used in the species descriptions most- ly follows GILES & WIRTH (1984) and Bose et al. (2003). Types and identified specimens are housed in the Insect collections of the Entomological Laboratory of the Uni- versity of Burdwan and will be submitted to the Nation- al Zoological Collections (NZC), Calcutta. 3. TAXONOMY Key to Indian species of Palpomyia Meigen * Kieffer types are considered lost, hence types of these species were not seen and the species names may be nom- ina dubia. 1. Wing membrane pale brown to grayish ..........2 Wine: membrane hyaline ...0..02. comes 7 2. Scutum with short anteromedially bristles and with- out tubercle Scutum without any bristles anteromedially and with tubercle 3. Male with narrow sternite IX having a deep caudo median excavation Gonostylus short, bud-like, with a blunt end SPE aeRO EEE Paste eee crassipalpis © 44 Sujit K. Das GUPTA et al.: Biting Midges of the Genus Palpomyia Meigen in India Fig. 1. A-J. Palpomyia pseudorivularis sp.n. 2. A) antenna, B) Palpus, C) Mandible, D-F) fore, mid, hind femora and tibiae, G) spines on fore femur, H) hind tibial comb, I) wing, J) spermathecae. Bonner zoologische Beitráge 56 (2007) 45 Male with a broad sternite IX, without caudomedian excavation. Gonostylus slender, with a hooked tip leia deminutipalpis S 4. Mid and hind femora with dark brown apical bands. Mid tibia with one apical spine Mid and hind femora pale, without pigmented bands. Mid tibia without apical spine 5. Palpal segment III of female with a subapical pit with two sensillae. Palpal segment V with a long api- ME ee ee implitheca Q Palpal segment III of female without sensillae. Palpal segment V with a short subapical spine Me tenis nedpasaics pseudorivularis Q 6. Female with inflated forefemur bearing 8-10 ventral SpInesion entire length .2.3..ssscscdecesercceecnssesvers magna Q Forefemur slender in both sexes, with 3 and 4 apical SPIHESSTESPECHVEIY. „iertssennansecenenhasnesaennnne stella 3,2 7. All femora of male armed with stout spines O lima ibas *leucopogon O Only fore femora armed with stout spines ........ 8 8. Female thorax shining red, bearing a pair of medio- lateralíspiules 00 demini *himalayae 2 Female thorax either dark brown or black and devoid Aspa ls ta od Se 9 9. Female dark brown with pale halter .. *albiditarsis Q Female with a deep black body and dark brown halter PE I A *rivularis Q Descriptions of the species Palpomyia crassipalpis Sinha, Das Gupta & Chaudhuri, 2003 (SmHA et al. 2003: 75). Palpomyia flexidigita Sinha, Das Gupta & Chaudhuri, 2003: 79. New synonym. Palpomyia novita Saha & Das Gupta, 2005: 63. New synonym. 9: Unknown. Material: Holotype 9, West Bengal, Panagarh Agriculture Farm, 20.vi.1990. leg. P. K. Chaudhuri. 2 7, Durgapur, 13.v11.2001, leg D.Sai; 1 9, Asansol, 18.viii.2001, leg. S. Sen. Remarks. SINHA et al. (2003) described the species from India. The midges in the present study conform fully with those of the above authors and the holotype of P. crassi- palpis. After critical study and a comparison of Palpomyia flexidigita and P. novita with P. crassipalpis it has been concluded that these species are synonymous with Palpomyia crassipalpis. In this context, it should be not- ed that Fig.1 of P crassipalpis was mistakenly printed as Fig. 3 of Sinha et al. (2003). The species may be diagnosed by the following combi- nation of character states: short palpal segments, anteri- or part of thorax hairy, 5 bristles on the scutellum, femo- ra with dark brown broad apical band, hind tibia dark brown with pale band at its apical 1/3rd, 5 forefemoral spines, 9 spines of hind tibial comb, thumb like hind tib- ial spur, grayish wing about three times longer than broad, r, 1.5 times longer than r,, aedeagus with longer recurved basal arms and slender distal arm, parameres fused and slender gonostylus ending with a bud Palpomyia deminutipalpis Sinha, Das Gupta & Chaud- huri, 2003 (SINHA et al. 2003: 79). Palpomyia barbara Saha £ Das Gupta, 2005: 62. New synonym. Q: Unknown. Material. 3 O, West Bengal, Asansol, 23.v11.2001, leg. G. Bhattacharyay;1 9, Kumardubi, 25.v11.2001, leg. D. Sai. Holotype 9, Belpahari, 8.v1.1991, leg. S. Sinha. Remarks. The present specimens conform with those of SINHA et al. (2003) in structures and the morphometrics. The diagnostic features of the species are: elongated pal- pal segment II with sensory pit, reduced palpal segment V, scattered hairs and streak on thorax, 7 fore femoral spines, hind tibial comb of 7 spines, r little longer than r,, elongated gonocoxite with broad base, short gonosty- lus with bud at the tip, triangular aedeagus having deep median excavation and heavily sclerotized parameres with fused basal arms On examination of the types and previous description of SINHA et al. (2003), it appears that Palpomyia deminuti- palpis and Palpomyia barbara are the same species and P. barbara is proposed as a new synonym of the former species due to similarities of morphological features as stated above. Palpomyia magna Tokunaga, 1966 (TOKUNAGA 1966: 130). Material: 2 9, West Bengal, Jorebanglow (2040m), 21.v11.2001, leg. U. Majumdar; 1 9, Tindharia (1320 m) 25.v11.2001, leg. S.K. Pradhan. Remarks. TOKUNAGA (1966) described the female of the species from New Guinea. In the course of the present study, three insects of the Himalayas are identified as con- forming fully to the description by TOKUNAGA (1966). The species may be diagnosed by: eyes separated as wide as one facet, 12 (6 large and 5 minute) mandibular teeth, scutellum with 4 large and many small bristles, fore fe- mur mostly with 10 spines, hind tibial comb of 8 spines, Sujit K. Das GUPTA et al.: Biting Midges of the Genus Palpomyia Meigen in India Fig. 2. A—J. Palpom via simplitheca sp.n. ?. A) flagellomeres I-XIII, B) Palpus, C) mandible, D-F) fore, mid , hind femora and tibiae, G) spines on fore femur, H) hind tibial comb, I) wing, J) spermathecae. Bonner zoologische Beiträge 56 (2007) 47 wing pale with brown apical part, r-m shorter than Y of the base of Rs and M, abdomen ochreous with pale bands on segments I-IV, dark brown bands on segments V-IX, small gland rod and two sub equal oval spermathecae. Palpomyia pseudorivularis Das Gupta, sp. n. (Figs 1A—J) Q. Head. Eyes separated completely. Antenna (Fig. 1A) with flagellomeres I-VIII yellow at bases, IX-XIII com- pletely brown with pale bases and 2x length of VII; lengths of flagellomeres XIII 8: 5: 5: 5: 5: 5: 5: 6: 10.5: 13.5: 14.5: 16: 18.5, AR (Antennal ratio) 1.65. Palpus (Fig. 1B) uniformly brown, palpal segment III with apical sensory pit, segment V elongated bearing a sub apical spine; length ratio of palpal segments I-V 3: 4: 7: 5: 10, PR (Palpal ra- tio) 3.0. Mandible (Fig. 1C) with 6 strong teeth. Thorax. Brown, very hairy with small blunt anterior tuber- cle, scutellum pale, with 8 large and many small bristles. Legs (Figs 1D—F). Coxae brown, trochanters yellow, fore femur (Fig. 1G) with 7 stout spines, mid and hind femo- ra with dark brown apical band, tibiae with dark brown basal and apical bands, mid tibia with | apical spine, tar- someres I-II light yellow, HI light brown, IV-V totally brown, mid tarsomeres I-II with 2 apical spines; length ratio of leg segments 23:19: 8: 4: 2.5; 2: 3 in fore leg, 28: Da AOA) WS: 2.5 in mid leg, 32: 27.5: 14: 7: 2.5: 2:3.in hind leg. TR (Tarsal ratio) of hind leg 2.0. Hind tibial comb (Fig. 1H) with 7 spines. Wing (Fig. 11): Wing length 2.85 (2.84-2.85, n=4), wing breadth 1.05 (n=3) mm. Membrane light brown with brown veins, R,,; ending well beyond the middle, r, 2.33 times longer than r, (21:9), costal length 2.31mm; CR (Costal ratio) 0.81. Halter brown. Abdomen. Tergites brown, cerci brown, gland rods dis- tinct. Spermathecae (Fig. 1J) large, unequal, 0.18 x 0.12 and 0.14 x 0.10 mm, and the third rudimentary distinct. Material. Holotype 9, Darjeeling (2180m) 20.x.1978, leg. Sikha Sarkar. Paratypes 2 9, Darjeeling, 12.1x.2002, leg. A. Mazumdar. Remarks. Due to its relatedness to P. rivularis Kieffer, 1911, the species is named P. pseudorivularis. It resem- bles P. rivularis Kieffer in the shape of the flagellomeres, tarsomere I and structure of the spermathecae. The color pattern of the hind tibia and the femoral spines of P. sub- spara (Coquillett, 1901) and P. tainana Kieffer, 1912 are somewhat similar to those of the new species, but the fol- lowing character states are distinctive:, dark palpal seg- ment III with an apical pit, 6 strong teeth of mandible, very hairy thorax, 8 bristles on the scutellum, dark brown col- or band of hind tibia, 6 stout femoral spines, 7 spines of hind tibial comb, length of r,, dark brown halter knob dis- tinct gland rod, structure of spermatheca and distinct rudi- mentary spermatheca. Palpomyia simplitheca Das Gupta, sp. n. (Figs 2A—J) Q. Head. Eyes separated as widely as 2 facets. Antenna (Fig. 2A) with flagellomeres I-VIII yellow basally, api- cally brown, [X—XHI brown with pale bases and almost equal or longer than VIII; length ratio of flagellomeres XS 3 52325233: Ola RI BE 18, AR 1.52. Palpus (Fig. 2B) uniformly brown; palpal segment III with a sensory pit having 2 sensillae, segment V long with a long stiff spine in addition to normal two; length ratio of palpal segments I-V 2.5: 4: 6.5: 4.5: 10, PR 3.25. Mandible (Fig. 2C) with 6 large strong teeth. Thorax. Dark brown with a few anterior hairs and a small blunt anterior tubercle, scutellum pale with 4 large and several small bristles. Legs (Figs 2D-F). Coxae brown, trochanters yellow, fore femur (Fig. 2G) with 6 short and stout apical spines, mid and hind femora with dark brown apices, tibiae with dark brown basal and apical bands, color band of hind tibia nar- row, tarsomeres I-II pale yellow, II little brown, IV-V brown; length ratio of leg segments 18.5: 16.5: 8: 3: 2: 1.5: Sim fore leg, 23: 20: 11: 3: 2: 1,5; 2.5 ın mid leg, 28: 24: 13: 5: 2: 1.5: 3 in hind leg. TR of hind leg 2.6. Hind tibial comb (Fig. 2H) with 8 spines. Wing (Fig. 21). Wing length 2.38 (2.35—2.39, n=4), wing breadth 0.84 (0.84-0.85, n=4) mm. Membrane light brown, R ¿,5 ending well beyond the middle, M, arising before r-m cross vein; r, 2.35 times longer than r,, costal length 1.93 mm. CR 0.8. Halter brown with light brown knob. Abdomen. Tergites brown, cerci brown. Spermathecae (Fig.2 J) simple, small, unequal, 0.091 by 0.067 mm and 0.076 by 0.062 mm. Material. Holotype 9, Darjeeling, 23 .VI1.1968, leg. S.K. Das Gupta. Paratypes 2 9, Darjeeling, 23, vii. 1968, leg. S. K. Das Gupta; 19, 28.v11.2001, leg. U. Majumdar. Remarks. The species is named P. simplitheca due to the simple spermatheca. In the number and nature of mandibular teeth, the wing, and the spermathecae, the species resembles the females of P flaviceps (Johannsen, 1908) and P jimmensis Tokunaga, 1966. The color pat- 48 Sujit K. Das GUPTA et al.: Biting Midges of the Genus Palpomyia Meigen in India tern of legs of the new species and of P. lineata (Meigen) appears more or less similar. It appears to be closer to P. pseudorivularis but the following combination of charac- ter states is unique: flagellomeres IX-XI brown at base, yellow at the apices and almost equal in length to fla- gellmere VIII, 6 large teeth of the mandible, thorax with a few anterior hairs and a small blunt anterior tubercle, 8 large bristles and several small bristles on scutellum, col- or pattern of femora and tibiae, narrow color band on hind tibia, 8 spines on hind tibial comb, length of r,, and a sim- ple, small spermatheca Palpomyia stella Tokunaga, 1966 (TOKUNAGA 1966: 136). Material. 3 Y 2 9, Burdwan, 17.v1.1999, leg. P.K.Chaud- huri; 2 9, 2 9, Durgapur, 12.v.2000, leg. D. Sai; 1 9, Ut- tarapara, 01.v111.2001, leg. U. Majumdar; | 7, Chinsura, 21.vii.2003, leg. D. Sai Remarks. TOKUNAGA (1966) originally described this species from New Guinea and New Britain. A few biting midges in our study are identified as Palpomyia stella be- cause they appear indistinguishable. The species may be identified by the following combination of characters: small size, very pale ochreous palpus, swollen segment III, presence of conical scutal tubercle, scutellum with 4 long bristles and 15 short bristles, absence of definite col- or pattern on legs, 3 spines on fore femur, 3 spines on the hind tibial comb, pale wing with C extending beyond the end of R4,5, 1, 2.25 times longer than r, slender and ta- pered gonocoxite, gonostylus swollen arched, with 3-4 small setae, trapezoid aedeagus, fused parameres with star- like median lobe little longer than basal arms and the tip rounded, bearing barb-like hook; female abdominal ter- gites I-II with pale median linear stripes, absence of gland rods and presence of two unequal oval spermathecae. Acknowledgements. We are thankful to the Ministry of Envi- ronment & Forests, Govt. of India for funding the project of the Dipteran fauna of India and to the Heads of the Department of Zoology, University of Burdwan and Presidency College, Cal- cutta for laboratory facilities. Our sincerest gratitude owes to Dr. Art Borkent of the Royal British Columbia Museum, Canada for critical appraisal of the manuscript and for his useful suggestions for its improvement. REFERENCES BORKENT, A. & WIRTH, W. W. 1997. World species of biting midges (Diptera: Ceratopogonidae). Bulletin of the American Museum Natural History 233: 1-257. BORKENT, A. 2008. Updated World species of Biting Midges (Diptera: Ceratopogonidae). http://www.inhs.uiuc.edu/re- search/FLYTREE/Borkent.html Bose, M., Das GGUPTA, S. K., MAZUMDAR, A. & CHAUDHURI, P. K. 2003. Biting midges of the genus Atrichopogon Kieffer (Diptera: Ceratopogonidae) from India. Tijdschrift voor En- tomologie 146: 259-296. COQUILLET, D. W. 1901. New Diptera in the U.S. National Mu- seum. Proceedings of the United States National Museum 23: 593-618. EDWARDS, F. W. 1926. On the British biting midges (Diptera, Ce- ratopogonidae) Transactions of the Entomological Society of London 74: 389-426. GILES, F. E. & WIRTH, W. W. 1984. Two new species of Orien- tal biting midges (Diptera: Ceratopogonidae). Proceedings of the Entomological Society of Washington 86: 210-213. GROGAN, W. L & WirTH, W. W. 1975. A revision of the genus Palpomyia Meigen of North eastern North America (Diptera: Ceratopogonidae). Agricultural Experiment Station, Univer- sity of Maryland 5076: v+49 pp. GROGAN, W. L. & WirTH, W. W. 1979. The North American predaceous midges of the genus Palpomyia Meigen (Diptera: Ceratopogonidae). Memoirs of the Entomological Society of Washington 8: 1-125. JOHANNSEN, O. A. 1908. New North American Chironomidae. Pp. 264-285 in Felt, P.: 2314 report of the State Entomologist on injurious other insects of the State of New York, 1907. NewYork State Museum, Museum Bulletin 124: 5-541 KIEFFER, J. J. 1910. Etude Sur les Chironomides Des Indes Ori- entales avec description de quelques nouvelles especes d'Egypte. Memoirs of the Indian Museum 2: 181-242. KIEFFER, J. J. 1911. Les chironomides (Tendipedidae) de I’Hi- malaya et d'Assam. Records of the Indian Museum 6: 319-349. KIEFFER, J. J. 1912. Tendipedidae (Chironomidae) (Dipt.). Sup- plementa Entomologica 1: 27-43. MEIGEN, J. W. 1818. Systematische Beschreibung der bekann- ten europäischen zweiflügeligen Insketen 1, xxxv1+333pp., Aachen. REMM, H. 1976. A synopsis of the Palpomyia of the U.S.S.R. (Diptera, Ceratopogonidae). Loodusuurijate Seltsi Aastaraa- mat 64: 172-197. SAHA, N. C. & Das Gupta, S. K. 2005. Two New species of the genus Palpomyia Meigen (Diptera: Ceratopogonidae) from lower Damodar valley of Eastern India. Geobios 32(1): 61-64. SINHA, S., DAS GGUPTA, S. K. & CHAUDHURI, P. K. 2003. Preda- ceous midges of the genus Palpomyia Meigen (Diptera: Ce- ratopogonidae) from West Bengal, India. Proceedings of the Zoological Society, Calcutta 56(2): 75—80. TTOKUNGA, M. 1966. Biting Midges of the Palpomyiinae from New Guinea (Diptera: Ceratopogonidae). Pacific Insects, Monograph 8: 101-152. Authors’ addresses: Professors Sujit K. DAs GUPTA, De- partment of Zoology, Presidency College, Calcutta 700 073, India; Abhijit MAZUMDAR & Prasanta K. CHAUDHURI, Department of Zoology, University of Burdwan, Burdwan 713 104, India; E-Mail: chaudhuri_pk@yahoo.co.in; abhijitau@rediffmail.com. Received: 5.12.2007 Revised: 21.03.2008 Accepted: 11.04.2008 Corresponding editor: N. Dorchin Bonner zoologische Beiträge | Band 56 (2007) Heft 1/2 Seiten 49-54 Bonn, März 2009 Second Find of the Recently Discovered Amazonian Giant Peccary, Pecari maximus (Mammalia: Tayassuidae) van Roosmalen et al., 2007: First Record from Bolivia Jifi MORAVEC!) & Wolfgang BOHME?) National Museum (Natural History Museum), Prague, Czech Republic 2)Zoologisches Forschungsmuseum A. Koenig, Bonn, Germany Abstract. We report on the discovery of a large peccary in the Amazon rainforest of northwestern Bolivia in the area of “Bolpebra” close to both the Peruvian and the Brazilian border, which is documented by several photo- graphs. Its size, the relatively small head, long legs, and relatively thin and short fur without white-coloured mar- Kings leave no doubt that this is the second find of the Amazonian giant peccary, Pecari maximus, which was first discovered and described from Rio Arıpuana, Amazonian Brazil, by VAN ROOSMALEN et al. (2007). Our second re- cord for the species is at the same time the first for Bolivia, and the close geographic proximity to SE Peru ma- kes its occurrence in this country also very likely. We suggest that this unique giant peccary should serve as a flag- ship species for protecting the entire region of its potential distribution range in a frontier-crossing World Herita- ge Site. Keywords. Pecari maximus, Bolivia, Amazon rainforest. 1. INTRODUCTION Only few years ago, a giant peccary species was discov- ered in Amazonian Brazil, the fourth species in the Neotropical artiodactyle family Tayassuidae (VAN ROOS- MALEN et al. 2007). Being the largest of the four extant peccary species, it was named Pecari maximus. First mo- lecular data showed that it is the sister species of the col- lared peccary (Pecari tayacu), while the white-lipped pec- cary (Tayassu pecari) and the Chacoan peccary (Catagonus wagneri) form the sister clade of the two for- mer. The description of P maximus was one of the spec- tacular discoveries of a new large mammal species and took place only three decades after the discovery of an- other Neotropical pig, viz. Catagonus wagneri. The lat- ter species had originally been described as a fossil tax- on, and later it was discovered to survive in the Chaco of Paraguay (WETZEL et al. 1975; WETZEL 1977 a, b, 1981). This (re)discovery was long preceded by information pro- vided by Carl BERKHAN, a German farmer resident in the Bolivian Chaco, who reported to M. EISENTRAUT in 1930 on the existence of a third and even fourth peccary species in the Chaco (see EISENTRAUT 1986; BÓHME & HUTTER- ER 1999; BÖHME & STIELS 2007). Only six decades later the presence of C. wagneri was proven also in Bolivia, one decade after its initial discovery as a living taxon in Paraguay (EISENTRAUT 1986). In the case of P. maximus from Amazonian Brazil, it took only two years since its discovery before 1t was also found in a neighbouring country. This sighting was made in 2005 and is reported and discussed below. 2. COMPARATIVE DESCRIPTION OF A BOLI- VIAN SPECIMEN OF PECARI MAXIMUS WITH SOME ECOLOGICAL NOTES In January 2005, the senior author spent 16 days in a re- mote area of the Bolivian side of the triangle where Bo- livia, Peru and Brazil meet (“Bolpebra”). Close to a set- tlement named Bioceanica (11°08’S, 69°22’W), local hunters had killed an obviously young female peccary which was subsequently skinned, and some photographs were taken for documentation (Figs 1-3). The head-body length of this specimen was approximately 105 cm (the hunter’s knife in Fig. 3 measured 38 cm for comparison,). The legs were much longer than in the two sympatric species P. tayacu and T. pecari, while the head was rela- tively smaller than in females of these species. The fur was thin, leaving much of the skin visible and showing some dark mottling, but in the neck and middorsal region, the hairs were longer and darker. There was no trace ofa white collar typical for P. tayacu, nor of whitish lips as in 7. pecari (rather lighter bristles occurred in the lower part Pecari maximus > covered Amazonian Giant Peccary. Dis f the Recently g BOuME: Second Find o Moravec & Wolfgan ri 50 Ji a. Photograph J. MORAV © [an] S 3) [e] 3 © 2) = ~ 3 le Pecari maximus, shot g. 1. Young fema F Lateral view of the head of the same specimen. Photograph J. MORAVEC. Fig. 2. Bonner zoologische Beiträge 56 (2007) Fig. 3. Local hunter beginning to skin same specimen. Photograph J. MORAVEC. of the cheek area only). The brown dark mottled fur colouration differed from the strongly speckled dark black- ish-grey fur colouration of P. tayacu as well as from the blackish brown longer fur of 7: pecari. In the figures the differences in fur colour and bristle-haired texture are ev- ident although the freshly killed female pig lay in a for- est creek and was therefore wet. All characteristics given above are diagnostic for P. max- imus from Rio Aripuana in Brazil, compared with its on- ly congener P. tayacu (Fig. 4). It is characterized “in be- ing much larger but less robust, with much longer legs and a proportionally small head only slightly bigger than that of P. tayacu. Most of the body is thinly bristle-haired, over- all colour brown mixed with dirty white, a black mid-dor- sal mane running from the ears as far as the rudimentary tail. Nasal disc pinkish, relatively small and soft” (VAN ROOSMALEN et al. 2007). According to this diagnosis, the Bolivian female specimen described here belongs with- out any doubt to Pecari maximus and represents thus the second documented record for this new species. At the same time, it is the first record from outside Brazil, viz. from Bolivia. The distance between the type locality of P. maximus (left bank of the Rio Aparuana, at the mouth of its tributary Rio Arauazinho, 06°16’S, 60°20’W) is about 1000 km away from our new locality (11°08’S, 69°22’W). However, VAN ROOSMALEN et al. (2007) cited already an old report by an American rubber cutter, J. C. Yungjohann, who had lived in the Rio Xapuri area from 1906-1919 which is on- ly 80-100 km NE of the Bolivian site at Bioceanica. Sim- ilarly as the German farmer in the Bolivian Chaco men- tioned above, YUNGJOHANN also reported already one cen- tury prior to the “official”, zoological discovery on a third species of “bush pigs”: “There is a great, big one porcao, they travel in pairs, and are very lively” (YUNGJOHANN 2003). Obviously, this truly cryptozoological information is now proven by the new record of P maximus described here. Also the characterization of P maximus by Yungjohann as living in pairs fits the information given by van Roos- MALEN et al. (2007) that this species — in strong contrast to the large herds of both P. tayacu and 7. pecari (SOWLS 1984) — lives “in small family groups that consist of an adult pair with or without 1-2 offspring”. Also the young female from Bioceanica was, according to the hunters, part of a small group of only four specimens. The area of Bioceanica is characterized by undulating ter- rain, with the elevation ranging from ca. 250 to 300 m a.s.l. It is covered by tall, slightly selectively logged terra firme rain forest characterized by an abundance of Brazil nut 52 Jifí Moravec & Wolfgang BOHME: Second Find of the Recently Discovered Amazonian Giant Peccary, Pecari maximus y u, x = - Fig. 4. Live Pecari maximus from the type locality. Photograph L. FRENZ. trees Bertholletia excelsa. The upper canopy closes at the height of 30-40 m. The densest understory is associated with moist depressions along forest brooks. 3. CONCLUSION The assumption by the discoverers of Pecari maximus that it might be more widely distributed than known so far has been corroborated by the new, distant find of this species in NW Bolivia. As to its conservation status according to IUCN criteria, it must certainly be regarded as “data de- ficient”. But possible severe threats have already been list- ed by VAN ROOSMALEN et al. (2007). However, it could well serve as an impressive flagship species for protect- ing the entire interfluve range between Rio Madeira and Rio Tapajos which yielded several impressive new mam- mal discoveries and is certainly also full of other, still un- described vertebrate species. Therefore, we strongly sup- port the appeal by VAN ROOSMALEN et al. (2007) to encour- age UNESCO to urge the Brazilian and Bolivian govern- ments to declare the entire region a — frontier-crossing — World Heritage Site, including also the rain forest part of neighbouring Peru. Acknowledgements. We thank Lothar Frenz, Hamburg, for the permission to use a photograph of a live Pecari maximus spec- imen taken by him at the type locality in Brazil. The senior au- thor is indebted to James Aparicio, Collección Boliviana de Fau- na (CBF), La Paz, for continuing support during his field research trips to Bolivia. The work of JM was supported by the Ministry of Culture of the Czech Republic, project MK00002327201. The junior author thanks his two mammological colleagues at ZFMK, Bonn, Rainer Hutterer and Gustav Peters, for sharing their ex- perience with him, and for constructive comments. REFERENCES BÖHME, W. & HUTTERER, R. 1999. Leben und Werk von Mar- tin Eisentraut (1902-1994). Bonner zoologische Beiträge 48: 367-382. BÖHME, W. & STIELS, D. 2007. Totgesagte leben länger: Wieder- entdeckungen ausgestorben geglaubter Landwirbeltiere. Koenigiana 1: 21-39. EISENBERG, J. F. & REDFORD, K. H. (eds.). 1999. Mammals of the Neotropics. The Central Neotropics, vol. 3: Ecuador, Pe- ru, Bolivia, Brazil. University of Chicago Press, Chicago and London, i-x+609 pp. EISENTRAUT, M. 1986. Über das Vorkommen des Chaco-Peka- ri, Catagonus wagneri, in Bolivien. Bonner zoologische Bei- träge 37: 43-47. Sowls, L. K. 1984. The peccaries. University of Arizona Press, Tucson, 251 pp. un Ww Bonner zoologische Beitráge 56 (2007) BRAZIL Fig. 6. Map of central South America showing the distribution records of Pecari maximus. Drawing J. MORAVEC. 54 Jiri Moravec & Wolfgang BÖHME: Second Find of the Recently Discovered Amazonian Giant Peccary, Pecari maximus VAN ROOSMALEN, M. G. M., FRENZ, L., VAN HOOFT, P., DE LONGH, H. H. & Letrs, H. 2007. A new species of living peccary (Mammalia: Tayassuidae) from the Brazilian Amazon. Bon- ner zoologische Beitrage 55: 105-112. WETZEL, R. M. 1977 a. The Chacoan peccary, Catagonus wag- neri (Rusconi). Bulletin of the Carnegie Museum 3: 1-36. WETZEL, R. M. 1977 b. The extinction of peccaries and a new case of survival. Annals of the New York Academy of Sci- ence 288: 538-544. WETZEL, R. M. 1981. The hidden Chacoan peccary. Carnegie Magazin 55: 25-32. WETZEL, R. M., DuBOs, R. E., MARTIN, L. R. & Myers, P. 1975. Catagonus, an “extinct” peccary, alive in Paraguay. Science 189: 379-381. YUNGJOHANN, J. C. 2003. White Gold: The Diary of a rubber cut- ter in the Amazon 1906-1916. Synergetic Press, Santa Fe. Authors’ addresses: Prof. Dr. Wolfgang BOHME (corre- sponding author), Zoologisches Forschungsmuseum Ale- xander Koenig, Adenauerallee 160, D-53113 Bonn, Ger- many; E-mail: w.boehme.zfmk@uni-bonn.de; Dr. Jiri Mo- RAVEC, National Museum (Natural History Museum), Zoo- logical Department, Vaclavské nam. 68, C-115 79 Praha 1, Czech Republic; E-mail: jiri_ moravec@nm.cz. Received: 20.06.2008 Revised: 29.06.2008 Accepted: 10.10.2008 Corresponding editor: R. van den Elzen Bonner zoologische Beiträge Band 56 (2007) Heft 1/2 Seiten 55-99 | Bonn, März 2009 On the Phylogeny and Taxonomy of the Genus Uromastyx Merrem, 1820 (Reptilia: Squamata: Agamidae: Uromastycinae) — Resurrection of the Genus Saara Gray, 1845 Thomas M. WiLms!).4), WOLFGANG BÖHME?), Philipp WAGNER2), Nicola LUTZMANN2) & Andreas SCHMITZ?) Zoologischer Garten Frankfurt, Bernhard-Grzimek-Allee 1, D-60316 Frankfurt am Main, Germany; E-Mail: thomas.wilms@stadt-frankfurt.de; Zoologisches Forschungsmuseum A. Koenig, Adenauerallee 160, D- 53113 Bonn, Germany; 3)Muséum d’Histoire naturelle, C. P. 6434, CH-1211 Genéve 6, Switzerland: WCorresponding author Abstract. We assessed the taxonomic relationships within the genus Uromastyx Merrem, 1820 using morphologi- cal and genetic methods, resulting in the resurrection of the genus Saara Gray, 1845 for Saara hardwickii, S. as- mussi and S. loricata and in changes of the taxonomic rank of Uromastyx nigriventris, U. aegyptia leptieni and U. shobraki. A synopsis of all taxa considered to be valid is provided, including differential diagnosis, description and data on their respective distribution. A key for the species of Saara and Uromastyx is presented. Keywords. Reptilia; Sauria; Agamidae; Uromastycinae; Uromastyx; Saara; Saara hardwickii, Saara asmussi new comb.; Saara loricata new comb.; Uromastyx aegyptia leptieni new status; Uromastyx nigriventris new status; Uromastyx sho- braki new status; Phylogeny; Taxonomy; Morphology. 1. INTRODUCTION Within the Palearctic genus Uromastyx Merrem, 1820 a total of 17 species are considered to be valid by WILMS & SCHMITZ (2007) and WILMS & BOHME (2007). Some of the species respective subspecies belonging to that genus have been described quite recently (e. g. Uromastyx dis- par maliensis Joger & Lambert, 1996; Uromastyx occi- dentalis Mateo et al., 1998; Uromastyx leptieni Wilms & Bohme, 2000; Uromastyx alfredschmidti Wilms & Bohme, 2001; Uromastyx y. yemenensis Wilms & Schmitz, 2007, and Uromastyx y. shobraki Wilms & Schmitz, 2007) re- flecting a continuing scientific interest in the phylogeny and taxonomy of these animals. Uromastyx spp. are medium sized to large lizards inhab- iting the old world desert belt from North Africa to north western India. All species are either ground dwellers or saxicolous, with some species climbing occasionally on trees. Uromastyx are predominantly herbivorous, feeding on the scarce vegetation in their desert environment. Eco- logically these animals are largely limited by the availabil- ity of food and by the availability of appropriate thermal refuges. Uromastyx spp. are currently listed on Appendix II of CITES. Internationally more than 367 000 specimens have been traded legally in the pet trade between 1977 and 2005 (Knapp 2004, Wırms 2007a). But the consumption of spiny-tailed lizards in their countries of origin may be con- siderably higher due to the fact, that Uromastyx are heav- ily hunted for food and for the production of souvenirs and traditional medicine (WILMS 2007a). The main aim of the present paper is to evaluate the phy- logenetic relationships within the taxa of the genus Uro- mastyx and to establish a hypothesis of the taxonomy of this group, based on a synthesis of morphological and ge- netic characters. Taxonomic History The taxonomic history of the lizards currently assigned to the genus Uromastyx dates back to the second half of the 18th century [description of Lacerta aegyptia ForsskAL, 1775; for more detailed information on the his- tory of this taxon see WILMS & BOHME 2000 a. For a dis- cussion on the spelling of PEHR FoRSSKAL’s family name see Frits & THULIN (1984)]. The genus name Uromastyx was coined by MERREM in his work ‘Versuch eines Systems der Amphibien — Tentamen Systematis Amphibiorum’ (MERREM 1820). Of the seven species included in this first synopsis of the genus only 56 Thomas M. WILMs et al.: On the Phylogeny and Taxonomy of the Genus Uromastyx Merrem, 1820 one is belonging to Uromastyx as it is currently defined [Uromastyx spinipes (Daudin, 1802) = Uromastyx aegyp- tia (Forsskäl, 1775)]. Between 1822 and 1885 a total of five new genera (Mas- tigura Fleming, 1822; Centrocercus Fitzinger, 1843; Saara Gray, 1845; Centrotrachelus Strauch, 1863; Aporoscelis Boulenger, 1885) were erected for different members of the genus Uromastyx of which only Aporoscelis and Cen- trotrachelus were considerably in use (e. g. ANDERSON 1894, 1896, 1901; BLANFORD 1874, 1881; VON BEDRIA- GA 1879; MURRAY 1884; SCORTECCI 1933; NINNI 1933; PARKER 1942; HAAS & WERNER 1969). Aporoscelis was used in the rank of a subgenus by JOGER (1987). The name Centrocercus Fitzinger, 1843 is preoccupied by Centro- cercus Swainson, 1832 (Aves, Phasianidae) and is there- fore not available. The main taxonomic problem within Uromastyx was the proper delimitation of taxon bound- aries on the specific and subspecific level, which led in the past to considerable confusion on the identity of di- verse taxa (for more detailed information see WILMS & BÖHME 2000 a, 2000 b, 2001). Beside studies based on external morphology (e. g. MERTENS 1962; Moopy 1987; WiLms € BÖHME 2000 a, 2000 b; WILMS € BÖHME 2001; WILMS & SCHMITZ 2007) and immunology (JOGER 1987), some recent papers also adress this issue by employing molecular genetic meth- ods (AMER & KUMAZAWA 2005; WILMS & SCHMITZ 2007; HARRIS et al. 2007). Nevertheless some aspects of the tax- onomy of these highly specialized desert lizards still re- main unclear. On the basıs of external morphology and immunological distances it is well established, that several species groups within Uromastyx are recognizable, but the relationships and species compositions of these groups are still under debate (JOGER 1986; Moopy 1987; WıLms 2001; AMER & KUMAZAWA 2005; WILMS & SCHMITZ 2007). 2. MATERIAL AND METHODS Morphological sampling and analysis 621 specimens of the genus Uromastyx, including the type material of the relevant taxa have been examined. The specimens are deposited in the following collections (In- stitutional abbreviations in parenthesis): The Natural His- tory Museum, London (BMNH); Naturhistorisches Mu- seum Wien (NMW); Museo Zoologico de „La Specola“, Firenze (MZUF); Museum d Histoire Naturelle, Geneve (MHNG); Museum National d’Histoire Naturelle, Paris (MNHN); Museum für Tierkunde, Dresden (MTKD); Na- tional Museum, Museum of Natural History Prague (NMP6V); Naturmuseum und Forschungsinstitut Senck- enberg, Frankfurt a. M. (SMF); Zoologisches Forschungsmuseum A. Koenig, Bonn (ZFMK); Zoologi- sches Museum der Universität Hamburg (ZMH); Muse- um für Naturkunde, Humbold-Universität, Berlin (ZMB) and Zoologische Staatssammlung München (ZSM). For a list of examined specimens see Appendix II. For each specimen 25 external characters (16 meristic, 6 metric, 3 qualitative) have been routinely recorded: snout- vent length (SVL), length of tail (TL), head width between the anterior margins of the ear openings (HW), head length from the tip of the snout to the anterior margin of the ear opening on the left side (HL), width of tail between the 4th and 5 whorl (TW), maximum tail width at the sth whorl (TW „ax); number of tail whorls (W), number of scales beneath the 4'h toe on the left side (SD), number of gular scales (from mental to a line between the anterior margins of the ear openings (G)), number of scales around mid-body (MBS), number of scales between gular- and inguinal fold (V; ventrals), number of scales around the Sth whorl (SW), number of preanofemoral pores (PP; left and right), number of enlarged scales at the anterior mar- gin of the ear opening (LS; left and right), number of scales between suboculars and supralabials (SO; left and right), number of scales from the mid of the lower end of the ear opening to the mental scale (HS; left and right), number of scales from the upper to the lower end of the left ear opening (ES; approximately three scale rows be- fore the anterior margin of the ear opening), number of scales from the upper end of the left ear opening to the first enlarged subocular scale (PES), presence or absence of enlarged tubercular scales at the flanks (TF; absent = 0 / present = 1), enlarged tubercular scales at the dorsum (TD; absent = 0 / present = | / arranged in rows = 2), in- tercalary scales between the whorls present or absent (IS; absent = 0 / 1-2 unkeeled present = 1 / 2-6 keeled pres- ent = 2). Measurements were taken to the nearest 0.5 mm using a calliper. To obtain morphological outgroup data from the closest relatives of Uromastyx several vouchers of the genus Leiolepis from the collection of the ZFMK were exam- ined. Statistical analyses of morphological data The Excel 2000 and SPSS (10.0) statistical packages were used to run the analyses. Hierachical Cluster analysis and Principal Component Analysis (PCA) have been selected to evaluate the morphological data and to explore the phe- netic relationships between the taxa examined. Bonner zoologische Beitráge 56 (2007) 57 Phylogenetic analysis of morphological data Phylogenetic analysis was carried out on the basis of twen- ty-five external characters (16 meristic, 6 metric, 3 qual- itative). To assign a polarity to these characters (plesiomor- phy vs. apomorphy), ingroup and outgroup comparisons were applied (WATROUS & WHEELER 1981; MADDISON et al. 1984). Species of the genus Leiolepis were used as out- group, because this genus forms the morphologically and genetically proposed sister clade to Uromastyx (PETERS 1971; BÖHME 1988; SCHMITZ et al. 2001; AMER & Ku- MAZAWA 2005). Within the genus Leiolepis seven taxa are distinguished: L. belliana HARDWICKE & GRAY, 1827; L. guttata CUVIER, 1829; L. revesii GRAY, 1831; L. peguen- sis PETERS, 1971; L. triploida PETERS, 1971; L. guenther- petersi DAREVSKY & KUPRIYANOVA, 1993 and L. boehmei DAREVSKY & KUPRIYANOVA, 1993 of which three are ‘ag- amospecies’ (L. triploida, L. guentherpetersi and L. boehmei; DAREVSKY & KUPRIYANOVA 1993), which do not require fertilisation of female gametes to produce off- spring. For thirteen of the twenty-five characters polarity was unanimously assignable. These characters (ten two-state and three multistate) were defined for one outgroup (Leiolepis) and all twenty-three taxa in this study. A char- acter matrix (Table 1) was designed using Nexus Data Ed- itor (PAGE 2001) and analysed in PAUP* v4.0b10 (Swor- FORD 2002) using both neighbour-joining (NJ) and max- imum parsimony (MP) algorithms. MP was run using a heuristic search and 2000 bootstrap pseudoreplicates. De- tail of the character definition and coding is provided in Appendix III. Genetic sampling Samples of muscle tissue were taken from fresh specimens as well as from preserved specimens kept in the collec- tion ofthe ZFMK, Bonn. New voucher specimens are now also kept in the herpetological collection ofthe ZFMK and the National Museum, Museum of Natural History Prague (NMP6V) (for a complete list of voucher speci- mens see Table 2). DNA was extracted from the tissue samples using QuiAmp tissue extraction kits (Quiagen) or a modified Chelex-Protocol (WALSH et al. 1991; SCHMITZ 2003). The primers 16sar-L (light chain; 5” - CGC CTG TTT ATC AAA AAC AT - 3”) and 16sbr-H (heavy chain; 5” - CCG GTC TGA ACT CAG ATC ACGT - 3”) of PALUMBI et al. (1991) were used to amplify a section of the mitochondr- ial 16S ribosomal RNA gene. PCR cycling procedure was as described in SCHMITZ et al. (2005). To get a better resolution within two identified clades of very closely related taxa (compare below), 12S rRNA da- ta for representatives of those clades were added and sep- arate trees were produced. Therefore, in these cases we amplified a section of the mitochondrial 12S ribosomal RNA gene using the primers 12SA-L (light chain; 5” - AAA CTG GGA TTA GAT ACC CCA CTA T - 3’) and 12SB-H (heavy chain; 5” - GAG GGT GAC GGG CGG TGT GT - 3”) of KOCHER et al. (1989). Cycling procedure was again identical as described in SCHMITZ et al. (2005). PCR products were purified using Qiaquick purification kits (Qiagen). Sequences (including complimentary strands for assuring the accuracy of the sequences) were obtained using an automatic sequencer (ABI 377). Se- quences were aligned using ClustalX (THOMPSON et al. 1997; default parameters) and manually checked using the original chromatograph data in the program BioEdit (HALL 1999). For the full dataset we performed neighbour-join- ing (NJ), and Bayesian reconstructions (PP), while for the two extended dataset we also calculated maximum parsi- mony trees. We used PAUP* 4.0b10 (SwoFFORD 2002) to compute the neighbor-joining tree, maximum parsimony tree and the uncorrected pairwise distances for all se- quences. For the additional MP analysis of the combined 16S and 12S datasets, we used the heuristic search algo- rithm of PAUP* (SwOFFORD 2002) with 100 random ad- ditions per replicate and the TBR (tree bisection-recon- nection) branch swapping option. Additionally, we used bootstrap analyses with 2000 pseudoreplicates to evalu- ate the relative branch support in the phylogenetic analy- sis. For the Bayesian analysis parameters of the model were estimated from the data set using MrModeltest 2.2 (NYLANDER 2004) and the analyses were performed with MrBayes, version 3.0b4 (HUELSENBECK & RONQUIST 2001). The comparison between the different likelihood scores for each model showed that the GTR + I model (YANG 1994) was determined to be the optimal model for the data set. For the Bayesian analyses we ran two MCMC analyses for 10° generations each. The initial 100000 (10%) trees were disregarded as “burn-in”. We consider probabilities of 95 % or greater to be significantly sup- ported. The exact parameters used for the Bayesian analy- ses followed those described in detail by REEDER (2003). Sixty-four 16S sequences comprising 555 bp (lengths re- ferring to the aligned sequences including gaps) as well as thirty-two 12S sequences comprising 434 bp were ob- tained. Sequences have been submitted to GenBank; for accession numbers see Tab. 2. Tympanocryptis tetra- porophora Lucas & Frost, 1895 (Agamidae: Amphiboluri- nae), Agama impalearis Boettger, 1874 (Agamidae: Agaminae), A. planiceps Peters, 1862 (Agamidae: Agam- inae), Leiolepis b. belliana Hardwicke & Gray, 1827 (Agamidae, Leiolepidinae), L. r. reevesii Gray, 1831 Thomas M. WiLms et al.: On the Phylogeny and Taxonomy of the Genus Uromastyx Merrem, 1820 58 Tympanocryptis tetraporophora E105.13 1.00 Agama planiceps AF355476 100 Agama impalearis AIM41468 Leiolepis b. belliana AF378379 1.00 [ 99 Leiolepis r. reevesii AF378376 100 11.00 Leiolepis guentherpetersi AF378378 100 Leiolepis guttata AF378377 79 1.00 99 1.00 100 1.00 100 93 1.00 100 1.00 90 1.00 100 1.00 100 1.00 100 1.00 100 _10 Fig. 1. Cladogram of the tree recovered by the analyses based on 555 lues at the nodes are Bayesian posterior probabilities (values below 0.5 replicates (values below 50 % not shown). Uromastyx asmussi E107.5 Uromastyx loricata E107.3 Uromastyx hardwickii E Uromastyx hardwickii E Uromastyx hardwickii E Uromastyx hardwickii E Uromastyx hardwickii E meh beck ah ab a ee ee N =) 23 wd od acanthinura-group aegyptia-group Uromastyx benti E111.2 pee Uromastyx benti E106.3 100 Uromastyx benti E111.4 88 Uromastyx benti E111.13 1.00 Uromastyx yemenensis E111.12 0.93/ 100 Uromastyx yemenensis E106.18 0.96 Uromastyx yemenensis E106.19 : Uromastyx yemenensis £106.20 83 0.99 Uromastyx shobraki E111.1 99 Uromastyx shobraki E111.3 70 Uromastyx shobraki E111.6 Uromastyx shobraki E111.7 1.00 Uromastyx ocellata E106.6 100 Uromastyx ocellata E106.7 Uromastyx ornata philbyi E110.20 0.97 Uromastyx ornata E106.11 100 E Uromastyx ornata E106.8 Uromastyx ornata E106.9 Uromastyx macfadyeni E112.1 Uromastyx macfadyeni E112.3 Uromastyx princeps E106.24 Uromastyx princeps E106.25 Uromastyx thomasi E105.4 Uromastyx thomasi E105.11 Uromastyx thomasi E105.12 555 bp of the 16S mitochondrial RNA gene. Upper (bold) va- not shown); lower values are neighbor-joining bootstrap Bonner zoologische Beiträge 56 (2007) 59 (Agamidae, Leiolepidinae), L. guentherpetersi Darevsky & Kupriyanova, 1993 (Agamidae, Leiolepidinae) and L. guttata Cuvier, 1829 (Agamidae, Leiolepidinae) were used as outgroup. 16S Sequences for all species used as out- group, with the exception of 7. tetraporophora, have been obtained vom GenBank. 3. RESULTS Results of the phylogenetic analysis of genetic data Uromastyx sensu lato and Leiolepis group together in a large clade supported by a neighbour joining bootstrap val- ue of 79 (Fig. 1) and is the sister group to a clade includ- ing Agama planiceps and A. impalearis. Within this clade Leiolepis and the ingroup are separated in fully support- ed subclades (PP: 1.00 / NJ: 100 for Leiolepis; PP: 1.00 / NJ: 100 for Uromastyx sensu lato). The ingroup itself forms again two well separated clades: in the first one U. hardwickii groups with the sister species U. asmussi and U. loricata, while in the second all other taxa of the genus Uromastyx are present (Uromastyx sensu stricto). Both clades are supported at least by very high and significant NJ bootstrap values (U. hardwickii, U. asmussi, U. lori- cata clade: PP: 1.00 / NJ: 100; Uromastyx s. s.: PP: <0.95 / NJ: 93). Genetic distances (uncorrected p-distances, 16S rRNA gene) from U. hardwickii, U. asmussi and U. loricata to all other taxa are as follows: hardwickii: 10.2-14.2 %, as- mussi: 8.6-13.0 %, loricata: 8.2-12.1 %. Within Uromastyx sensu stricto five well supported clades are recognizable but the direct relationships of these clades are not resolved, as they are forming an unresolved poly- tomy. Uromastyx hardwickii, U. asmussi and U. loricata clade The clade including these three afore mentioned species shows a substructure with two principal subclades. Beside the clade consisting of the sister taxa U. asmussi and U. loricata (NJ: 78), asecond well supported clade (PP: 1.00 / NJ: 100) comprising all five U. hardwickii-specimens. This latter clade also shows another clear separation with taxa-units of three and two hardwickii-specimens respec- tively and both these terminal clades are significantly sup- ported by at least one bootstrap value (ZFMK 83794, 83795, 83797: NJ: 99; ZFMK 83796, sample without voucher specimen: PP: 1.00 / NJ: 98). We preliminarily assigned the second subcluster exclusively to Uromastyx hardwickii, but data suggest that in fact two taxa may be involved (see also discussion). Genetic distances between the taxa of the Uromastyx hard- wickii, U. asmussi and U. loricata clade are as follows: asmussi-loricata: 2.9 %; asmussi-hardwickii: 5.8-6.5 %; loricata-hardwickii: 6.1-6.7. Distance between the two identified subclades within hardwickii is rather low at 0.9%. Uromastyx sensu stricto clade Based on the genetic data four of the five clearly recog- nizable clades are strongly supported by bootstrap values: Uromastyx acanthinura group (PP: 1.00 / NJ: 99); U. ae- gyptia group (PP: 1.00 / NJ: 100); U. ocellata group (PP: 1.00 / NJ: 90) and U. thomasi (PP: 1.00 / NJ: 100). The fifth clade comprising U. macfadyeni and U. princeps is only very weakly supported. To get a better resolution within the U. acanthinura and the U. aegyptia clades 12S ıRNA data were added and separate trees were produced. Uromastyx acanthinura group Based on the genetic data the U. acanthinura clade (Fig. 1), including the taxa geyri, acanthinura, nigriventris, dis- par, flavifasciata and maliensis (alfredschmidti was not included in this analysis due to the non-availability of DNA samples), is very well supported by bootstrap val- ues (PP: 1.00 / NJ: 99). Intraspecific genetic distances within all taxa of the U. acanthinura group was 0.0-0.4 % (exception U. geyri: 0.9 %). Between the taxa of this group, genetic distances are 0.2-1.4 %. On the basis of these data, decisions on the rank of the taxa in question were not possible. To further enhance the resolution of the tree, 12S rRNA data were combined with the 16S rRNA data and new trees were produced using U. ornata as out- group (Fig. 2). The newly calculated tree shows the geyri clade basal to all other taxa within the U. acanthinura group. This clade is maximally supported (PP: 1.00 / NJ: 100 / MP: 100) and forms the sister taxon to all other members of the U. acanthinura group, which form a clade significantly supported by bootstrap values (NJ: 88 / MP: 100). This clade shows a very well supported substruc- ture with nigriventris being the sister taxon (PP: 1.00 / MP: NJ: 100 / MP: 100) of the clade including acanthinura and U. dispar spp. On the basis of this tree, acanthinura is the sister taxon to the clade comprising the taxa dispar, flav- ifasciata and maliensis with both clades being significant- ly supported by at least NJ and MP bootstrap values (acan- thinura clade: PP: 0.97/0.95 / NJ: 100 / MP: 100; dispar clade: PP: 0.75/0.80 NJ: 85 / MP: 90). As it was not pos- sible to win a 12S DNA-sequence from the only available representative of maliensis and we still wanted to include all described taxa in our analyses, we filled the missing 12S sequence information with “N”s and calculated the phylogenetic trees both with and without the inclusion of maliensis. This was done to check if the inclusion of the 60 Thomas M. WiLms et al.: On the Phylogeny and Taxonomy of the Genus Uromastyx Merrem, 1820 Uromastyx ornata E106.11 U. acanthinura acanthinura E105.21 0.97/0.95 100/100 U. acanthinura acanthinura E105.22 70/- U. acanthinura acanthinura E105.23 var U. acanthinura acanthinura E107.15 IS U. dispar maliensis E106.26 U. dispar flavifasciata E105.15 51/ | U. dispar flavifasciata E105.27 56 | U. dispar flavifasciata E106.22 | U. dispar flavifasciata "obscura" E133.2 | U. dispar flavifasciata "obscura" E133.7 | U. dispar flavifasciata "obscura" E133.8 100/98* 62/- U. dispar flavifasciata "obscura" E133.9 U. dispar flavifasciata "obscura" E133.10 U. dispar flavifasciata "obscura" E133.11 x U. dispar flavifasciata "obscura" E111.21 88/100 63 7 70/- U. dispar flavifasciata "obscura" E111.22 a U. dispar flavifasciata "obscura" E133.3 oo. a U. dispar flavifasciata "obscura" E133.6 85/90* U. dispar flavifasciata E110.8 ST U. dispar flavifasciata E110.9 1.00 U. d. dispar E106.2 U. d. dispar E110.19 U. acanthinura nigriventris E107.14 1.00 100/100 U. acanthinura nigriventris E106.4 65/- un : U. acanthinura nigriventris E106.5 1.00 U. geyri E105.24 1a9za0l U. geyri E105.25 10 Fig. 2. Cladogram of the tree recovered by the analyses based on 989 bp of the combined 16S and 12S mitochondrial RNA ge- nes. Upper values at the nodes are Bayesian posterior probabilities (values below 0.5 not shown); lower values on the right are maximum-parsimony bootstrap replicates; lower values on the left are neighbor-joining bootstrap replicates (values below 50 % not shown). The line connecting to Uromastyx dispar maliensis is dotted to incorporate the fact that we were not able to geta 12S sequence for this species and that we had to fill up the alignment with "N"s to include the species in the calculation. Bonner zoologische Beiträge 56 (2007) 61 incomplete sequence would alter the tree topologies. As this was not the case, we added the sequence and have marked its calculated position with a broken line. Within the dispar clade three fairly well supported sub- units are recognizable (for exact bootstrap values see Fig. 2), corresponding to the currently valid subspecies dispar, flavifasciata and maliensis, while the recently described “obscura”-form is included in and identical (no genetic difference) with flavifasciata. Intraspecific genetic distances within the terminal taxa (lumping data for the subspecies of U. dispar) are: acan- thinura: 0.0-0.1%, nigriventris: 0.0-0.1%, geyri: 0.4%, dispar: 0.0-0.7%. Distances between the taxa are: acan- thinura-geyri: 4.57-4.69 %, nigriventris-geyri: 4.46-4.57 %, dispar-geyri: 4.21-4.99 %, acanthinura-nigriventris: 2.0-2.3 %, acanthinura-dispar: 1.542,44 %, nigriventris- dispar: 1.72-3.08 %. Uromastyx aegyptia group The calculation for the extended dataset for the taxa of the U. aegyptia-group produced an identical topology (tree not shown) for all three algorithms, with the following struc- ture [numbers are bootstrap values (PP/NJ/MP) for the fol- lowing nodes; significant values in bold; values below 0.90 (PP) or under 50 (NJ/MP) (*) not shown]: (Uromastyx ornata), */100 (Uromastyx a aegyptia, */59/55 (Uromastyx a. microlepis, 0.94/55/52 (Uro- mastyx a. microlepis, 1.00/93/94 (Uromastyx leptieni, Uro- mastyx leptieni)))) Within taxa genetic distances are extremely low: mi- crolepis: 0.10%, leptieni: 0.13 % (for aegyptia only a sin- gle specimen was sequenced), while between the differ- ent taxa genetic difference were comparatively much high- er: aegyptia-microlepis: 0.3-0.4 %, microlepis-leptieni: 0.3-0.6 %, aegyptia-leptieni: 0.7-0.9 %. Uromastyx ocellata group The Uromastyx ocellata group constitutes a further well supported clade (Fig. 1) which is itself again subdivided: the first main clade comprises the taxa ornata (including the single specimen of philbyi) and ocellata (PP: 1.00 / NJ: 100). Both of the nominal taxa are clearly separate species-units (PP: 1.00 / NJ: 100). The second clade comprises a well supported substructure, consisting of three subclades which correspond to the taxa benti, yemenensis and shobraki (PP: 1.00 / NJ: 90). Each of these taxa is fully supported (benti: PP: 0.98 / NJ: 100; yemenensis: PP: 1.00 NJ: 100; shobraki: PP: 0.99 / NJ: 99). Intraspecific genetic difference is very low: benti: 0.0-0.4 %, yemenensis: 0.0-0.2, shobraki: 0.0-0.2, ocellata: 0.2 %, ornata (without philbyi): 0.0 %. As expected, the in- terspecific genetic distances are much higher: benti-yeme- nensis: 2.2-2.7 %, benti-shobraki: 2.2-2.9 %, yemenen- sis-shobraki: 1.8-2.0 %, benti-ocellata: 6.5-7.2 %, ben- ti-ornata: 5.8-6.3 %, yemenensis-ocellata: 7.2-7.4 %, yemenensis-ornata: 6.5 %, shobraki-ocellata: 7.0-7.4 %, shobraki-ornata: 6.5—7.0 %, ocellata-ornata (including philbyi): 3.6-4.0 %. Genetic difference between ornata and philbyi is 0.7 %. Uromastyx macfadyeni / Uromastyx princeps clade This is the only major clade (Fig. 1) which is not signif- icantly supported on its basal node; it therefore compris- es two clearly separated species units (each with PP: 1.00 / NJ: 100), whose direct relationships remain unclear. In- traspecific genetic difference is: macfadyeni: 0.0 %, prin- ceps: 0.2 %. Between those two taxa, the genetic differ- ence is 9.0-9.5%. Uromastyx thomasi clade Uromastyx thomasi forms a separate, well supported clade of its own (PP: 1.00 / NJ: 100). Intraspecific genetic difference is 0.2-0.3 %. Results of the multivariate analyses of the taxa of the genus Uromastyx A distance phenogram based on the average values of 18 characters for all taxa of the genus Uromastyx (number of taxa = 22; Uromastyx occidentalis data were not avail- able; for definition of variables see Table 3) was calcu- lated using the complete linkage method (Fig. 3). The re- sulting distance phaenogram shows two distinct main clus- ters (OTU I £ OTU II), of which one includes hardwickii, loricata and asmussi (OTU Il), while the second cluster represents all remaining taxa of the genus (OTU I). With- in this second cluster five subcluster based on phenetic similarity are recognizable — the first cluster contains ae- gyptia, microlepis and leptieni; the second ocellata, yeme- nensis, Shobraki and benti, the third dispar, maliensis, flav- ifasciata, acanthinura and nigriventris; the fourth princeps and the fifth clade contains alfredschmidti, geyri, thomasi, macfadyeni, ornata and philbyi. Because of these morphological findings based on aver- age values, we suggest, that the genus Uromastyx s.l. con- sist of two clades which are different. To further evaluate the phenetic relationships within the genus we applied a principal component analysis (PCA) on data obtained from 481 individuals (Variables: V1-V17; see Table 4). The dis- 62 Thomas M. WiLms et al.: On the Phylogeny and Taxonomy of the Genus Uromastyx Merrem, 1820 Rescaled Distance Cluster Combine ornata philbyi macfadyeni thomasi geyri alfredschmidti princeps maliensis flavifasciata acanthinura nigriventris dispar benti yemenensis shobraki ocellata aegyptia leptieni microlepis loricata hardwickii asmussi OTU | OTU Il Fig. 3. Distance phenogram resulting from cluster analysis of average values of Variable V1—V 18 (see Tab. 4) of the taxa of Uro- mastyx sensu lato (Hierarchical cluster using complete linkage, Tschebyscheff distances and z-transformation). crepancy between the total number of specimens used in this study and the number of specimens subject to the sta- tistical analysis is because of the elimination of incom- plete datasets. In the projection of the first two principal components all specimens of hardwickii cluster seperately as well as all specimens of asmussi and loricata respectively. Both clus- ters are clearly separated from all specimens of the remain- ing Uromastyx taxa (Fig. 4; for factor loadings on princi- pal components see Table 5), and correspond to the clus- ters identified as OTU I and OTU II in the hierarchical cluster. OTU II contains two clearly separated subclusters with all U. hardwickii clustering together as well as U. as- mussi and U. loricata. The finding of two phenetic clus- ters clearly outside the Uromastyx sensu stricto cluster as well as the identification of two well supported genetic clades raise the question of a polyphyletic origin of the genus Uromastyx sensu lato. To evaluate the phenetic relationships and to discriminate the species or species groups within Uromastyx sensu stricto, data of all taxa (without U. occidentalis) were sub- ject of six PCAs (Variables: V1—V15; see Table 6). Be- tween the subsequent PCAs, data of taxa clustering out- side the respective main clusters were removed. As a re- sult of this procedure seven entities containing single species or phenetically similar taxa were recovered: l. benti, yemenensis, shobraki, princeps (Fig. 5; for fac- tor loadings on principal components see Table 7) 2. ocellata (Fig. 6; for factor loadings on principal com- ponents see Table 8) 3. thomasi (Fig. 7; for factor loadings on principal com- ponents see Table 9) 4. aegyptia, microlepis, leptieni (Fig. 8; for factor load- ings on principal components see Table 10) Bonner zoologische Beitráge 56 (2007) 63 Principal component 1 Principal component 2 Fig. 4. Projection of the first two principal components from a PCA run on 481 individuals assignable to OTU 1 and OTU 2 (0 = Uromastyx sensu stricto; A = Uromastyx loricata;) O = Uromastyx asmussi; @ = Uromastyx hardwickii). 44 34 21 ocellata 14 os = o °F E i=) a E 41] >} = = 2] = = pa Ban Sl + : + — 4 -2 0 2 4 6 Principal component 3 Fig. 6. Projection of the first and third principal component from a PCA run on 354 individuals assigned to Uromastyx sen- su strico without yemenensis, benti, shobraki and princeps (@ = Uromastyx spp., A = Uromastyx ocellata). 5. dispar, flavifasciata, maliensis (Fig. 9; for factor load- ings on principal components see Table 11) 6. acanthinura, nigriventris (Fig. 10; for factor loadings on principal components see Table 12) 7. alfredschmidti, geyri, ornata, philbyi, macfadyeni (Fig. 10) Principal component 1 Principal component 3 Fig. 5. Projection of the first and third principal component from a PCA run on 431 individuals assigned to Uromastyx sen- su stricto (OTU 1) (0 = Uromastyx yemenensis, A = Uroma- styx princeps; Y = Uromastyx benti, \\ = Uromastyx shobraki; B = Uromastyx spp.). an ~ E 0] : Es thomasi S a Eu S 2 = 2 =) = ie adi : — nn 4 3 2 -1 0 1 2 3 Principal component 4 Fig. 7. Projection of the first and fourth principal component from a PCA run on 331 individuals assigned to Uromastyx sen- su strico without yemenensis, benti, shobraki, princeps and ocel- lata (@ = Uromastyx spp.; W = Uromastyx thomasi). These seven clusters are based on external similarities and therefore do not exclusively reflect phylogenetic relation- ships but also identify phenetic similarities based on ho- moplasious character states. To evaluate phenetic relation- ships within the clades identified by genetic analysis, sep- arate PCAs were applied to the data sets of the taxa. 64 Thomas M. WiLms et al.: On the Phylogeny and Taxonomy of the Genus Uromastyx Merrem, 1820 aegyptia, microlepis 3 leptieni 24 1 ~ >: 04 z v E = e: E S a 8.2 3 = in a 3 , , - 2 -1 0 1 2 3 Principal component 2 Fig. 8. Projection of the first two principal components from a PCA run on 317 individuals assigned to Uromastyx sensu stri- co without yemenensis, benti, shobraki, princeps, ocellata and thomasi (@ = Uromastyx spp., 4 = Uromastvx a. microlepis; Y = Uromastyx a. aegyptia; @ = Uromastyx a. leptieni). Uromastyx acanthinura group The taxa of the U. acanthinura group cluster in three sub- sequent PCAs (PCA 5, 6 & 7). This indicates, that the mor- phology of the taxa of this group is to some degree dif- ferent to the other species of the genus (see also discus- sion regarding cluster 7 also containing taxa not belong- ing to the U. acanthinura group). PCAs carried out exclusively on the data of the U. acan- thinura group revealed, that U. gevri and U. alfredschmidti cluster outside of the remaining taxa. Separation of acan- thinura, nigriventris, dispar, flavifasciata and maliensis by means of PCA was not possible (data not shown; Vari- ables: VI-V15). Uromastyx aegyptia group All taxa in this study belonging to this group cluster in one single PCA (PCA 4). Phenetical relationships within the taxa of the U. aegyptia group (excluding U. occiden- talis) have already been assessed by WiLms € BÖHME (2007). Analysis revealed that male specimens could be assigned according to the a priori specimen classification using cluster analysis and PCA. For females taxon dis- crimination was not possible. Uromastyx ocellata group The taxa of the U. ocellata group are included in several clusters of the previous PCAs (PCA 1, 2 & 7). This indi- ~ = Y E e Bt] dispar, maliensis, = flavifasciata o - = 2] = = u a 21 a 3 2 1 0 1 2 3 4 Principal component 2 Fig. 9. Projection of the first two principal components from a PCA run on 265 individuals assigned to Uromastyx sensu stri- co without yemenensis, benti, shobraki, princeps, ocellata, tho- masi, aegyptia, microlepis and leptieni (i = Uromastyx spp., 4 = Uromastyx dispar maliensis; + = Uromastyx d. flavifasciata, Y = Uromastyx d. dispar). acanthinura, nigriventris nm E o 5 14 a E 3 | ornata, philbyi, = * | macfadyeni el alfredschmidti, 3 2 4 0 i g > Principal component 2 Fig. 10. Projection of the first two principal components from a PCA run on 223 individuals assigned to Uromastyx sensu stri- co without vemenensis, benti, shobraki, princeps, ocellata, tho- masi, aegyptia, microlepis, leptieni, maliensis, dispar and fla- vifasciata (M = Uromastyx spp., A = Uromastyx acanthinura; v = Uromastyx nigriventris). cates, that some taxa of this group are readily distinguish- able from other taxa of the genus (see also discussion re- garding cluster 1 £ 7 containing not only taxa belonging to the U. ocellata group). Bonner zoologische Beitráge 56 (2007) 65 A PCA carried out only on data of the specimens belong- ing to the U. ocellata group (data not shown) revealed, that two taxa cluster completely separate (ocellata, ben- ti) while shobraki and yemenensis form a common clus- ter as well as ornata and philbyi (Variables: V1-V15). Uromastyx macfadyeni / Uromastyx princeps clades Both species cluster in different PCAs (No. 1 £ 7) and are clearly separated in the PCA carried out solely on da- ta of both species (data not shown; variables: V1—V15). Uromastyx thomasi group U. thomasi already clustered completely separate in one of the subsequent PCAs applied to data of the whole genus (PCA 3). Results of the phylogenetic analysis of morphological data A NJ analysis was carried out (tree not shown) and a MP heuristic parsimony analysis resulted in 254 shortest trees (L: 254, CI: 0.762, RI: 0.857, RC: 0.653) (not shown) whose 50% majority-rule consensus shows the following structure [numbers are bootstrap values (NJ/MP) for the following nodes; significant values in bold; values under 50 not shown *]: (((((acanthinura,(((aegyptia, microlepis)70/56, lep- tieni)61/64, occidentalis)56/64, alfredschmidti,(benti, macfadyeni, ocellata,(ornata, philbyi)55/*, shobraki, yemenensis)53/71, dispar, flavifasciata, geyri, maliensis, nigriventris)56/* (princeps, thomasi)100/98)70/71,(as- mussi, loricata)94/*)53/*, hardwickii)100/100, Leiolepis) Even though the resolution of the phylogenetic analysis of the morphological data is not surprisingly rather lim- ited, the node separating Leiolepis from Uromastyx sen- su lato is fully supported by bootstrap values (100/100), and more importantly the node separating Uromastyx sen- su stricto from U. hardwickii, U. loricata and U. asmus- si is also well supported in both analyses (70/71). Synthesis and discussion of the morphological and ge- netic results As pointed out by AMER & KUMAZAWA (2005) the rela- tionship between Leiolepis and Uromastyx has been sub- ject to scientific discussions. Based on morphology both genera possess autapomorphies supporting the monophy- ly of this clade within the Acrodontia and their position as the sister taxon to all remaining agamids (Moopy 1980; BÖHME 1982). Studies based on molecular data sets failed to support this monophyly (Macey et al. 1997, 2000) or did not place this clade as the sister taxon of the remain- ing agamids (HONDA et al. 2000). We used members of the Agaminae (Agama planiceps, A. impalearis) and Am- phibolurinae (Tympanocryptis tetraporophora) as out- groups in our analysıs and found a weakly supported monophyly of the clade consisting of Leiolepis and Uro- mastyx. This result is consistent with the phylogeny es- tablished by AMER & KUMAZAWA (2005) also based on mtDNA. ANANJEVA et al. (2004, 2007) integrated morphological and molecular data and established a classification of agamid lizards by distinguishing six monophyletic line- ages on subfamily level: Uromastycinae Theobald, 1868; Leiolepidinae Fitzinger, 1843; Amphibolurinae Wagler, 1830; Hydrosaurinae Kaup, 1828; Draconinae Fitzinger, 1826; Agaminae Spix, 1825. We follow this concept of ANANJEVA et al. and regard the Leiolepidinae and Uro- mastycinae as separate lineages. Our observations based on morphological and genetic da- ta show a clear and well supported substructure within Uromastyx 8.1. Both of these entities warrant recognition on genus level. For the clade comprising the taxa of the irano-turanian subregion (hardwickii, asmussi, loricata) the genus name Saara GRAY, 1845 1s available. We there- fore resurrect Saara as the sister genus of Uromastyx. Af- ter the resurrection of the genus Saara for the species of the irano-turanian region, two genera are now placed with- in the Uromastycinae: Saara and Uromastyx. After the ex- clusion of the species of the genus Saara, Uromastyx is now monophyletic comprising 20 nominal taxa. An early separation of hardwickii from the other species of the genus Uromastyx was already proposed by JOGER (1986) based on immunological distances and AMER & KUMAZAWA (2005) based on molecular data. JOGER (1986) furthermore established a close phylogenetic rela- tionship between hardwickii and loricata. This author sug- gested that Uromastyx should be divided into several sub- genera (one of them being the clade of hardwickii and lor- icata), but did not impose formal taxonomic changes with the exception of the resurrection of the name Aporoscelis for the two broad tailed species (U. thomasi and U. prin- ceps). As Moopy (1987) pointed out, applying this con- cept would have caused the genus Uromastyx to be para- phyletic. The separation between Saara hardwickii and the species of the Afro-Arabian radiation of Uromastyx was estimated at 25—29 Mya (AMER & KUMAZAWA 2005) which is in general accordance with the estimates made by JOGER (1986). Within Saara a clear substructure is rec- ognizable with S. asmussi and S. loricata forming sister clades which are themselves the sister taxa to S. hard- wickii. 66 Thomas M. WiLms et al.: On the Phylogeny and Taxonomy of the Genus Uromastyx Merrem, 1820 The situation and relationships within Uromastyx are not as clear as in Saara. Genetically, we recognize five species groups within the genus of which four are at least partly supported by morphological data (U. acanthinura group, U. aegyptia group, U. ocellata group, U. thomasi group). The remaining group (cluster containing U. princeps and U. macfadyeni), feebly recognized on the basis of the mo- lecular data set, is not supported by morphological data. Differences in the composition of genetically based clus- ters and morphologically based groups might mainly be the result of a convergent evolution of the taxa involved due to similar ecological or climatic environments. Well supported by morphological analysis are the U. ae- gyptia and the U. thomasi groups. While U. thomasi clus- ters completely separate in the PCA analysis, all taxa of the U. aegyptia group cluster together according to the ge- netic results (hierarchical cluster, PCA analysis, PAUP analysis of morphological data). It is therefore well estab- lished, that U. thomasi and the U. aegyptia group form phylogenetic entities of their own. This is especially re- markable for U. thomasi, because this species has in for- mer studies been placed in a clade together with U. prin- ceps (JOGER 1986; Moopy 1987; WILMs 2001) with which it also clusters in the PAUP analysis of morphological da- ta (this study). The present study is the first including DNA samples of both broad tailed Uromastyx species and there- fore recovers a biased morphological interpretation in the phylogenetic relationship of these two taxa. The overall similarity between U. thomasi and U. princeps is most possibly based on the extraordinary short tail in those taxa which was misinterpreted as an autapomorphy for this group instead of an independantly evolved analogous char- acter state. From our point of view the phylogenetic af- filiation of U. princeps and U. macfadyeni is probable, though this is not conclusive due to the low bootstrap val- ues (PP: 0.77 / NJ: 51). AMER & KUMAZAWA (2005) found a sister group relationship of U. macfadyeni with species of the U. acanthinura clade (U. geyri, U. acanthinura, U. dispar), which we cannot confirm based on our own da- ta. Nevertheless the relationships within the U. acanthinu- ra group in this previous (AMER & KUMAZAWA 2005) and in the present study are in good accordance. Within the North African Uromastyx acanthinura group seven taxa are recognized, of which all but U. alfred- schmidti were available for genetic analysis. Based on 12S and 16S rRNA data geyri is the sister taxon of the clade comprising acanthinura, nigriventris and U. dispar ssp. As reported earlier nigriventris is the sister taxon of the two remaining taxa, which form themselves strongly sup- ported clades. Morphologically acanthinura and nigriventris as well as dispar, flavifasciata and maliensis form clusters in sub- sequent PCA analysis. These taxa cluster in the S'h and 6th PCA cycle respectively. All specimens of geyri and al- Jredschmidti remained in a cluster together with ornata, philbyi and macfadyeni, for which a further resolution was not possible based on the PCA methodology. It is evident, that the assingnment of geyri and alfredschmidti to the three taxa mentioned above is because of a superficial morphological similarity within the taxa in question, which is due to similar ecological adaptations (convergent evolution) and not due to phylogenetic relationships (all of them are predominantly rock dwelling species). Another PCA was carried out exclusively on specimens belonging to the taxa of the U. acanthinura clade. In this PCA geyri and alfredschmidti clustered together and out- side of the remaining taxa, with only a very small area of overlap between the respective clusters (data not shown). It was not possible to separate the remaining taxa with a further PCA. On the basis of the morphological data we consider dis- par, flavifasciata and maliensis as being closely related entities, as well as geyri and alfredschmidti. The taxa acan- thinura and nigriventris show a certain morphological similarity, which led in the past to the conclusion to tread both taxa as subspecies of a single species (WILMS & BOHME 2001). We suppose, that the U. acanthinura clade in North Africa represents a relatively recent radiation within the genus (see also WILMS 2001). This hypothesis is supported by the relatively low level of genetic difference within all taxa of this group, which is generally between 0.0 and 1.4 % difference in the 16S rRNA gene (within Uromastyx, on- ly one further group shows a similarily low degree of sep- aration: the U. aegyptia group) as well as the overall sim- ilarity concerning scalation characters. By including da- ta for the 12S rRNA gene the resolution of taxa discrim- ination was significantly enhanced, resulting in genetic distances suitable to distinguish between the taxa involved. As has been shown earlier in this study, acanthinura as well as nigriventris exhibit a very low intraspecific genet- ic distance of 0.0-0.1 %, while dispar shows a respective distance up to 0.7 %. We have therefore assessed the in- ternal distances within the nominal taxa dispar, flavifas- ciata and maliensis, which proved to be: dispar 0.0 %, flavifasciata 0.0-0.2 % and maliensis 0.0 (only one se- quence available). The respective distances between those taxa are: dispar-flavifaciata: 0.53-0.74 %, dispar-malien- sis: 0.58 %, maliensis-flavifasciata: 0.39-0.58 %. We therefore recognize dispar, flavifasciata and maliensis as valid taxa belonging to one species, Uromastyx dispar, but being differentated on subspecific level. Bonner zoologische Beiträge 56 (2007) 67 To evaluate a further taxonomic problem, we have includ- ed several melanistic specimens of flavifasciata from northern Mauritania in this study. These animals have been described as Uromastyx flavifasciata obscura by MATEO et al. (1998), and the validity of this taxon was under de- bate ever since (WILMS & BÖHME 2001; GENIEZ et al. 2004). The genetic difference between these animals and typical U. dispar flavifasciata 1s 0.0-0.2 %. We therefore consider obscura to be synonymous with flavifasciata (see also WILMS & BÖHME 2001). As a synthesis of our morphological and molecular data we consider five evolutionary entities within the U. acan- thinura group as valid on specific level: U. alfredschmidti, U. geyri, U. acanthinura, U. nigriventris and U. dispar. This result is in general accordance with the results of AMER & KUMAZAWA (2005) and HARRIS et al. (2007). The second group within the genus Uromastyx compris- ing several nominal taxa and only showing a weak mor- phological and genetic differentiation is the Uromastyx ae- gyptia group. Within this group four nominal taxa are known: aegyptia, leptieni, microlepis and occidentalis. We hypothesize that the origin of the U. aegyptia group is Afrıca and that the Arabian radiation of this group has on- ly recently dispersed into the Arabian Peninsula. The clari- fication of the evolutionary scenario of the U. aegyptia group would require the incorporation of U. occidentalis in the genetic analysis and the resolution of the relation- ships between all identified species groups. Á sister group relationship between the U. acanthinura and the U. ae- gyptia group as postulated on the basis of morphological data (Moopy 1987; WiLms 2001; this study) would bring the groups together, which represent the most recent evo- lutionary lineages. Despite the overall similarity of the taxa of the U. aegyp- tia group, it is possible to differentiate between them on the basis of morphological characters (WILMS & BÖHME 2001, 2007). Genetically, they exhibit the following in- tertaxon distances: aegyptia-microlepis: 0.3-0.4 %, mi- crolepis-leptieni: 0.3-0.6 % and aegyptia-leptieni: 0.7-0.9 %. These p-distances based on 12S and 16S rRNA are very low compared to those between Uromastyx species in gen- eral, but are similar to those shown by the taxa assigned to U. dispar as subspecies in the present study. We there- fore recognize Uromastyx aegyptia as a polytypic species with three subspecies (aegyptia, leptieni, and microlepis). Because of the significant geographic distance between the Arabian U. aegyptia and the African U. occidentalis we suppose, that both are good species. The results of the analysis of morphological as well as mo- lecular data for the U. ocellata group have been published elswhere (WILMS & SCHMITZ 2007). This group consists of six taxa which represent five evolutionary entities: ben- ti, yemenensis, shobraki, ocellata, ornata. In the context of the current data, we recognize the subspecies of U. yemenensis as valid at specific rank because of the in- traspecific genetic distances which are similar between all taxa of the subclade comprising benti, yemenensis and shobraki. 4. TAXONOMY DEFINITION AND RESURRECTION OF THE GENUS SAARA GRAY, 1845 1845 Saara Gray, Cat. Spec. Liz. Coll. brit. Mus.: 262. — Type species: Uromastyx hardwickii GRAY, 1827 Original definition: Head very short, broad, much arched. Body depressed, with a fold on each side of the back. Scales minute, equal. Tail short, broad, depressed; upper part with cross bands of compressed, conical scales, sep- arated by other rings of granular and smooth square scales; beneath covered with square, smooth, imbricate scales. Femoral pores distinct (GRAY 1845). Diagnosis: Acrodont dentition, with the premaxillary bone forming in adult specimens a sharp, tooth- like structure replacing the incisive teeth. Tail scalation arranged in dis- tinct whorls, which are separated by 1-6 rows of inter- calary scales dorsally. Species: Saara asmussi, S. hardwickii, S. loricata. Distribution: The species of the genus Saara are distrib- uted in eastern Iraq, Iran, Afganistan, Pakistan and north- western India. Taxonomy: As shown in the present study, Saara hard- wickii represents most probably a polytypic species, whose taxa are genetically distinct. Further study on the taxon- omy of Saara hardwickii 1s required to evaluate the dis- tribution and morphological characters of the taxa in- volved. SYNOPSIS OF THE SPECIES OF THE GENUS SAARA GRAY, 1845 Saara asmussi (Strauch, 1863) new comb. [Common name: Persian Spiny-tailed Lizard] Centrotrachelus asmussi STRAUCH, 1863; Bull. Acad. Sci. St. Pétersbourg, 6: 479. 68 Thomas M. WILMs et al.: On the Phylogeny and Taxonomy of the Genus Uromastyx Merrem, 1820 Uromastix asmussi — BOULENGER 1885; Cat. Liz. brit. Mus., 1: 409. Uromastyx asmussi — MERTENS 1956; Jh. Ver. vaterl. Na- turk. Wúrttemb., 111: 93. Holotype: ZISP 3029 (Zoological Museum, Academy of Sciences, Russian Academy of Sciences, St. Petersburg), male, Seri-Tschah (Eastern Persia), coll. Keyzerling, 1858-1859. Differential diagnosis: The species asmussi belongs to the genus Saara. This taxon is distinguished from Saara hardwickii by having 1-2 rows of unkeeled intercalary scales separating each tail whorl dorsally (2-6 keeled in- tercalary scales in S. hardwickii). S. asmussi is distin- guished from S. /oricata in having fewer preanofemoral- pores (8-13 in S. asmussi vs. 14-20 in S. loricata). Subspecies: None Description: Maximum total length 475 mm, maximum SVL 265 mm. 170-201 scales around mid-body, 94-103 scales between gular- and inguinal fold, 40-53 gular scales, 21-27 scales counted from the mid of the lower end of the ear opening to the mental scale. On both sides 5-7 scales between supralabial and enlarged subocular scale. 25-30 scales around 5!h whorl. 23-26 tail whorls. 11-13 scales beneath 4th left toe. 8-13 preanofemoral pores on either side. Colouration: Head, shoulders and forelegs coloured light grey to blue. Hindlegs yellowish grey to blue. Tail dull grey-olive with yellowish spines or completely blue. Back light ocker yellow up to the tailroot; some tubercules on the back are coloured orange. The belly is yellowish white with dark spots on the breast. At low temperatures the back is blackgrey. For pictures of live animals see ANDERSON (1999). Distribution: Saara asmussi lives in the dry areas of Iran, Afghanistan and Pakistan. In Iran the species lives in the following provinces: Esfehan, Kerman, Khorasan and Baluchistan-Sistan (ANDERSON 1974, 1999). In Pakistan the species is known from Baluchistan (MINTON 1966, KAHN 1980). The presence in Afghanistan obviously is limited to the southern part of the country in the border- ing area with Iran and Pakistan (for map see ANDERSON 1999 and WILMs 2001). Saara hardwickii (Gray, 1827) [Common name: Indian Spiny-tailed Lizard] Uromastix hardwickii Gray, 1827; in HARDWICKE & GRAY, Zo0L.J.3:219. Uromastix griseus Cuvier, 1829; Regne animal, Ed. 2, 2: 34. Uromastix reticulatus Cuvier, 1829; (nomen nudum; syn. fide BOULENGER 1885), Regne animal, Ed. 2, 2: 34. Uromastyx grisseus — GRAY 1831; GRAY (ex errore) in GRIFFITH, Animal Kingdom of Cuvier 9 Synops. Spec.: 62. Centrocercus griseus — FITZINGER 1843; (non Centrocer- cus SWAINSON 1831 = Aves), Syst. Rept. 1: 18, 86. Uromastyx similis Fitzinger, 1843; (nomen nudum; syn. fide BOULENGER 1885), Syst. Rept., 1: 86 Saara hardwickii — GRAY 1845; Cat. Spec. Liz. Coll. brit. Mus.: 262. Uromastyx hardwickii — KAHN 1980; Biologica 26 (1/2): 133. Uromastyx hardwickú — SHARMA 1992; Cobra, Madras Snake Park Trust 10: 8 (error typographicus). Holotype: BMNH 1946.8.14.44, male, Plains of Kanouge, Hindustan, India, pres. General Hardwicke, without date. Differential diagnosis: The species hardwickii is the type species of the genus Saara. This taxon is distinguished from S. asmussi and S. loricata by having 2-6 keeled in- tercalary scales separating each tail whorl dorsally (1-2 rows of unkeeled intercalary scales in S. asmussi and S. loricata). Subspecies: None Description: Maximum total length 438 mm, maximum SVL 233 mm. 190-275 scales around mid-body, 112-157 scales between gular- and inguinal fold, 32-46 gular scales, 24-42 scales counted from the mid of the lower end of the ear opening to the mental scale. On both sides 6-9 scales between supralabial and enlarged subocular scale. 40-52 scales around 5h whorl. 28-39 whorls. 15-21 scales beneath 4'h left toe. 12-19 preanofemoral pores on either side. Colouration: The colouration of the back is yellow brown, with dark dots or with a vermiculation. The belly is whitish. The throat is scattered with dark dots. The front sides of the upper thighs on both sides show a black spot at the base of the frontlegs. The pattern of the juveniles consists of black dots, which are arranged in a regular way on the back. For pictures of live specimens see WILMS (2005). Bonner zoologische Beiträge 56 (2007) 69 Distribution: Saara hardwickii is widely distributed in the dry areas of northwest India and Pakistan. In Afghanistan this species lives at least in the border area with Pakistan (near Jalalabad; WiLms 2001). Saara loricata (Blanford, 1874) new comb. [Common name: Iraqi Spiny-tailed Lizard] Centrotrachelus loricatus Blanford, 1874; Proc. zool. Soc. London, 1874: 660. Centrotrachelus asmussi — MURRAY 1884; Ann. Mag. Nat. Hist. 14 (Ser.5): 101. Uromastix loricatus — BOULENGER 1885; Cat. Liz. brit. Mus., 1: 409. Uromastix costatus Muller, 1885; Verh. natforsch. Ges. Wien 7: 292 £ 713 (syn. fide BOULENGER, Zool. Rec. 1885). Uromastyx asmussi loricatus — MERTENS 1956; Jh. Ver. vaterl. Naturk. Wúrttemb., 111: 93. Uromastyx loricatus — CLARK, CLARK & ANDERSON 1966; Occ. Pap. Calif. Acad. Sci. 55: 6. Centrotrachelis loricatus — HAAS & WERNER 1969; Bull. Mus. Comp. Zool., 138 (6): 341. Uromastyx loricata — WıLms 1995; Dornschwanzagamen: 95. Holotype: BMNH 1946.8.11.59, female, Bushir, Iraq, pres. P.L. Sclater, without date. Differential diagnosis: This taxon is distinguished from Saara hardwickii by having 1-2 rows of unkeeled inter- calary scales separating each tail whorl dorsally (2-6 keeled intercalary scales in S. hardwickii). S. loricata is distinguished from S. asmussi in having more pre- anofemoralpores (8-13 in S. asmussi vs. 15-20 in S. lor- icata). Subspecies: None Description: Maximum total length 520 mm, maximum SVL 290 mm. 183-234 scales around mid-body, 101-110 scales between gular- and inguinal fold, 32-45 gular scales, 24-36 scales counted from the mid of the lower end of the ear opening to the mental scale. On both sides 4-8 scales between supralabial and enlarged subocular scale. 23-33 scales around 5h whorl. 22-26 tail whorls. 11-13 scales beneath 4th left toe. 15-20 preanofemoral pores on each side. Colouration: Head, limbs, back and tail brown, yellow- grey or creme coloured with small intermixed brown dots. Back sometimes vividly red coloured. The belly is yellow brown or yellowish white (KALAF 1959; HAAS & WERN- ER 1969). For pictures of live animals see ANDERSON (1999). Distribution: Saara loricata lives in the arid areas of Iraq and southwest Iran. In Iran the following provinces are in- habitated: Kurdestan-Kermanshah, Kuzestan-Lorestan and Fars (ANDERSON 1974). DEFINITION OF THE GENUS UROMASTYX MERREM, 1820 1820 Uromastyx MERREM, Tent. Syst: 56. — Type species (fide FITZINGER 1843): Stellio spinipes Daudin = Uro- mastyx aegyptia (ForsskAl) Original definition: Cauda squamis magnis crassis ac- uleatis verticillata (Tail annulated by large, thick and spiny scales) (MERREM 1820). Diagnosis: Acrodont dentition, with the premaxillary bone forming in adult specimens a sharp, tooth- like structure replacing the incisive teeth. Tail scalation arranged in dis- tinct whorls, which are not separated by intercalary scales dorsally. Species: Uromastyx acanthinura, U. aegyptia, U. alfred- schmidti, U. benti, U. dispar, U. geyri, U. nigriventris, U. ornata, U. ocellata, U. occidentalis, U. princeps, U. mac- fadyeni, U. shobraki, U. thomasi, U. yemenensis. Distribution: The species of the genus Uromastyx are dis- tributed in all North African countries bordering the Sa- hara desert (Algeria, Chad, Egypt, Libya, Mali, Maurita- nia, Morocco, Niger, Sudan, and Tunesia) as well as in Ethiopia, Eritrea, Djibouti and Somalia in Africa and in all countries on the Arabian Peninsula. In the north Uro- mastyx occures in Israel, Jordan, Syria, Iraq and to the east on a narrow stripe along the Arabian Gulf in Iran (up to the city of Bandar Abbas). Taxonomy: Within the genus Uromastyx several taxono- mic problems remain unresolved. Further studies should focus on the phylogenetic relationships between and with- in the different species groups (eg. U. o. ornata and U. o. philbyi) as well as on the evaluation of the taxonomic placement of U. princeps, U. macfadyeni, U. alfred- schmidti and U. occidentalis within the genus. 70 Thomas M. WiLms et al.: On the Phylogeny and Taxonomy of the Genus Uromastyx Merrem, 1820 SYNOPSIS OF THE SPECIES OF THE GENUS UROMASTYX MERREM, 1820 Uromastyx acanthinura Bell, 1825 [Common name: North African Spiny-tailed Lizard] Uromastyx acanthinurus Bell, 1825; Zool. J., 1:457. Uromastix mutabilis — FISCHER 1885; Zool. Garten 26: 272. Uromastix acanthinurus — BOULENGER 1885; Cat. Liz. Brit. Mus. Vol. 1: 406. Uromastix acanthinurus nigerrimus — HARTERT, 1913; No- vitat. Zool. Tring 20: 79. Uromastyx acanthinurus acanthinurus — MERTENS 1962; Senckenberg. biol. 43: 426. Uromastyx acanthinura acanthinura — WıLms 1995; Dorn- schwanzagamen: 57. Holotype: OUM 7845 (Oxford University Museum of Na- tural History), N. Africa (brought by Capt. Lyon RN), Bell & Hope Collection. Differential diagnosis: U. acanthinura ıs distinguished from U. thomasi and U. princeps by the longer and nar- rower tail (50.27-74.42 % of SVL in U. acanthinura vs. 25.00-36.16 % in U. thomasi and 34.62-52.55 % in U. princeps); from the species of the U. ocellata group and from U. macfadyeni by the arrangement of the annuli of the tail: last 8-21 forming a continuous scale row each (U. ocellata group and U. macfadyeni) vs. 2-5 whorls form- ing a continuous scale row in U. acanthinura; from U. ae- gyptia and U. occidentalis by the lower scale counts around midbody (238-322 in U. aegyptia, 297-301 in U. occidentalis vs. 146-195 in U. acanthinura), from U. geyri and U. alfredschmidti by the shorter tail (50.27-74.42 % of SVL in U. acanthinura vs. 65.45-98.06 % in U. geyri and 79.31-87.26 % in U. alfredschmidti). Diagnostic char- acters between U. acanthinura and the subspecies of U. dispar are: Lower number of scales around midbody [145-195 (mean. 165.6) in U. acanthinura vs. 187-227 (mean: 205.0) in U. d. dispar]; lower number of ventrals [74-96 (mean. 83.1) in U. acanthinura vs. 88-118 (mean: 104.5) in U. d. flavifasciata| and lower number of sub- digital scales [9-15 (mean: 12.7) in U. acanthinura vs. 15-18 (mean: 16.4) in U. d. maliensis]. U. acanthinura is differentiated from U. nigriventris by being much less colourful and lacking red, green and citreous colouration. Subspecies: None Description: Maximum total length 430 mm, maximum SVL 253 mm. 146-195 scales around midbody, 74-96 scales between gular- and inguinal fold, 25-45 gular scales, 22-38 scales counted from the mid of the lower end of the ear opening to the mental scale. On both sides 4-8 scales between supralabial and enlarged subocular scale. 26-32 scales around 5!h whorl. 16-20 tail whorls. 9-15 scales beneath 4'h left toe. 10-16 preanofemoral pores on each side. Colouration: Pattern and colouration of U. acanthinura is not very variable. There is a sexual dimorphism with males being black with white or yellowish dots and fe- males being beige to silvergrey with small dark spots. For pictures of live specimens see SCHLEICH et al. (1996) and WILMS (2005). Distribution: U. acanthinura lives at the northern edge of the Sahara, but penetrades deep into the central Sahara along wadis or along plateaus and mountain chains. Geo- graphically it occurs in the dry areas of eastern Algeria, Tunesia and northwest Libya. For detailed discussion of the distribution of this taxon and for a distribution map see WILMS (2005). Further studies are needed to assess the geographic distribution of U. acanthinura and U. nigriven- tris in the area between the Great Ergs (Grande Erg Oc- cidental and Grande Erg Oriental). Uromastyx aegyptia Forsskal, 1775 [Common names: Egyptian Spiny—tailed Lizard] Differential diagnosis: U. aegyptia is distinguished from U. thomasi and U. princeps by the longer tail (60.18-102.83 % of SVL in U. aegyptia vs. 25.00-36.16 % in U. thomasi and 34.62-52.55 % in U. princeps); from the species of the U. ocellata group and from U. mac- fadveni by the arrangement of the annuli of the tail: last 8-21 forming a continuous scale row each (U. ocellata group and U. macfadyeni) vs. 2-8 whorls forming a con- tinuous scale row in U. aegyptia; from the species of the U. acanthinura group by more scales around midbody (238-322 in U. aegyptia vs. 142-231 in the species of the U. acanthinura group). U. aegyptia is distinguished from U. occidentalis by having preanofemoral pores. Subspeeies: We recognize three ofthe closely related taxa within the U. aegyptia clade as subspecies of one single species: U. aegyptia aegyptia, U. a. microlepis and U. a. leptieni. The phylogenetic relationship of the nominal species U. occidentalis requires further studies based on new material. Uromastyx aegyptia aegyptia ( Forsskal, 1775) Lacerta aegyptia Forsskal, 1775; Descr. Anim. Itin. ori- ent. 13. Bonner zoologische Beitráge 56 (2007) 71 Lacerta harbai Forsskal, 1775; Descr. Anim. Itin. orient.: 9 (? syn. fide MERREM 1820). Stellio spinipes Daudin, 1802; Hist. nat. gen. part. Rept. 4:31. Uromastyx spinipes — MERREM 1820; Tent. Syst. Amph.: 56. Lacerta herbai — MERREM 1820; Tent. Syst. Amph.: 56 (nomen substitutum pro Lacerta harbai Forsskal, 1775). Mastigura spinipes — FLEMING 1822; Philos. Zool., 2: 277. Uromastix spinipes — BOULENGER 1885; Cat. Liz. brit. Mus. 1: 407. Uromastix aegyptius — ANDERSON 1896; Contrib. Herpetol. Arabia: 79, 85. Uromastyx aegyptia — FLOWER 1933; Proc. zool. Soc. Lon- don 1933: 779. Uromastyx aegyptius — WERNER 1982; Comun.,Wildl. Res. Rep. 13: 155. Herp. Uromastyx aegyptius aegyptius — ARNOLD 1987; Proc. Symp. Fauna Zoogeogr. Middle East. 28: 249. Uromastyx aegyptia — SCHATTI & GASPARETTI 1994; Fau- na of Saudi Arabia 14: 369. Uromastyx aegyptia aegyptia — Wırms 1995; Dorn- schwanzagamen: 71. Neotype: ZFMK 44216, adult male, Suez at the road to Cairo, Egypt, coll. I. REHAK, VII. 1982 (designated by WILMS & BOHME 2000 a). Differential diagnosis: The nominotypic subspecies is distinguished from U. a. microlepis by having enlarged tubercular scales scattered over the scalation of the flanks and by lower scale counts. It is distinguished from U. a. leptieni by a different juvenile colour pattern and a high- er number of ventrals (see WILMS & BOHME 2000 a). Description: Maximum total length exceeding 700 mm. 247-322 scales around midbody, 126-158 scales between gular- and inguinal fold, 33-59 gular scales, 24-31 scales from the mid of the lower end of the ear opening to the mental scale. On both sides 4-7 scales between supralabi- al and enlarged subocular scale. 29-46 scales around Sth whorl. 20-23 tail whorls. 16-20 scales beneath 4" left toe. 14-20 preanofemoral pores on either side. Colouration: U. aegyptia has the ability of a physiolog- ical colour change. At high temperatures the animals show a light brown to light grey coloration with a black throat and small black dots on the neck. Some individuals have an entirely black to dark blue colouration of the head. At low temperatures the animals show a dark grey, nearly black, colouration. Juveniles have characteristic transverse rows of yellow to orange ocelli on their back. The main colouration of the body is greyish brown. For pictures of live specimens see WILMS (2005). Distribution: The nominotypic subspecies inhabits noth- ern Egypt east of the river Nile, the Sinai Peninsula, Palestina and extreme northwestern Saudi Arabia (Wadi Sawawin / Jabal as Sinfa). The border between the ranges of the taxa aegyptia and microlepis is obviously east of Wadi Araba in Palestina and Jordan and east of Wadi Sawawin in the Jabal as Sinfa region of Saudi Arabia. Uromastyx aegyptia microlepis Blanford, 1874 Uromastix microlepis Blanford, 1874; Proc. zool. Soc. London, 1874. Uromastyx microlepis — SCHMIDT 1939; Field Mus. nat. Hist. Zool. 24: 59. Uromastyx aegyptius — SCHMIDT 1941; Field Mus. nat. Hist. Zool. 24 (16): 162. Uromastyx aegyptius microlepis — MERTENS 1956; Jh. Ver. vaterl. Naturk. Wúrttemb., 111: 93. Uromastix aegyptius — KEVORK & AL-UTHMAN 1972; Bull. Iraq Nat. Hist. Mus. 5 (2): 26. Uromastyx aegyptius — Moopy 1987; Proc. 4th General Meeting of the Societas Europaea Herpetologica: 287. Uromastyx aegyptia — SCHATTI & GASPARETTI 1994; Fau- na of Saudi Arabia 14: 369. Uromastyx aegyptia microlepis — WıLms 1995; Dorn- schwanzagamen: 72. Lectotype: BMNH 1946.8.14.55, adult male, Basrah, Iraq, leg. Capt. Phillips, without date (designated by WILMS & BÖHME 2000 a). Differential diagnosis: Uromastyx a. microlepis is dis- tinguished from U. a. aegyptia by lacking enlarged tuber- cular scales scattered over the scalation of the flanks and by smaller scales. It is distinguished from U. a. leptieni by a different juvenile colour pattern and a higher num- ber of ventrals (see WILMS & BÖHME 2000 a). 72 Thomas M. WILMs et al.: On the Phylogeny and Taxonomy of the Genus Uromastyx Merrem, 1820 Description: Maximum total length exceeding 700 mm. 255-391 scales around mid-body, 149-193 scales between gular- and inguinal fold, 38-65 gular scales, 27-49 scales counted from the mid of the lower end of the ear open- ing to the mental scale. On both sides 5-8 scales between supralabial and enlarged subocular scale. 30-43 scales around 5!h whorl. 20-24 tail whorls.14-23 scales beneath 4th left toe. 13-21 preanofemoral pores on either side. Colouration: U. aegyptia microlepis has the ability of physiological colour change. At high temperatures the an- imals show a light brown to yellow or greenish coloration with a black throat and small black dots on the neck and dorsum. Some individuals have an entirely black to dark blue colouration of the head. At low temperatures the an- imals show a dark grey, nearly black, colouration. For pic- tures of live specimens see WILMS (2005) and SINDALCO & JEREMCENKO (2008). Juveniles have characteristic transverse rows of yellow to orange ocellae on their back. The main colouration of the body is greyish brown. Distribution: Uromastyx aegyptia microlepis lives in the deserts and semideserts of Arabia (Saudi Arabia, Yemen, Oman, United Arab Emirates, Qatar, Kuwait), in Jordan, Syria, Iraq and coastal Iran. Uromastyx aegyptia leptieni Wilms & Böhme, 2000 new status Uromastyx leptieni Wilms & Böhme, 2000; Herpetozoa 13(3/4): 142. Uromastyx leptiens — HARRIS, VACONCELOS & BRITO 2007; Amphibia-Reptilia 28 (2007): 1 (error typographicus). Holotype: ZFMK 52398, adult female, Wadi Sij1, United Arab Emirates (UAE), coll. R. LEPTIEN, VI. 1983. Differential diagnosis: Uromastyx a. leptieni is distin- guished from aegyptia and microlepis by a different ju- venile colour pattern and a lower number of ventrals (see WILMS & BÖHME 2000 a). Description: Maximum total length 675 mm. 238-294 scales around mid-body, 112-130 scales between gular- and inguinal fold, 40-47 gular scales, 30-37 scales count- ed from the mid of the lower end of the ear opening to the mental scale. On both sides 5-7 scales between supral- abial and enlarged subocular scale. 32-37 scales around 5th whorl. 22-24 tail whorls. 17-21 scales beneath 4" left toe. 12-19 preanofemoral pores on either side. Colouration: Main colour olive-beige with small dark dots. Neck, head and throat black. In some specimens the throat is marbled with black and orange. Ventral part of the front-legs, chest and belly marbled with grey. Ventral parts of the hind legs and first half of the tail grey. Colouration of juveniles red brown to dark brown with a dark brown to black net-like pattern. For pictures of live specimens see WILMS (2005) and WILMS & BÖHME (2007). Distribution: Uromastyx aegyptia leptieni is known from east of the Hajar al-Gharbi mountains in northern Oman (from the vicinity of Muscat up to the Musandam penin- sular), and from north-eastern United Arab Emirates. The westernmost locality is near Abu Dhabi Airport (24° 27” N 54°38’E). For detailed distribution maps for the taxa assigned here to the species Uromastyx aegyptia on sub- specific level see WiLms & BOHME (2007). Uromastyx alfredschmidti Wilms & Böhme, 2001 [Common name: Schmidt's Spiny-tailed Lizard] Uromastyx acanthinurus — JOGER 1981; Bonn. Zool. Beitr. 32 (34): 323. Uromastyx alfredschmidti Wilms & Böhme, 2001; Zool. Abh. 51 (1): 95. Holotype: ZFMK 24643, adult male, Tassili N’Ajjer, Tamrit Plateau (1600 m), approx. 30 km northeast Djanet, Algeria, leg. Dr. G. Wangorsch, 22.07.1974. Differential diagnosis: U. alfredschmidti is distinguished from U. thomasi and U. princeps by the longer and nar- rower tail (79.31-87.26 % of SVL in U. alfredschmidti vs. 25.00-36.16 % in U. thomasi and 34.62-52.55 % in U. princeps); from the species of the U. ocellata group and U. macfadyeni by the arrangement of the annuli of the tail: last 8-21 forming a continuous scale row each (U. ocel- lata group and U. macfadyeni) vs. 2-3 whorls forming a continuous scale row in U. alfredschmidti; from U. aegyp- tia and U. occidentalis by the lower scale counts around midbody (238-322 in U. aegyptia, 297-301 in U. occi- dentalis vs. 138202 in U. alfredschmidti), from U. acan- thinura, U. nigriventris and U. dispar by its longer tail (79.31-87.26 % of SVL in U. alfredschmidti vs. 50.27-74.42 in U. acanthinura, 47.83-70 % in U. dispar and 43.48-75.14 % in U. nigriventris). From U. geyri it is distinguished by differences in the scalation of the flanks (enlarged triangular and imbricate scales in U. alfred- schmidti vs. enlarged tubercular scales in U. geyri), as well as the complete black colouration of adult males in U. al- fredschmidti. Subspecies: None Bonner zoologische Beiträge 56 (2007) 73 Description: Maximum total length 429 mm, maximum SVL 230 mm. 138-202 scales around midbody, 68-94 scales between gular- and inguinal fold, 26-42 gular scales, 17-36 scales counted from the mid of the lower end of the ear opening to the mental scale. On both sides 3-6 scales between supralabial and enlarged subocular scale. 28-32 scales around Sth whorl. 21-24 tail whorls. 12-15 scales beneath 4th left toe. 13-21 preanofemoral pores on each side. Colouration: Adult males are entirely black. One female from the type-series has a lightbrown colour, with throat and rear part of the abdomen being ivory-coloured with a light brown reticulation. The top of the tail is coloured dark brown. Adult females can also be totally black. For pictures of live specimens see WILMS (2005) and SINDAL- CO & JEREMCENKO (2008). Distribution: U. alfredschmidt is restricted to the Tamrit plateau, the Hoggar Mountains in southern Algeria and the Akkakus region in southwestern Libya (for distribution map see WILMS & BOHME 2001) Uromastyx benti (Anderson, 1894) [Common name: Yemeni Spiny-tailed Lizard] Aporoscelis benti Anderson, 1894; Ann. Mag. nat. Hist., London, (6) 14: 376. Uromastix (Aporoscelis) benti— ANDERSON 1896; Contrib. Herpetol. Arabia: 33. Uromastix simonyi Steindachner, 1899; Anz. Akad. Wiss. Wien. math. naturwiss. Kl., 36: 143 Uromastyx benti — PARKER 1938; Ann. Mag. Nat. Hist. (11) 1: 486. Aporoscelis benti — SCHMIDT 1939; Field Mus. nat. Hist. Zool. 24: 59. Uromastix philbyi — Haas & BATTERSBY 1959; Copeia 1959: 202 (syn. fide ARNOLD 1986). Uromastyx thomasi — AL-BADRY & AL-SAFADI 1982; Proc. Egypt. Acad. Sci 34: 66 (syn. fide SCHATTI 1989). Uromastyx (Aporoscelis) benti — JOGER 1987; Proc. Symp. Fauna Zoogeogr. Middle East. 28: 260. Uromastyx ocellata benti — SCHÄTTI & GASPARETTI, 1994; Fauna of Saudi Arabia 14: 369. Uromastyx ocellata — SCHATTI & DESVOIGNES 1999; The Herpetofauna of southern Yemen and the Sokotra Archipelago: 39. Lectotype: BMNH 1946.8.11.72, adult male, Wadi Hadramaut, Yemen, leg. Dr. J. Anderson, without date (designated by WILMS & BÖHME 2000 b). Differential diagnosis: Uromastyx benti 1s distinguished from U. thomasi and U. princeps by the significantly longer tail. From all remaining species of the genus (with the exception of the U. ocellata group and U. macfadyeni) by the arrangement of the annuli of the tail: last 8-21 form- ing a continuous scale row each (U. ocellata group and U. macfadyeni) vs. 2-5 whorls forming a continuous scale row (all other Uromastyx species). From U. ocellata, U. ornata and U. macfadyeni the species differs in lacking femoral- and preanalpores. Uromastyx benti differs from U. shobraki and U. yeme- nensis in having larger scales around midbody (188.92 +/- 13.22 in U. shobraki, 197.44 +/- 20.9463 in U. vemenen- sis vs. 160.05 +/- 8.98 in U. benti) and larger ventrals (86.64 +/- 4.88 in U. shobraki, 88.25 +/- 6.98 in U. yeme- nensis vs. 74 +/- 4.02 in U. benti), but also in significant genetic differences. Subspecies: None Description: Maximum total length 360 mm, maximum SVL 196 mm. 143-187 scales around midbody, 66-86 scales between gular- and inguinal fold, 23-33 gular scales, 19-27 scales counted from the mid of the lower end of the ear opening to the mental scale. On both sides 4-7 scales between supralabial and enlarged subocular scale. 28-38 scales around Sth whorl. 22-26 tail whorls. 11-15 scales beneath 4'h left toe. No preanofemoral pores. Colouration: Ground colour of the males back, tail and hind legs yellowish brown. Tail without distinct pattern, hind legs with a turquoise and orange colouration. Back with a pattern consisting of dark brown lines and dots, as well as 7-9 rows of ocellae (ivory coloured with dark brown edges); Dorsal side of the front legs anthracite coloured with orange and green colour elements. Hands yellowish brown. Head orange, dark brown and black mar- bled. Underside of the head anthracite coloured with some orange dots. Ventral parts of forelegs and chest marbled with grey. Belly with narrow grey/anthracite crossbands. The females are much paler in colouration, with a yellow- ish brown ground colour and a pattern of small dark brown lines and dots. For pictures of live specimens see WILMS & BÖHME (2007) and SINDALCO & JEREMCENKO (2008). Distribution: Uromastyx benti occurs in southern and southeastern Yemen, from the vicinity of Azzan eastwards to the Hadramaut Valley and along the coast of the Ara- bian Sea. In the Sultanate of Oman this species is only known from the vicinity of Mirbat in south-western Oman (SEUFER et al. 1998, WiLms & HULBERT 2000). 74 Thomas M. WiLms et al.: On the Phylogeny and Taxonomy of the Genus Uromastyx Merrem, 1820 Uromastyx dispar Heyden, 1827 [Common name: Southern Saharan Spiny-tailed Lizard] Differential diagnosis: U. dispar is distinguished from U. thomasi and U. princeps by the longer and narrower tail (43.83-70 % of SVL in U. dispar vs. 25.00-36.16 % in U. thomasi and 34.62-52.55 % in U. princeps); from the species of the U. ocellata group and U. macfadyeni by the arrangement of the annuli of the tail: last 8-21 form- ing a continuous scale row each (U. ocellata group and U. macfadyeni) vs. 2-5 whorls forming a continuous scale row in U. dispar; from U. aegyptia and U. occidentalis by the lower scale counts around midbody (238-322 in U. aegyptia, 297-301 in U. occidentalis vs. 164-231 in U. dispar), from U. geyri and U. alfredschmidti by the shorter tail (43.83-70 % of SVL in U. dispar vs. 65.45-98.06 % in U. geyri and 79.31-87.26 % in U. al- fredschmidti). Diagnostic characters between U. acan- thinura, U. nigriventris and the subspecies of U. dispar are: Lower number of scales around midbody [145-195 (mean. 165.6) ın U. acanthinura, 139-208 (mean: 170.63) in U. nigriventris vs. 187-227 (mean: 205) in U. d. dis- par]; lower number of ventrals [74-96 (mean. 83.1) in U. acanthinura, 66-99 (mean: 83.98) in U. nigriventris vs. 88-118 (mean: 104.5) in U. d. flavifasciata] and lower number of subdigital scales [9-15 (mean: 12.7) in U. acanthinura, 9-17 (mean: 13.15) in U. nigriventris vs. 15-18 (mean: 16.4) in U. d. maliensis]. For a detailed dis- cussion of the differences between acanthinura, nigriven- tris, dispar, flavifasciata and maliensis see WILMS & Bónme (2001). Subspecies: We recognize three of the closely related taxa within the U. acanthinura clade as subspecies of U. dis- par: U. d. dispar, U. d. flavifasciata and U. d. maliensis. Uromastyx dispar dispar Heyden, 1827 Uromastyx dispar Heyden, 1827; Atl. Reise nórdl. Afr. Rept.: 5. Uromastix acanthinurus — WAKE & KLUGE 1961; Contr. Sci. No. 40: 11. Uromastyx acanthinurus dispar — MERTENS 1962; Senckenb. biol. 43: 430. Holotype: SMF 10417, female, Desert near Ambukol and Dongola, Sudan, coll. E. Rúppel, 1826. Differential diagnosis: Discrimination between the sub- species of U. dispar is possible only by means of coloura- tion of adult males. Adult U. d. dispar males are distin- guished from adult U. d. flavifasciata males by lacking transversal stripes on the animals back and from adult U. d. maliensis males by the less pronounced black coloura- tion of the body. Description: Maximum total length 376 mm, maximum SVL 231 mm. 187-227 scales around midbody, 79-103 scales between gular- and inguinal fold, 30-43 gular scales, 24-33 scales counted from the mid of the lower end of the ear opening to the mental scale. On both sides 4-8 scales between supralabial and enlarged subocular scale. 30-36 scales around 5'h whorl. 16-21 tail whorls. 12-18 scales beneath 4' left toe. 11-18 preanofemoral pores on each side. Colouration: Adult males of U. d. dispar show a black colouration of limbs, heads and tails. Dorsal colouration is yellow or yellowish green. Females are sand coloured with small black dots and occasionally 4-5 grey bars at the flanks. For pictures of live specimens see WILMS et al. (2003) and WILMs (2005). Distribution: U. dispar dispar is found in the desert areas west of the Nile in Sudan and in the Tibesti and Ennedi mountains in Chad. The northernmost locality is Wadi Halfa at the border beween Sudan and Egypt. This taxon has not been found in Egypt yet (contra SALEH 1997). In the Ennedi Mountains, U. d. dispar is known from Fada, while it is known from Bardai and Zouar in the Tibesti Mountains. Between the Ennedi and Tibesti this taxon is known from Ouniaga/Erdi. The westernmost location is Zouar (western Chad). Uromastyx dispar flavifasciata Mertens, 1962 Uromastyx acanthinurus flavifasciatus Mertens, 1962; Senckenb. biol. 43: 427. Uromastyx acanthinura acanthinura — WILMS 1995; Dorn- schwanzagamen: 57. Uromastyx acanthinura flavifasciata — SCHLEICH, KÄSTLE & KABISCH 1996; Amph. & Rept.of North Africa: 309. Uromastyx flavifasciata flavifasciata — MATEO, GENIEZ, LÓPEZ-JURADO & Bons 1998; Rev. Esp. Herp. 12: 104. Uromastyx flavifasciata obscura MATEO, GENIEZ, LÓPEZ- JURADO & Bons, 1998; Rev. Esp. Herp. 12: 104. Uromastyx dispar flavifasciata — WILMS & BÖHME 2001; Zool. Abh. 51 (1): 88. Uromastyx flavifasciata — GENIEZ, MATEO, GENIEZ & PETHER 2004; Amph. & Rept. ofthe Western Sahara: 94. Holotype: SMF 58032, male, approx. 50 km north of Dakar, Senegal (For the reliability of the type locality see BÖHME 1978), coll. N. Heidrich, 01.11.1961. Differential diagnosis: Adult U. d. flavifasciata males can be distinguished from U. d. dispar and U. d. maliensis Bonner zoologische Beitráge 56 (2007) 19 males by their black body colouration with 5—7 wide, clearly-defined yellow, white or red dorsal crossbands. Oc- casionally these crossbands can be reduced or be even completely absent. Description: Maximum total length 455 mm, maximum SVL 280 mm. 164-231 scales around midbody, 88-118 scales between gular- and inguinal fold, 37-48 gular scales, 27-36 scales counted from the mid of the lower end of the ear opening to the mental scale. On both sides 5-6 scales between supralabial and enlarged subocular scale. 31-37 scales around Sth whorl. 19-21 tail whorls. 14-18 scales beneath 4' left toe. 13-17 preanofemoral pores on each side. Colouration: Adult males of Uromastyx dispar flavifas- ciata show black body colouration with 5-7 wide, clear- ly-defined yellow, white or red dorsal crossbands, which can occassionally, be absent. Females are sand colored with small black dots and ocelli at their backs. For pic- tures of live specimens see WILMS et al. (2003) and WILMS (2005). Distribution: Uromastyx dispar flavifasciata lives in the Western Sahara south of 28° northern latitude, in Mauri- tania and in southwestern Algeria. Uromastyx dispar maliensis Joger & Lambert, 1996 Uromastix acanthinurus — ANDERSSON 1935; K. Vet. O. Vitterh. Samh. Handl. Ser. B. 4 (10): 9. Uromastyx acanthinurus — PAPENFUSS 1969; Wasman Jour. Biol. 27 (2): 286. Uromastyx sp. — JOGER 1986; Studies in Herpetology: 187. Uromastyx maliensis Joger & Lambert, 1996; J. Afr. Zool. 110 (1): 24. Uromastyx dispar maliensis — WILMS & BOHME 2001; Zool. Abh. 51 (1): 89. Holotype: HLMD RA 1545, female, 40 km southeast of Gao, Mali, coll. H. Rudolf, without date. Differential diagnosis: Adult Uromastyx dispar malien- sis males differ from adult dispar males by the more pro- nounced black colouration of the body and from adult flav- ifasciata males by lacking transversal crossbands on the dorsum. Description: Maximum total length 383 mm, maximum SVL 232 mm. 177-224 scales around midbody, 86-112 scales between gular- and inguinal fold, 34-46 gular scales, 30-40 scales counted from the mid of the lower end of the ear opening to the mental scale. On both sides 5-9 scales between supralabial and enlarged subocular scale. 30-38 scales around 5h whorl. 16-20 whorls. 15—18 scales beneath 4' left toe. 11-17 preanofemoral pores on each side. Colouration: Dorsal colouration in adult male maliensis consists of yellow ocelli on a dark ground colour. The ocel- li may merge partially, though they never form crossbands. Adult females are brownish black with a beige-yellow to yellow dorsal coloration, which may have dark brown to brownish black vermiculation or ocelli. For pictures of live specimens see JOGER & GRAY (1997), WILMS & MULLER (1998) and WiLms (2005). Distribution: U. dispar maliensis lives in northwestern Mali, in the Tilemsi Valley, on the edge of the Adrar des Iforas and in southwestern Algeria (Taoudrart in Tanezrouft). U. dispar maliensis und U. geyri occur sym- patrically in the region of the Adrar des Iforas (JOGER & LAMBERT 1996). The northernmost locality of U. d. maliensis is Gara Djenoum / Hoggar Mountains (WILMS & BÖHME 2001). Uromastyx geyri Müller, 1922 [Common name: Geyr’s Spiny-tailed Lizard] Uromastix temporalis Valenciennes, 1854; C. R. Acad. Sci. 39:89; Uromastix acanthinurus nigerimmus — GEYR VON SCHWEPPENBURG 1917; J. Ornith. 65 (3): 286 (error typo- graphicus). Uromastix geyri MÜLLER, 1922; Naturwiss. Beobachter 63:.193. Uromastyx acanthinurus geyri — MERTENS 1962; Senckenb. biol. 43: 430. Uromastyx geyri — JOGER 1981; Bonn. zool. Beitr. 32 (3— 4): 324. Uromastyx acanthinura geyri — WıLms 1995; Dorn- schwanzagamen: 61. Uromastyx (acanthinura) geyri— JOGER & LAMBERT 1996; Jour. Afr. Zool. 100(1): 24. Neotype: ZFMK 9230 (designated by MÜLLER 1951), ma- le, Gara Djenoum, Ahaggar Mts. Algeria, S Algeria, coll. Frhr. Hans Geyr von Schweppenburg, 10 March 1914. Differential diagnosis: U. geyri is distinguished from U. thomasi and U. princeps by the longer and narrower tail (65.45-98.06 % of SVL in U. geyri vs. 25.00-36.16 % in U. thomasi and 34.62-52.55 % in U. princeps); from the 76 Thomas M. Wırms et al.: On the Phylogeny and Taxonomy of the Genus Uromastyx Merrem, 1820 species of the U. ocellata group and U. macfadyeni by the arrangement of the annuli of the tail: last 8-21 forming a continuous scale row each (U. ocellata group and U. mac- fadveni) vs. 2-5 whorls forming a continuous scale row in U. geyri; from U. aegyptia and U. occidentalis by the lower scale counts around midbody (238-322 ın U. ae- gyptia, 297-301 in U. occidentalis vs. 142-196 in U. geyri), from U. acanthinura, U. nigriventris and U. dis- par by its longer tail (65.45-98.06 % of SVL in U. geyri vs. 50.27-74.42 in U. acanthinura, 47.83-70 % in U. dis- par and 43.48-75.14 % in U. nigriventris). From U. al- fredschmidti it is distinguished by differences in the sca- lation of the flanks (enlarged triangular and imbricate scales in U. alfredschmidti vs. enlarged tubercular scales in U. geyri), as well as the complete black colouration of adult males in U. alfredschmidti. Subspecies: None Description: Maximum total length 355 mm, maximum SVL 193 mm. 142-196 scales around midbody, 69-93 scales between gular— and inguinal fold, 22-40 gular scales, 19-28 scales counted from the mid of the lower end of the ear opening to the mental scale. On both sides 34 scales between supralabial and enlarged subocular scale. 23-32 scales around 5!" whorl. 20-23 tail whorls. 13-17 scales beneath 4th left toe. 13-20 preanofemoral pores on each side. Colouration: Uromastyx geyri shows only a limited vari- ability. The animals are either beautifully vermilion red or shiny yellow. The pattern consists of brown to black ornaments, which form a non continuous reticulated pat- tern, and of transversal rows of eyed-like dots. For pic- tures of live specimens see LOHR (2004) and WILMS (2005). Distribution: Endemic to the Hoggar- and Air mountains, to the Adrar des Iforas in northeastern Mali and southern Algeria as well as to the Tassili N’ Ajjer in the vicinity of Amguid. For a distribution map of the species see WILMS (2005). Uromastyx macfadyeni Parker, 1932 [Common name: Macfadyen’s Spiny-tailed Lizard] Uromastix ocellatus — TORNIER 1905; Zool. Jahrb. Syst., 22 (4): 372 (syn. fide PARKER 1932). Uromastix ocellatus — NEUMANN 1905; Zool. Jahrb. Syst., 22 (4): 392 (syn. fide PARKER 1932). Uromastix macfadyeni Parker, 1932; Proc. zool. Soc. Lon- don, 1932: 353. Uromastyx macfadyeni — LANZA 1983; Monit. zool. ital. 8: 208. Uromastyx ocellata macfadyeni — LANZA 1988; Biogeo- graphia 14: 420. Uromastyx ocellata ocellata — SCHÄTTI & GASPERETTI 1994; Fauna of Saudi Arabia 14: 369. Holotype: BMNH 1946.8.14.54, male, near Berbara British Somaliland, Somalia, pres. & coll. V. S. Bryan, without date. Differential diagnosis: Uromastyx macfadyeni is distin- guished from U. thomasi and U. princeps by the signifi- cantly longer tail. From all remaining species of the genus (with the exception of the U. ocellata group) by the arrangement of the annuli of the tail: last 8-21 forming a continuous scale row each (U. ocellata group and U. mac- fadyeni) vs. 2-5 whorls forming a continuous scale row (all other Uromastyx species). From U. benti, U. shobra- ki and U. yemenensis the species differs in possessing pre- anofemoralpores and from U. ocellata in having enlarged scales on the anterior margin of the ear opening. From U. ornata it is distinguished by a different ratio between tail width at the 5'h tail whorl and between 4th and 5th whorl (tail width between the 4'h and 5th whorl equivalent to 63-79 % of maximum tail width at the 5h whorl in U. or- nata vs. tail width between the 4'h and Sth whorl equiva- lent to 56-62 % of maximum tail width at Sth whorl in U. macfadyeni). Subspecies: None Description: Maximum total length 221 mm, maximum SVL 120 mm. 157-182 scales around midbody, 78-93 scales between gular- and inguinal fold, 29-32 gular scales, 22-27 scales counted from the mid of the lower end of the ear opening to the mental scale. On both sides 3-4 scales between supralabial and enlarged subocular scale. 23-26 scales around 5!h whorl. 22-23 tail whorls. 14-16 scales beneath 4th left toe. 13-15 preanofemoral pores on each side. Colouration: Males have an either yellowish, greenish or bluish ground colour, dorsally with a brown or black net- like pattern. The inner areas of these patterns are bright- ly yellow, brown or bluish. Belly blue or green, partially white. Females are much paler in colouration. Distribution: Uromastyx macfadyeni is known only from the area between Berbera and Heis (20 miles west of Mait) on the Gulf of Aden (Somalia). Bonner zoologische Beiträge 56 (2007) 77 Uromastyx nigriventris Rothschild & Hartert, 1912 new status [Common name: Moroccan Spiny-tailed Lizard] Uromastix acanthinurus nigriventris Rothschild & Har- tert, 1912; Novitat. Zool. 18: 468. Uromastyx acanthinurus werneri Müller, 1922; Naturwis- senschaftlicher Beobachter 63: 201. Uromastyx acanthinurus var. pluriscutata Fejérváry, 1927; Ann. Mag. Nat. Hist. 20 (9): 514. Uromastyx acanthinurus acanthinurus — BONS & GENIEZ 1996; Amphibiens et Reptiles du Maroc: 126 Holotype: BMNH 1969.2074, male, Tilrhempt between Laghouat and Ghardaia, Algeria, coll. W. Rothschild and E. Hartert, without date (for remarks see WILMS & BÖHME 2001). Differential diagnosis: U. nigriventris is distinguished from U. thomasi and U. princeps by the longer and nar- rower tail (43.48-75.14 % of SVL in U. nigriventris vs. 25.00-36.16 % in U. thomasi and 34.62-52.55 % in U. princeps); from the species of the U. ocellata group and U. macfadyeni by the arrangement of the annuli of the tail: last 8-21 forming a continuous scale row each (U. ocel- lata group and U. macfadyeni) vs. 2-5 whorls forming a continuous scale row in U. nigriventris; from U. aegyp- tia and U. occidentalis by the lower scale counts around midbody (238-322 in U. aegyptia, 297-301 in U. occi- dentalis vs. 139-208 in U. nigriventris), from U. geyri and U. alfredschmidti by the shorter tail (43.48-75.14 % of SVL in U. nigriventris vs. 65.45-98.06 % in U. geyri and 79.31-87.26 % in U. alfredschmidti). Diagnostic charac- ters between U. nigriventris and the subspecies of U. dis- par are: lower number of scales around midbody [139-208 (mean: 170.63) in U. nigriventris vs. 187-227 (mean: 205) in U. d. dispar]; lower number of ventrals [66-99 (mean: 83.98) in U. nigriventris vs. 88-118 (mean: 104.5) in U. d. flavifasciata] and lower number of subdigital scales [9-17 (mean: 13.15) in U. nigriventris vs. 15-18 (mean: 16.4) in U. d. maliensis]. U. nigriventris is differentiated from U. acanthinura by being much more colourful, with vividly red, green and citreous coloured specimens. Subspecies: None Description: Maximum total length 415 mm, maximum SVL 250 mm. 139-208 scales around midbody, 66-99 scales between gular- and inguinal fold, 26-46 gular scales, 22-35 scales counted from the mid of the lower end of the ear opening to the mental scale. On both sides 3-6 scales between supralabial and enlarged subocular scale. 25-36 scales around 5!" whorl. 16-21 tail whorls. 9-17 scales beneath 4th left toe. 11-18 preanofemoral pores on each side. Colouration: In Uromastyx nigriventris colouration is very variable, with red, yellow, green and orange coloured specimens. Old adult males show frequently a black colouration of head and belly. For pictures of live speci- mens see WILMS (2005). Distribution: Uromastyx nigriventris is widespread in Morocco east and south of the Atlas Mountain Chain. In western Algeria it ranges in the Sahara Atlas and in the regions northwest, northeast and southwest of the Great Western Erg. In Morocco the southern distribution limits for this taxon are Oued Draa (Draa-valley) and Djebel Ouarkziz (see also discussion of the distribution of this taxon in WILMS & BÖHME 2001). Uromastyx ornata Heyden, 1827 [Common name: Ornate Spiny-tailed lizard] Differential diagnosis: Uromastyx ornata is distin- guished from U. thomasi and U. princeps by the signifi- cantly longer tail. From all remaining species of the genus (with the exception of the U. ocellata group and U. mac- fadyeni) by the arrangement of the annuli of the tail: last 8-21 forming a continuous scale row each (U. ocellata group and U. macfadyeni) vs. 2-5 whorls forming a con- tinuous scale row (all other Uromastyx species). From U. benti, U. shobraki and U. yemenensis the species differs in possessing preanofemoralpores and from U. ocellata in having enlarged scales on the anterior margin of the ear opening. From U. macfadyeni it is distinguished by a dif- ferent ratio between tail width at the 5th tail whorl and be- tween 4th and 5th whorl (tail width between the 4" and 5th whorl equivalent to 63-79 % of maximum tail width at the Sth whorl in U. ornata vs. tail width between the 4th and Sth whorl equivalent to 56-62 % of maximum tail width at 5‘ whorl in U. macfadyeni). Subspecies: Based on the high morphological and genet- ical similarity between ornata and philbyi we consider both taxa to be conspecific and assign them as subspecies to Uromastyx ornata: U. ornata ornata and U. ornata philbyi. Uromastyx ornata ornata Heyden, 1827 Uromastyx ornatus Heyden, 1827; Atlas Reise nórdl. Afr., Rept:: 1. Uromastix ornatus — ANDERSON 1896; Contrib. Herpetol. Arabia: 79. 78 Thomas M. WiLms et al.: On the Phylogeny and Taxonomy of the Genus Uromastyx Merrem, 1820 Uromastix oronatus — FARAG & BANAJA 1980; Bull. Fac. Sci. K.A.U. 4: 12 (error typographicus). Uromastyx ocellatus ornatus — ARNOLD 1986; Fauna of Saudi Arabia 8: 393. Uromastyx ocellata ornata — LANZA 1988; Biogeographia 14: 420. Uromastyx ocellata ornata — SCHÄTTI & GASPARETTI 1994; Fauna of Saudi Arabıa 14: 369. Uromastyx ocellata — SCHÄTTI & DESVOIGNES 1999; The Herpetofauna of south. Yemen and the Sokotra Archipel- ago: 39. Holotype: SMF 10403, Mohila = Al Muwaylih, Saudi Arabia, leg. E. RUPPELL, 1828. Differential diagnosis: Uromastyx o. ornata is distin- guished from U. o. philbyi by its narrower tail (ratio tail length divided by maximum tail width at the 5‘ whorl is 3.61-5.3 in ornata vs. 3.03-3.96 in philbyi). Description: Maximum total length 368 mm, maximum SVL 196 mm (BMNH 97.10.28.199). 149-185 scales around mid-body, 75-99 scales between gular- and in- guinal fold, 22-31 gular scales, 21-27 scales counted from the mid of the lower end of the ear opening to the men- tal scale. On both sides 3-5 scales between supralabial and enlarged subocular scale. 19-25 scales around Sth whorl. 20-23 tail whorls. 11-15 scales beneath 4th left toe. 7-14 preanofemoral pores on each side. Colouration: U. ornata ıs a very variable species. Colour of the males is green, blue or red, with a irregularly red- dish brown net-like pattern and yellow spots on the back. Sometimes yellow cross-bands are present. Ventrum with dark pattern. Females are not as colourful as males. They are light brown with dark brown spots and sometimes light yellow or light red spots. Belly without pattern, yellow- ish or white. For pictures of live specimens see WILMS et al. (2002), WiLms (2005) and WILMS & BÖHME (2007). Distribution: For discussion and map of the distribution range see WILMS & BÖHME (2007). Uromastyx ornata philbyi Parker, 1938 Uromastyx philbyi Parker, 1938; Ann. Mag. nat. Hist. (11) 1: 484. Uromastyx ocellatus philbyi — ARNOLD 1986; Fauna of Saudi Arabia 8: 416. Uromastyx ornatus philbyi — ARNOLD 1987; Proc. Symp. Fauna Zoogeogr. Middle East. 28: 249. Uromastyx ocellata philbyi — SCHÄTTI & GASPARETTI 1994; Fauna of Saudi Arabia 14: 369. Uromastyx ocellata — SCHÄTTI & DESVOIGNES 1999; The Herpetofauna of southern Yemen and the Sokotra Archipelago: 39. Holotype: BMNH 1946.8.14.65 (former number: BMNH 1938.2.1.1), male, between Makkah and Shabwa, south- ern Hejaz, between Mountains and Rub al Khali, Saudi Arabia, coll. H. St. J. B. PHILBy, without date. Differential diagnosis: see under U. o. ornata. Description: Maximum total length 341 mm, maximum SVL 205 mm (MZUF 27906). 138-193 scales around midbody, 69-96 scales between gular- and inguinal fold, 17-31 gular scales, 18-22 scales counted from the mid of the lower end of the ear opening to the mental scale. On both sides 3-5 scales between supralabial and enlarged subocular scale. 22-29 scales around 5'h whorl. 17-22 tail whorls. 11-14 scales beneath 4'th left toe. 7-14 pre- anofemoral pores on each side. Colouration: Similar to Uromastyx o. ornata. For pictures of live specimens see WILMS (2007 b). Distribution: For discussion and map of the distribution areas of this taxon see WILMS & BÖHME (2007). Uromastyx ocellata Lichtenstein, 1823 [Common name: Eyed Spiny-tailed Lizard] Uromastyx ocellatus Lichtenstein, 1823; Verz. Doubl. zo- ol. Mus. k. Univ. Berlin: 107. Uromastix ornatus — BOULENGER 1885; Cat. Liz. Brit. Mus.,1: 406. Uromastix ocellatus — ANDERSON 1898; Zool. Egypt, 1 Rept. Batr.: 127. Uromastyx ocellata ocellata — LANZA 1988; Biogeogra- phia 14: 420. Syntypes: ZMB 809, Nubia; ZMB 811-13, Nubia; ZMB 810, Syria; all specimens leg. Hemprich & Ehren- berg. After DENZER et al. (1997), ZMB 811-13 are lost which we cannot confirm at least for ZMB 811. Differential diagnosis: Uromastyx ocellata is distin- guished from U. thomasi and U. princeps by the signifi- Bonner zoologische Beiträge 56 (2007) 79 cantly longer tail. From all remaining species of the genus (with the exception of the U. ocellata group and U. mac- fadyeni) by the arrangement of the annuli of the tail: last 8-21 forming a continuous scale row each (U. ocellata group and U. macfadyeni) vs. 2-5 whorls forming a con- tinuous scale row (all other Uromastyx species). From U. benti, U. shobraki and U. yemenensis the species differs in possessing preanofemoralpores and from U. ornata and U. macfadyeni in lacking enlarged scales on the anterior margin of the ear opening. Subspecies: None Description: Maximum total length 276 mm, maximum SVL 174 mm. 189-256 scales around midbody, 95-113 scales between gular- and inguinal fold, 29-42 gular scales, 23-33 scales counted from the mid of the lower end of the ear opening to the mental scale. On both sides 4-6 scales between supralabial and enlarged subocular scale. 24-39 scales around 5'h whorl. 22-29 tail whorls. 14-19 scales beneath 4th left toe. 12-17 preanofemoral pores on each side. Colouration: Showing a distinctive sexual dichromatism. Ground colour of males either beautifully red with a black vermiculation, olive green with red dots or red with green dots. On the back 7-8 transversal rows of yellow or white, black edged ocellae. Sides of the neck, throat and breast light green or lively blue coloured. The belly is monochro- matic yellow or white. Females are by far not as lively coloured as the males, with an either pale brownish, green, redish or grey groundcolour. For pictures of live specimens see WILMS (2005) and FRAHM (2006). Distribution: This species occures in the dry areas west of the red sea in the following countries: Somalia (Bora- ma district), Djibouti, Eritrea, Sudan and southeastern Egypt, where the southernmost locality is in the Borama district (northwestern Somalia) and the northernmost in the Wadi Gul'an (Egypt). According to LARGEN & SPAWLS (2006) this species lives also in Ethiopia near the border to Somalia. Uromastyx occidentalis Mateo, Geniez, López-Jurado & Bons, 1998 [Common name: Western Giant Spiny-tailed Lizard] Uromastyx occidentalis Mateo, Geniez, Löpez-Jurado & Bons, 1998 Holotype: DB.ULPGC-5 (Departamento de Biologia, Universidad de Las Palmas de Gran Canaria), Aagtel Ag- mumuit, between Yeloua and Mades (Adrar Souttouf, Western Sahara) (21° 52’N, 15° 31’W), coll. M. Hası, 25. June 1995. Differential diagnosis: U. occidentalis is distinguished from U. thomasi and U. princeps by the longer tail; from the species of the U. ocellata group and U. macfadyeni by the arrangement of the annuli of the tail: last 8-21 form- ing a continuous scale row each (U. ocellata group and U. macfadyeni) vs. less than 7 whorls forming a contin- uous scale row in U. occidentalis; from the species of the U. acanthinura group by more scales around midbody (297301 in U. occidentalis vs. 142-231 in the species of the U. acanthinura group). U. occidentalis is distinguished from U. aegyptia by lacking preanofemoral pores. Subspecies: None Description: Maximum total length 536 mm, maximum SVL 228 mm. 297-301 scales around midbody, 121-122 scales between gular- and inguinal fold. On both sides 7 scales between supralabial and enlarged subocular scale. 23 whorls. No preanofemoral pores. Colouration: Colouration in life not known. For picture of the holotype see MATEO et al. (1998) and WiLms (2005). Distribution: Known only from the type locality and from Udei Sfa (45 km west of Maatal Laj, 22°22’N 15°32’W; GENIEZ et al. 2004). Uromastyx princeps O’Shaughnessy, 1880 [Common name: Princely Spiny-tailed Lizard] Uromastix princeps O’Shaughnessy, 1880; Proc. zool. Soc. London, 1880: 445. Aporoscelis princeps — BOULENGER 1885; Cat. Liz. Brit. Mus.,1: 410. Uromastix princeps scorteccii — CHERCHI 1954; Atti. Soc. ital. Sci. Nat. Milano, 93: 540. Uromastyx princeps — LANZA 1983; Monitore zool. ital. (new Series) Suppl. 18: 208 Uromastyx scortecci — MooDY 1987; Proc. 4t* General Meeting of the Societas Europaea Herpetologica: 286. Holotype: BMNH 1946.814.56, male, Zanzibar, coll. Sir J. Kirk, without date (see comments on type locality in WILMS 2001). Differential diagnosis: With the exception of Uromastyx thomasi, U. princeps 1s distinguished from all other taxa in the genus by its significantly shorter tail. From U. thomasi it is distinguished by the absence of pre- anofemorai pores. Subspecies: None 80 Thomas M. WILMS et al.: On the Phylogeny and Taxonomy of the Genus Uromastyx Merrem, 1820 Description: Maximum total length 265 mm, maximum SVL 180 mm. 150-226 scales around mid-body, 77-128 scales between gular- and inguinal fold, 28-43 gular scales, 22-34 scales counted from the mid of the lower end of the ear opening to the mental scale. On both sides 2-5 scales between supralabial and enlarged subocular scale. 20-27 scales around Sth whorl. 9-14 tail whorls. 14-18 scales beneath 4!" left toe. No preanofemoral pores. Colouration: This species displays a pronounced sexual dimorphism. The ground colour of the body is olive grey to green with small brown markings. Males have a yel- lowish red to green dorsum with small, scattered black spots. The venter 1s yellowish with blue grey marbling in the area of the chest and throat. The tail is yellow green or red in colour. Females are grey brown dorsally with a light red shimmer and small black spots. The venter is im- maculate white. For pictures of live specimens see WILMS & HULBERT (1995) and WiLms (2005). Distribution: Uromastyx princeps is found in the Soma- lian provinces of Sanaag, Bari, Nogal and Mudug. Uromastyx shobraki Wilms & Schmitz, 2007 new sta- tus [Common name: Shobrak’s Spiny-tailed Lizard] Uromastyx ocellata benti — SCHÄTTI & GASPARETTI 1994; Fauna of Saudi Arabia 14: 369. Uromastyx ocellata — SCHÄTTI & DESVOIGNES 1999; The Herpetofauna of southern Yemen and the Sokotra Archipelago: 39. Uromastyx yemenensis shobraki Wilms & Schmitz, 2007; Zootaxa 1394: 16. Holotype: ZFMK 48681, adult male, Mafraq Mocca (Mafraq al-Mukha), km 13.5, Republic of Yemen, leg. B. Schätti, 5.-6.1V.1988. Differential diagnosis: Uromastyx shobraki is distin- guished from U. thomasi and U. princeps by the signifi- cantly longer tail. From all remaining species of the genus (with the exception of the U. ocellata group) by the arrangement of the annulı of the tail: last 8-21 forming a continuous scale row each (U. ocellata group) vs. 2-5 whorls forming a continuous scale row (all other Uro- mastyx species). From U. ocellata, U. ornata and U. mac- fadyeni the species differs in lacking femoral- and preanal- pores. Uromastyx shobraki differs from U. benti in having smaler scales around midbody (188.92 +/- 13.22 in U. shobraki vs. 160.05 +/- 8.98 in U. benti) and smaller ventrals (86.64 +/-4.88 in U. shobraki vs. 74 +/-4.02 in U. benti). U. shobraki is differentiated from U. yemenensis not only by its larger maximum size (393 mm in U. shobraki vs. 337 mm in U. yemenensis) but also in different colour pattern and in significant genetic differences. Subspecies: None Description: Maximum total length 393 mm, maximum SVL 208 mm. 163-207 scales around midbody, 79-97 scales between gular- and inguinal fold, 25-33 gular scales, 23-31 scales counted from the mid of the lower end of the ear opening to the mental scale. On both sides 3-5 scales between supralabial and enlarged subocular scale. 32-39 scales around Sth whorl. 24-27 tail whorls. 15-17 scales beneath 4" left toe. No preanofemoral pores. Colouration: In preserved specimens dorsal surface of head, body and hindlimbs dark brown, tail lighter. Light brown roundish dots (diameter 4-5 scales) are present on the dorsum, tending to form transverse rows. In addition, irregular light brown dots are present on the whole dor- sum. Colour of the hands not different to the colour of the forearm. Head dark brown, with light brown pattern. Ven- tral side yellowish brown. Ventral side of head and chest marbled with anthracite and dark brown. For a picture of a live specimen see WILMS & BOHME (2007). Distribution: South-western Yemen. For a map of the dis- tribution area see WILMS & BOHME (2007). Uromastyx thomasi Parker, 1930 [Common name: Omani Spiny-tailed Lizard] Uromastix thomasi Parker, 1930; Ann. Mag. nat. Hist., London, (10) 6: 595. Uromastyx thomasi — ARNOLD 1980; J. Oman Stud. Spec. Rep. No. 2: 293. Holotype: BMNH 1946.8.14.43 (former number: BMNH 1930.6.30.2), male, Bu Ju’ay, Rub al Khali, Dhofar, Oman, coll. B. Thomas, without date. Differential diagnosis: With the exception of Uromastyx princeps, U. thomasi is distinguished from all other taxa in the genus by its significantly shorter tail. From U. prin- ceps it is distinguished by the presence of preanofemoral pores. Subspecies: None Description: Maximum total length approx. 24 cm, max- imum SVL approx. 19 cm. 125-150 scales around mid- body, 72-100 scales between gular- and inguinal fold, Bonner zoologische Beiträge 56 (2007) 81 25-36 gular scales between a hypothetical line between the anterior margins of the ears and the mental scale, 19-25 scales counted from the mid of the lower end of the ear opening to the mental scale. On both sides 2-4 scales between supralabial and enlarged subocular scale. 28-34 scales around 5!" whorl. 11-13 whorls. 13-18 scales beneath 4" left toe. 12-19 preanofemoral pores each side. Colouration: Yellowish green with a dark net-like pattern. A broad red vertebral-stripe runs from the neck to the first half of the tail. In some specimens an orange to red colouration of the head can occur. Ventral side yellowish or white. Neck and sides of the head of the juveniles striped (black and white). Colouration of the upper side of the body black with 6 lighter transversal bands. Be- tween those bands yellowish to orange coloured ocellae. Tail above brown with some large black spots. Belly and throat white. Ventral side of the tail white with black dots. For pictures of live specimens see WILMS et al. (2002) Distribution: Uromastyx thomasi lives in coastal Oman (for map see WILMS & BÖHME 2007). Uromastyx yemenensis Wilms & Schmitz, 2007 [Common name: South Arabian Spiny-tailed Lizard] Uromastyx ocellata benti — SCHÄTTI & GASPARETTI 1994; Fauna of Saudi Arabia 14: 369. Uromastyx ocellata — SCHÄTTI & DESVOIGNES 1999; The Herpetofauna of southern Yemen andthe Sokotra Archi- pelago: 39. Uromastyx y. yemenensis Wilms & Schmitz, 2007; Zootaxa 1394: 12. Holotype: ZFMK 47861, adult male, Abyan Governorate, vicinity of Lodar (= Lawdar), Republic of Yemen, leg. 1. Haikal, don. 1985. Differential diagnosis: Uromastyx yemenensis is distin- guished from U. thomasi and U. princeps by the signifi- cantly longer tail. From all remaining species of the genus (with the exception of the U. ocellata group) by the arrangement of the annuli of the tail: last 8-21 forming a continuous scale row each (U. ocellata group) vs. 2-5 whorls forming a continuous scale row (all other Uro- mastyx species). From U. ocellata, U. ornata and U. mac- fadyeni the species differs in lacking femoral— and pre- analpores. Uromastyx yemenensis differs from U. benti in having smaler scales around midbody (197.44 +/-20.9463 in U. yemenensis vs. 160.05 +/- 8.98 in U. benti) and smaler ventralia (88.25 +/-6.98 in U. yemenensis vs. 74 +/-4.02 in U. benti). U. yemenensis is differentiated from U. shobraki not only by its smaller maximum size (393 mm in U. shobraki vs. 337 mm in U. yemenensis) but also in different colour pattern and significant genetic differences. Subspecies: None Description: Maximum total length 337 mm, maximum SVL 185 mm. 146-227 scales around midbody, 73-100 scales between gular- and inguinal fold, 25-40 gular scales, 22-30 scales counted from the mid of the lower end of the ear opening to the mental scale. On both sides 4-6 scales between supralabial and enlarged subocular scale. 33-40 scales around 5 whorl. 23-27 tail whorls. 12-18 scales beneath 4" left toe. No preanofemoral pores. Colouration: Ground colour of the males back, tail and hind legs yellowish brown. Tail without distinct pattern, hind legs with very small dark brown dots. Back with a pattern consisting of dark brown lines and dots; five dis- tinct cross bands without or with very few pattern on the back. Dorsal side of the front legs anthracite coloured. Hands yellowish brown. Head yellowish brown, dark brown marbled. Underside of the head anthracite coloured with some yellowish brown dots. Ventral parts of forelegs and chest marbled with grey. Belly with narrow grey/an- thracite crossbands. The females are much paler in colouration. With a yellowish brown ground colour with a pattern of small dark brown lines and dots. Five pale cross bands on the back. The ground colour of the ven- tral side is a light yellowish brown. Distribution: South-western Yemen. For a map of the dis- tribution area see WILMS & BOHME (2007). KEY TO THE SPECIES OF THE GENERA SAARA GRAY, 1845 AND UROMASTYX MERREM, 1820 1 a — Tail whorls separated dorsally by 1-6 continuous TO WS ¡Obten Saara b -Tail whorls without dorsal intercalary scales EEE ED ER EUER Raros. Uromastyx Saara Gray, 1845 1 a-Tail with 29-36 primary whorls; 2-6 rows of keeled intercalary scales between whorls on dorsal surface of tail; dorsal scalation interspersed with irregular, only slightly enlarged, tubercular scales......................S. hardwickii b —Tail with less than 28 primary whorls; 1-2 rows of un- keeled intercalary scales between tail whorls on dorsal sur- face of tail; dorsal scalation with transverse rows of con- spicuously enlarged tubercular scales ........................... 2 82 Thomas M. WiLms et al.: On the Phylogeny and Taxonomy of the Genus Uromastyx Merrem, 1820 2 a — Slightly enlarged scales at front edge of ear open- ing; 8-13 preanofemoral pores on either side; 7-10 scales in a transverse row on the dorsal surface of the tail Dose nenne S. asmussi b -Without enlarged scales at the front edge of the ear opening; 15-20 preanofemoral pores on either side; 12 scales in a transverse row on the dorsal surface of the tail A rg S. loricata Uromastyx Merrem, 1820 1 a — Without preanofemoral pores ..........................- 2 b = With preanofemoral au. 2 a — Tail short, approx. 35-53 % of SVL; 9-14 whorls cai un. Y, princeps b - Tail long, approx. 71-94 % of SVL; 22-27 whorls 3 a — Body scales small, approx. 297-301 scales around midbody; 121-122 scales between gular and inguinal fold sn Ur Occidentalis b — Body scales larger, approx. 143-227 scales at mid- body; 66-100 scales between gular and inguinal fold 4 a— 143-187 scales around midbody (average 160.05 +/- 8.98), 66-86 ventral scales ............................U. benti b — 163-227 scales around midbody (average 192.53 +/- 16.63), 79-97 ventral scales ..............cocccccccccccnccnnneninnnnnns 5 5 a — Ground colour light brown with five distinct cross- bands. on heback ns U. vemenensis b — Ground colour dark brown with light brown dots tend- ing to form transverse rows on the back as id U. shobraki 6 a — Tail short, approx. 25-35 % of SVL, from above dis SINAC Ol as U. thomasi b — Tail long, approx. 48-103 % of SVL, from above a A es tamccamarsontice taste I 7 a — The last 12-21 tail whorls formed of continuous SCALE LOWS ie recae 8 b — The last 2—5 tail whorls formed of continuous scales EOS era avs A sang eisernen 10 8 a — Anterior margin of ear opening without enlarged A en A ae aun U. ocellata b — Anterior margin of ear opening with enlarged scales 9 a— 17-29 (very rarely 31) gular scales; tail width be- tween the 4 and Sth whorl equivalent to 63-79 % of maximum tail width at the 5th whorl .............. U. ornata b — 29-32 gulars; tail width between the 4'h and 54 whorl equivalent to 56-62 % of maximum tail width at 5 whorl nun... U. macfadyeni 10a —- 238-391 scales at midbody, 112-193 ventrals be- tween gular and inguinal fold ...................... U. aegyptia b - 138-227 scales at midbody, 68-112 ventrals between gular and inguinal fold ..........ee. a nl ll a — Tail with 20-24 whorls; tail length in adult spec- imens approx. 70-98 % Of SVL....2... 2 Me 12 b — Tail with 16-21 whorls; tail length approx. 48-75 % SYLT 8 12 a — Several transverse rows of enlarged scales along the flanks; max. total length 35.5 cm; never completely black Coloured «...c.ccssess00scs asses nesiosc IO CyT b — Flank scalation imbricate with enlarged triangular scales; max. total length 42.9 cm; adult males and occa- sionally females completely black........... U. alfredschmidti 13 a — 79-118 ventrals between gular and inguinal fold; 164-231 scales at midbody; 30-38 scales around 5h whorl ae nee U. dispar b — 66-99 ventrals between gular and inguinal fold; 139-208 scales at midbody; 25-36 scales form 5th whorl 14 a — Adult males black with ivory coloured or yellow- ish dots, adult females beige to silvergrey with small dark DON U. acanthinura b -Colouration very variable, with red, yellow, green and orange coloured specimens. Old adult males show fre- quently a black colouration of head and belly a U. nigriventris Bonner zoologische Beiträge 56 (2007) 83 Acknowledgements. We would like to thank H.H. Prince Ban- dar bin Saud, Secretary General, National Commission for Wildlife Conservation and Development (NCWCD), Riyadh, Saudi Arabia; H.E. Prof. Dr. A.H. Abuzinada, former Secretary General, NCWCD; Prof. Dr. I. A. Nader, NCWCD; Dr. H. Tat- wani, NCWCD; Mr. Ali bin Amer al Kiyumi, Director-General of Nature Reserves, Muscat, Oman and Mr. S. M. Al-Saady, Di- rector of Research, Muscat, Oman, for their hospitality and help during the field works of T. Wilms and for issuing the permits required. The first author would also like to thank Dr. M. Shobrak, Taif, Saudi Arabia; Mrs. I. Barcelo & Dr. C. Toureng, ERWDA, Abu Dhabi, UAE; Dr. I. Galal, Riyadh, Saudi Arabia; Mr. P. Paillat, formerly National Wildlife Research Center (NWRC), Taif, Saudi Arabia now National Avian Research Cen- ter (NARC), Sweihan, Abu Dhabi, UAE and the teams of the NWRC, NARC and the King Khalid Wildlife Research Center, Thumamah, Saudi Arabia, for their hospiality and for the sup- port provided. We are grateful to the curators of the following museums for their hospitality, cooperation and/or for the loan of important collec- tions: Dr. Jifi Moravec, National Museum, Museum of Natural History Prague, Czech Republic; Dr. E. N. Arnold, Dr. C. Mc- Carthy and Dr. B. Clarke, The Natural History Museum, Lon- don, UK; Dr. F. Tiedemann, Naturhistorisches Museum Wien, Vienna, Austria; Dr. M. Poggesi and Dr. C. Corti, Museo Zoo- logico de „La Specola“, Florence, Italy; Dr. I. Ineich, Museum National d’histoire naturelle, Paris, France; Mrs. M. Nowak- Kemp, Oxford University Museum of Natural History, Oxford, UK; Prof. F.J. Obst and Dr. U. Fritz, Museum für Tierkunde, Dresden, Germany; Dr. K. Klemmer and Dr. G. Köhler, Forschungsinstitut und Naturmuseum Senckenberg, Frankfurt a.M., Germany; Dr. J. Hallermann, Zoologisches Museum der Universität Hamburg, Germany; Dr. R. Günther and Dr. M.-O. Rödel, Museum für Naturkunde, Humboldt-Universität, Berlin, Germany; Dr. N. Ananjeva, Zoological Museum, Academy of Sciences, Russian Academy of Sciences, St. Petersburg, Russia; Dr. U. Gruber and Dr. F. Glaw, Zoologische Staatssammlung München, Germany. Additional material was made available by Dr. V. Mahnert and Dr. J. Mariaux, Museum d’Histoire naturelle, Geneva, Switzerland. Dr. P.A. Crochet, Dr. P. 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WILMS, T.; MÜLLER, H.D. & LÖHR, B. 2002: Bunte Juwelen im Terrarium — Erfahrungen bei der langjährigen Pflege und Ver- mehrung von Uromastyx ornata Heyden, 1827 bis zur F2-Ge- neration. Draco 3 (10): 41-49. Wits, T.; Rur, D. & LöHr, B. 2003: Zur Haltung und Nachzucht zweier Taxa aus dem Uromastyx-acanthinura- Komplex: Uromastyx geyri MÜLLER 1922 und Uromastyx dis- par flavifaciata Mertens, 1962 (Reptilia: Agamidae: Leiole- pidinae: Uromastyx). Draco 4 (14): 42-55. WILMS, T. & SCHMITZ, A. 2007: A new polytypic species of the genus Uromastyx MERREM 1820 (Reptilia: Squamata: Agami- dae: Leiolepidinae) from southwestern Arabia. Zootaxa 1394: 1-23. WILMS, T. & HULBERT, F. 1995: Uromastyx princeps. Sauria 17 (3): 12 WiLms, T. & HULBERT, F. 2000: On the herpetofauna of the Sul- tanate of Oman, with comments on the relationship between Afro-tropical and Saharo-sindian faunas. Bonner zoologische Monographien 46: 367-380. YANG, Z. H., GOLDMAN, N. & FRIDAY, A. 1994: Comparison of Models for Nucleotide Substitution Used in Maximum-Like- lihood Phylogenetic Estimation. Molecular Biology and Evo- lution 11: 316-324. Received: 18.06.2008 Revised: 27.01.2008 Accepted: 28.01.2008 Corresponding editor: R. van den Elzen 86 Thomas M. WiLms et al.: On the Phylogeny and Taxonomy of the Genus Uromastyx Merrem, 1820 APPENDIX I. Table 1. Character matrix for thirteen polarized characters (Outgroup Leiolepis and all twenty-three taxa in this study). For cha- racter coding see Appendix III. > a an = | = = S Taxon A B C D Sl acanthinura 1 0 0 0 l 0 0 0 0 l 1 0 1 aegyptia 1 0 l | 1 0 1 0 0 1 0 0 1 alfredschmidti 1 0 0 0 l 0 0 0 0 1 0 0 1 asmussi l 0 0 0 l 0 0 0 0 2 0 1 0 benti l 0 0 0 l 0 0 0 l l 0 0 1 dispar i 0 0 0 l 0 0 0 0 1 l 0 1 flavifasciata l 0 0 0 l 0 0 0 0 1 l 0 1 geyri | 0 0 0 | 0 0 0 0 1 0 0 1 hardwickii 0 0 0 l 0 0 0 0 0 2 0 1 0 leptieni l 0 l 0 | 0 l 0 0 l 0 0 l loricata l 0 0 0 | 0 0 0 0 2 0 1 0 macfadyeni l 0 0 0 l 0 0 0 l 1 1 0 1 maliensis l 0 0 0 | 0 0 0 0 l l 0 1 microlepis l 0 l l l 0 l 0 0 l 0 0 l nigriventris | 0 0 0 l 0 0 0 0 l l 0 l occidentalis l 0 l 0 0 0 - 0 1 ocellata l 0 0 0 1 0 0 0 l l 0 0 l ornata l il 0 0 l 0 0 0 l l 0 0 l philbyi l 1 0 0 l 0 0 0 l | l 0 1 princeps 2 0 0 0 1 l 0 1 0 0 2 0 l shobraki | 0 0 0 1 0 0 0 l 1 0 0 1 thomasi 2 0 0 0 l l 0 l 0 0 2 0 1 yemenensis l 0 0 0 1 0 0 0 l l 0 0 1 Leiolepis 0 0 0 0 0 0 0 0 0 0 0 0 0 Bonner zoologische Beitráge 56 (2007) Table 2. List of samples used for genetic analysis (geographic origin, locality and GenBank accession numbers). Species Geographic origin Tympanocryptis tetraporophora Australia E105.13 Saara asmussi Iran E107.5 Saara loricata Iran E107.3 Saara hardwickii unknown E111.17 Saara hardwickii unknown E111.18 Saara hardwickii unknown E111.19 Saara hardwickii unknown E111.20 Saara hardwickii unknown E112.2 Uromastyx acanthinura Tunesia E105.21 Uromastyx acanthinura Tunesia E105.22 Uromastyx acanthinura Tunesia E105.23 Uromastyx acanthinura Tunesia E107.15 Uromastyx nigriventris Morocco E106.4 Uromastyx nigriventris Morocco E106.5 Uromastyx nigriventris Morocco E107.14 Uromastyx dispar dispar Chad E106.2 Uromastyx dispar dispar Chad E110.19 Uromastyx dispar flavifasciata Mauritania E105.15 Uromastyx dispar flavifasciata Mauritania E105.27 Uromastyx dispar flavifasciata Mauritania E106.22 Uromastyx dispar flavifasciata Algeria E110.8 Uromastyx dispar flavifasciata Algeria E110.9 Uromastyx dispar flavifasciata Mauritanıa (obscura- phenotype) E111.21 Uromastyx dispar flavifasciata Mauritania (obscura-phenotype) E111.22 Uromastyx dispar flavifasciata Mauritania (obscura- phenotype) E133.2 Locality Mount Olga 30 km north of Bampur, direction to Zahedan (Pakistan) Voucher ZFMK 83840 NMP6V 73519 Chahak, approx. 15 km north ZFMK 87396 of Bandar-e-Genaveh, Province Busheer unknown unknown unknown unknown unknown unknown unknown unknown unknown unknown Guelmim Guelmim Zouar, Tibesti Mountains Zouar, Tibesti Mountains Captive bred; father from vicinity of Atar, mother from vicinity of Akjoujt Captive bred; father from vicinity of Atar, mother from vicinity of Akjoujt Atar Tindouf Tindouf Vicinity of Atar northern Mauritania 33 km southwest of Choum, er 0036°N/13.1347°W) ZFMK 83797 ZFMK 83795 ZFMK 83794 ZFMK 83796 No Voucher ZFMK 83816 ZFMK 83817 ZFMK 83818 No Voucher ZFMK 83820 ZFMK 83819 ZFMK 84438 ZFMK 84800 ZFMK 84437 ZFMK 85163 ZFMK 83824 ZFMK 73500 ZFMK 84261 ZFMK 84262 ZFMK 86473 ZFMK 86474 No Voucher GenBank Accession No. EF081041 (16s) FJ639585 (16s) FJ639586 (16s) FJ639587 (16s) FJ639588 (16s) FJ639589 (16s) FJ639590 (16s) FJ639591 (16s) FJ639630 (12s) FJ639592 (16s) FJ639631 (12s) FJ639593 (16s) FJ639632 (12s) FJ639594 (16s) FJ639633 (12s) FJ639595 (16s) FJ639634 (12s) FJ639596 (16s) FJ639635 (12s) FJ639597 (16s) FJ639636 (12s) FJ639598 (16s) FJ639637 (12s) FJ639599 (16s) FJ639638 (12s) FJ639600 (16s) FJ639639 (12s) FJ639605 (16s) FJ639640 (12s) FJ639601 (16s) FJ639641 ( FJ639602 ( FJ639642 ( FJ639603 ( FJ639643 ( FJ639604 (16s) FJ639644 (12s) FJ639606 (16s) FJ639645 (12s) FJ639610 (16s) FJ639648 (12s) 12s) 16s) 12s) 16s) 12s) 16s FJ639607 (16s) 88 Thomas M. WiLms et al.: On the Phylogeny and Taxonomy of the Genus Uromastyx Merrem, 1820 Species Uromastyx dispar flavifasciata (obscura- phenotype) E133.3 Uromastyx dispar flavifasciata (obscura-phenotype) E133.6 Uromastyx dispar flavifasciata (obscura-phenotype) E133.7 Uromastyx dispar flavifasciata (obscura-phenotype) E133.8 Uromastyx dispar flavifasciata (obscura- phenotype) E133.9 Uromastyx dispar flavifasciata (obscura-phenotype) E133.10 Uromastyx dispar flavifasciata (obscura-phenotype) E133.11 Uromastyx dispar maliensis E106.26 Uromastyx geyri E105.24 Uromastyx geyri E105.25 Uromastyx aegyptia aegyptia E106.21 Uromastyx aegyptia microlepis E117.7 Uromastyx aegyptia microlepis E117.11 Uromastyx aegyptia leptieni E106.27 Uromastyx aegyptia leptieni E.110.14 Uromastyx benti E106.3 Uromastyx benti E111.2 Uromastyx benti E111.4 Uromastyx benti E111.13 Uromastyx vemenensis Elll.12 Uromastyx yemenensis E106.18 Uromastyx yemenensis E106.19 Uromastyx vemenensis E106.20 Uromastyx shobraki Elll.l Uromastyx shobraki E111.3 Uromastyx shobraki E111.6 Uromastyx shobraki E111.7 Uromastyx macfadyeni E112.1 Geographic origin ia 7 Mauritania Mauritania Mauritania Mauritania Mauritania Mauritania Mali Niger Niger Egypt Saudi Arabia Saudi Arabia United Arab Emirates United Arab Emirates Oman Oman Oman Oman Yemen Yemen Yemen Yemen Yemen Yemen Yemen Yemen Somalia Locality Voucher Track Aghmakoum - El Beyed No Voucher (21°28’23” N/11° 33’ 24” W) south of Choum at 21.0027°N/13.1636°W south of Choum at 21.0027°N/13.1636°W 33 km southwest of Choum (21.0043°N/13.1324°W) 26 km northwest of Atar (20.7462°N/13.1293°W) track Atar - Choum (21.0003°N / 13.1598°W) 26 km northwest of Atar (20.7432°N/13.1183°W) unknown Kafadek, near Agadez Kafadek, near Agadez Sinai Peninsula Mahazat as Sayd Mahazat as Sayd Rimah / Al-Kaznah Wadi Siji Dhofar, vicinity of Mirbat Dhofar, vicinity of Mirbat Dhofar, vicinity of Mirbat Dhofar, vicinity of Mirbat Abian, southern Yemen unknown unknown unknown Mocca, northern Yemen Mocca, northern Yemen Between Mafraq and Mocca, northern Yemen Mocca, northern Yemen unknown No Voucher No Voucher No Voucher No Voucher No Voucher No Voucher ZFMK 71647 ZFMK 83821 ZFMK 83822 ZFMK 83792 ZFMK 86573 ZFMK 86567 No Voucher ZFMK 52398 Holotype ZFMK 83801 ZFMK 73681 ZFMK 83347 ZFMK 73680 ZFMK 47861 Holotype ZFMK 83805 ZFMK 83806 ZFMK 83807 ZFMK 73677 ZFMK 73676 ZFMK 48681 Holotype ZFMK 73675 ZFMK 84441 GenBank Accession No. FJ639649 (12s) FJ639608 (16s) FJ639650 (12s) FJ639612 (16s) FJ639651 (12s) FJ639613 (16s) FJ639652 (12s) FJ639614 (16s) FJ639653 (12s) FJ639609 (16s) FJ639646 (12s) FJ639615 (16s) FJ639647 (12s) FJ639611 (16s) FJ639616 (16s) FJ639654 (12s) FJ639617 (16s) FJ639655 (12s) FJ639618 (16s) FJ639656 (12s) FJ639619 (16s) FJ639658 (12s) FJ639620 (16s) FJ639657 (12s) FJ639621 (16s) FJ639659 (12s) FJ639622 (16s) FJ639660 (12s) FJ639623 (16s) EF081054 (16s) EF081055 (16s) EF081056 (16s) EF081057 (16s) EF081058 (16s) EF081059 (16s) EF08 1060 (16s) EF081061 (16s) EF081065 (16s) EF081066 (16s) EF081067 (16s) EF081068 (16s) EF081042 (16s) Species Uromastyx macfadyeni E112.3 Uromastyx ocellata E106.6 Uromastyx ocellata E106.7 Uromastyx ornata ornata E106.11 Uromastyx ornata ornata E106.8 Uromastyx ornata ornata E106.9 Uromastyx ornata philbyi E110.20 Uromastyx princeps E106.24 Uromastyx princeps E106.25 Uromastyx thomasi E105.4 Uromastyx thomasi E105.11 Uromastyx thomasi E105.12 Bonner zoologische Beitrage 56 (2007) Geographic origin Somalia u Sudan Sudan Egypt Egypt Egypt Saudi Arabia Somalia Somalia Oman Oman Oman Locality unknown unknown unknown Sinai Peninsula Sinai Peninsula Sinai Peninsula 19°05’N 41°50’E, Tihama Bossasso Bossasso Vicinity of Ras Hilf, Masirah Island Vicinity of Ras Hilf, Masirah Island Vicinity of Ras Hilf, Masirah Island Voucher ZFMK 84440 ZFMK 83798 ZFMK 83799 ZFMK 83815 ZFMK 83812 ZFMK 83813 ZFMK 84442 ZFMK 58985 ZFMK 58048 ZFMK 83830 ZFMK 83837 ZFMK 83838 89 GenBank Accession No. EF081043 (16s) EF08 1044 (16s) EF081045 (16s) EF081051 (16s) FJ639629 (12s) EF081052 (16s) EF081053 (16s) EF08 1046 (16s) FJ639624 (16s) FJ639625 (16s) FJ639626 (16s) FJ639627 (16s) FJ639628 (16s) Table 3. Variables used to calculate the distance phenogram (Fig. 3); for definition of appreviations see “Material and Methods”. Variable vi v2 V3 v4 V5 V6 V7 V8 v9 Definition W SD G MBS V SW PP left PP right LS left Variable v10 vil v12 v13 v14 v15 v16 v17 v18 Definition LS right SO left SO right HS left HS right ES * PES IS TE TD Table 4. Definition of variables used for the PCA separating OTU I and OTU I; for definition of abbreviations see “Material and Methods”. Variable v1 v2 v3 v4 V5 V6 V7 v8 v9 Definition W SD G MBS Vv SW PP left PP right LS left Variable v10 vil v12 v13 v14 v15 V16 v17 Definition SO left IS TD SO right HS left HS right ES * PES 90 Thomas M. WiLms et al.: On the Phylogeny and Taxonomy of the Genus Uromastyx Merrem, 1820 Table 5. Factor loading on the first four principal components (PC) from a correlation matrix of V1-V17 for individuals of OTU I and OTU H. Variable PC 1 PC 2 PC 3 PC 4 vi - 0.066 0.293 - 0.092 0.073 v2 0.189 - 0.062 - 0.159 0.041 v3 0.194 - 0.147 0.089 - 0.040 v4 0.199 - 0.092 - 0.009 - 0.050 v5 0.176 - 0.066 0.021 - 0.031 V6 0.022 0.205 - 0.133 0.157 v7 - 0.050 - 0.021 0.456 - 0.017 v8 - 0.050 - 0.018 0.458 - 0.027 v9 - 0.033 0.022 - 0.031 0.462 V10 - 0.035 0.029 - 0.014 0.458 Vil 0.045 0.131 0.017 0.022 v12 0.064 0.115 - 0.011 0.050 v13 0.162 - 0.035 0.016 - 0.023 v14 0.167 - 0.050 0.022 - 0.026 V15 0.129 0.068 - 0.153 - 0.042 V16 - 0.116 - 0.324 0.033 - 0.016 V17 - 0.128 0.264 0.166 - 0.206 Eigenvalues 7,264 2,512 1,939 1,560 Accumulated percent of trace 42,731 57,508 69,031 78,207 Table 6. Definition of variables used for the PCA separating species and species groups within Uromastyx (Figs 5-10); for defi- nition of abbreviations see “Material and Methods”. Variable Vi v2 v3 V4 V5 V6 V7 Definition W SD G MBS V SW PP left Variable v8 v9 V10 Vil v12 v13 v14 v15 Definition PP right LS left ES right SO left SO right HS left HS right ES * PES Bonner zoologische Beitráge 56 (2007) 91 Table 7. Factor loading on the first four principal components (PC) from a correlation matrix of V1-V15 for 431 individuals of Uromastyx. Variable PC Il o PC? PC 3 PC 4 PC 5 v1 0.274 - 0.216 0.598 0.246 0.581 v2 0.630 031 - 0.265 0.433 - 1.898E-02 v3 0.850 7.232E-02 - 0.219 9.518E-02 - 0.168 v4 0.841 - 0.148 - 0.133 0.346 7.417E-02 v5 0.824 - 6.121E-02 - 0.234 0.365 2.631E-02 v6 0.622 - 0.105 0.475 0.148 9.023E-02 v7 0.433 0.715 - 0.314 - 0.134 0.403 v8 0.439 0.702 - 0.316 0.137 0.419 v9 0.147 0.753 0.402 0.261 - 0.295 V10 0.166 0.785 0.358 0.239 - 0.288 Vil 0.742 -2.223E-02 0.308 - 0.430 8.690E-02 v12 0.755 -1.883E-02 0.281 - 0.439 - 4.109E-02 v13 0.857 - 5.430E-02 - 8.995E-02 - 0.220 - 0.253 v14 0.852 - 3.638E-02 - 9.704E-02 - 0.200 0.278 vis 0.649 - 0.488 0.115 - 8.413E-02 6.694E-03 Eigenvalues 6399-2615 IT Accumulated percent of trace 42.662 60.097 69.870 77.646 84.560 Table 8. Factor loading on the first three principal components (PC) from a correlation matrix of VI-V15 for 354 individuals of Uromastyx. Variable PC 1 PC 2 PC 3 vi 0.337 - 0.403 - 0.427 v2 0.728 - 5.468E-02 - 0.354 V3 0.884 0.147 1.143E-02 v4 0.872 - 2.405E-02 - 0.240 V5 0.860 4.975E-02 - 0.246 V6 0737 0.105 2.198E-02 v7 0.603 0.500 - 0.418 v8 0.611 0.442 - 0.446 v9 6.418E-02 0.831 0.311 v10 7.056E-02 0.835 0,323 Vil 0.715 - 0.282 0.449 v12 0.724 - 0.228 0.504 v13 0.848 - 7.847E-02 0.314 v14 0.852 -4.002E-02 0.308 V15 0.715 - 0.381 8.754E-02 Eigenvalues 7.238 2.318 1.658 Accumulated percent of trace 48.250 63.706 74.756 92 Thomas M. WiLms et al.: On the Phylogeny and Taxonomy of the Genus Uromastyx Merrem, 1820 Table 9. Factor loading on the first four principal components (PC) from a correlation matrix of V1-V15 for 331 individuals of Uromastyx. PC 1 PC 2 PES PC 4 Variable v1 0.363 0.221 4.585E-02 0.809 v2 0.727 0.130 - 0.320 0.144 V3 0.894 5.650E-02 - 1.491E-02 - 6.992E-02 v4 0.874 0.117 - 0.139 0.289 V5 0.861 0.150 - 0.167 0.235 V6 0.765 - 1.862E-02 - 0.121 - 0.329 v7 0.630 0.532 - 0.327 - 0.247 V8 0.626 0.520 - 0.335 - 0.256 v9 0.193 0.607 0.664 - 5.942E-02 v10 0.211 0.599 0.653 - 6.505E-02 vi 0.711 - 0.462 0.288 - 6.479E-03 v12 0.724 - 0.496 0.260 - 3.671E-02 v13 0.850 - 0.282 0.130 - 0.121 V14 0.855 - 0.245 0.146 - 0.104 V15 0.749 - 0.280 0.145 3.519E-02 Eigenvalues 7.463 2.064 1.462 1.089 Accumulated percent of trace 49.751 63.512 73.262 80.523 Table 10. Factor loading on the first three principal components (PC) from a correlation matrix of VI-V7 & V10-V15 for 317 individuals of Uromastyx (because V8 & V9 are coding for preanofemorapores, it is justified to exclude these variables in the PCAs dealing exclusively with species possessing preanofemoralpores). Variable o EG 0... PC2 _PC3 vi 0.271 0.620 - 0.564 v2 0.766 0.213 - 0.369 V3 0.879 9.939E-02 1.397E-02 v4 0.852 0.249 - 0.281 V5 0.849 0.265 - 0.270 V6 0.786 - 5.101E-02 4.997E-02 V7 0.137 0.684 0.601 V10 0.162 0.695 0.581 Vil 0.751 - 0.359 0.234 v12 0.764 - 0.406 0.226 v13 0.870 - 0.202 0.148 v14 0.873 - 0.161 0.158 v15 0.768 - 0.164 8.008E-02 Eigenvalues 6.799 1.912 1.468 Accumulated percent of trace 52.301 67.010 78.299 Bonner zoologische Beitráge 56 (2007) 93 Table 11. Factor loading on the first three principal components (PC) from a correlation matrix of VI-V7 & V10-V15 for 265 individuals of Uromastyx (because V8 & V9 are coding for preanofemorapores, it is justified to exclude these variables in the PCAs dealing exclusively with species possessing preanofemoralpores). PCS Variable PC 1 PC 2 v1 - 0.289 0.578 - 0.458 V2 0.486 0.328 - 0.502 V3 0.774 0.190 0.127 v4 0.677 0.310 - 0.328 V5 0.704 0.352 - 0.309 V6 0:757 0.151 4.432E-02 v7 - 5.536E-02 0.618 0.619 v10 - 3.002E-02 0.684 0.576 Vil 0.716 - 0.397 0.239 v12 0.764 - 0.384 0.210 v13 0.895 - 2.617E-02 9.960E-02 v14 0.890 1.378E-02 0.130 v15 0.747 - 8.992E-02 - 8.770E-02 Eigenvalues 5.694 1.884 1.534 Accumulated percent of trace 43.804 58.295 70.092 Table 12. Factor loading on the first three principal components (PC) from a correlation matrix of VI-V7 & V10-V15 for 223 individuals of Uromastyx (because V8 & V9 are coding for preanofemorapore, it is justified to exclude these variables in the PCAs dealing exclusively with species possessing preanofemoralpores). Variable PC 1 PC 2 PC3 vi - 0.554 0.490 -0.118 v2 - 9.640E-02 0.494 - 0.325 v3 0.743 0.315 -0.112 v4 0.396 0.515 - 0.282 v5 0.417 0.529 - 0.308 v6 0.662 0.239 - 4.877E-02 v7 0.140 0.497 0.741 v10 0.110 0.580 0.680 Vil 0.656 - 0.491 0.209 v12 0.707 - 0.495 0.147 V13 0.866 4.393E-02 - 0.100 v14 0.876 8.752E-02 - 9.225E-02 vis 0.563 0181 1.228E-02 Eigenvalues E 4.434 2283 1.405 Accumulated percent of trace 34.108 51.667 62.473 94 Thomas M. WILMs et al.: On the Phylogeny and Taxonomy of the Genus Uromastyx Merrem, 1820 APPENDIX II. LIST OF EXAMINED SPECIMENS Saara asmussi (Strauch, 1863) MHNP 1989.3005, unknown; ZFMK 7925, Afghanistan, Seistan, 50 km east Seranj; BMNH 1964.279, Pakistan, Kharan / Balut- schistan; BMNH 74.11.239, Iran, near Rigan / Narmashir; BMNH 79.8.15.18, Afghanistan, Ghorak; BMNH 79.8.15.30, Afghanistan, Ghorak. Saara hardwickii (Gray, 1827) NMW 21175, Pakistan, Sindh; NMW 21167:1, India, Katchh; NMW 21167:2, India, Katch; NMW 21167:3, India Katchh: NMW 21173, unknown, foothills of the western Himalaya; NMW 21169:1, Pakistan, Sindh; NMW 21169:2, Pakistan, Sindh; NMW 21169:3, Pakistan, Sindh; NMW 15121:2, India, Katchh; NMW 19981:2, Pakistan, Karachi: NMW 19981:3, Pa- kistan, Karachi; NMW 19981:4, Pakistan, Karachi; NMW 19981:5, Pakistan, Karachi; MHNP 1962.726, Pakistan, Gizri; MHNP 1962.727, Pakistan, Gizri, MHNP 1962.728, Pakistan, Gizri; ZSM 327/79, Pakistan, Uthal; ZSM 20/1912, Pakistan, Habb; ZSM 7/1912, Pakistan, Wajara; ZSM 7/1912, Pakistan, Wajara; ZFMK 21453, Pakistan, Mokran coast; ZFMK 8616, Af- ghanistan, Nimla to Djalalabad; ZFMK 21454, Pakistan, Mo- kran coast; ZFMK 21455, Pakistan, Mokran coast; ZFMK 22103, Pakistan, Mokran coast; BMNH 1933.4.1.23, India, Thar Parkar/Rajputana; BMNH 1933.4.1.24, India, Thar Parkar/Raj- putana; BMNH 91.9.11.9, Pakistan, Sindh; BMNH 60.3.19.1006, India, Goojerat; BMNH 1973.447, Pakistan, Karachi; BMNH 1946.8.14.44, India, Plains of Kanouge/Hindustan; BMNH 98.12.22.10, Pakistan, Karachi. Saara loricata (Blanford, 1874) NMW 21177, Iran, Bushir; NMW 21179:1, Iran, Bushir; NMW 21179:2, Iran, Bushir; ZSM 2/1966, Iraq, Chankin / southeast Bagdad; ZFMK 22072, Iran, Ahwaz; ZFMK 44906, Iraq, Kir- kuk; ZFMK 40594, Iraq, Kirkuk; ZFMK 40593, Iraq, Kirkuk; ZFMK 40592, Iraq, Kirkuk; ZFMK 40591, Iraq, Kirkuk; ZFMK 40590, Iraq, Kirkuk; BMNH 1933.4.1.25, Iran, Bushir; BMNH 87.9.22.19, Iran, Bushir; BMNH 1905.10.14.21, Iran, 30 mls northwest Ahwaz. Uromastyx acanthinura Bell, 1825 BMNH 1907.4.6.14, Algeria, Biskra; BMNH 1907.4.6.15, Al- geria, Biskra, BMNH 1912.11.9.4.6., Algeria, Fort Miribel; BMNH 1912.11.9.50, Paratype of Uromastix acanthinurus ni- gerrimus, Algeria, Oued Mya; BMNH 1912.11.9.51, Paratype of Uromastix acanthinurus nigerrimus, Algeria, Ain Guettara; BMNH 1938.7.5.10, Algeria, Biskra; BMNH 1938.7.5.11, Al- geria, Bistra; BMNH 1938.7.5.12, Algeria, Bistra; BMNH 1938.7.5.13, Algeria, Biskra; BMNH 1964.2075, Libya, Ain Uif Jebel Nefrousa; BMNH 1969.2088, Algeria, Bistra; BMNH 1969.2090, Algeria, Bistra; BMNH 1969.2091, Algeria, Bistra; BMNH 1969.2106, Algeria, Touggourt; BMNH 1969.2107, Al- geria, Touggourt; BMNH 1969.2108, Algeria, Touggourt: BMNH 71.4.16.52, Algeria, Bistra; BMNH 91.4.5.41, Algeria, Bistra; BMNH 91.4.5.42, Algeria, Biskra; BMNH 91.5.439, Tu- nesia, Duirat; BMNH 91.5.440, Tunesia, Duirat; BMNH 96.2.29.1, Tunesia; Duirat; MZUF 13757, Algeria, Touggourt (Tourghuf); MZUF 21666, Somalia, unreliable locality!; MZUF 25125, Tunesia, Tamerza/Gafsa; MZUF 25126, Tunesia, Ta- merza/Gafsa; MZUF 743, Libya, Bu Ngem; MZUF 744, Libya, Cirenaica (Barqa), NMW 21192:1, Tunesia, Gafsa; NMW 21192:2, Tunesia, Gafsa; NMW 21192:3, Tunesia, Gafsa; NMW 21197:1, Algeria, Bistra; NMW 21198:1, Tunesia, Gafsa; NMW 21198:10, Tunesia, Gafsa; NMW 21198:2, Tunesia, Gafsa; NMW 21198:3, Tunesia, Gafsa; NMW 21198:4, Tune- sia, Gafsa; NMW 21198:5, Tunesia, Gafsa; NMW 21198:6, Tu- nesia, Gafsa; NMW 21198:7, Tunesia, Gafsa; NMW 21198:8, Tunesia, Gafsa; NMW 21198:9, Tunesia, Gafsa; NMW 21202: 1, Algeria, Biskra; NMW 21202:2, Algeria, Bistra; NMW 21207, Tunesia, Gafsa; NMW 21208, Tunesia, El Guietar; NMW 22116, Libya, Tripolis; SMNS 602: 1, Algeria, Biskra; ZFMK 2707, Tu- nesia, Gabes (Quderef); ZFMK 2708, Tunesia, Gabes-Matmata; ZFMK 2709, Algeria, Biskra; ZFMK 2710, Algeria, Biskra; ZFMK 2711, Algeria, Biskra; ZFMK 2714, Algeria, Biskra; ZMH-R4507, Algeria, Biskra; ZMH-R4508, Algeria, Biskra; ZMH-R4509, Algeria, Biskra; ZSM 112/1983, Libya, south of Tripolis; ZSM 18/1968 (1), Libya, Jebel el Soda; ZSM 18/1968 (2), Libya, Jebel el Soda; ZSM 18/1968 (3), Libya, Jebel el Soda; ZSM 18/1968 (4), Libya, Jebel el Soda; ZSM 181/36, Libya, Je- bel el Soda; ZSM 26/1951 (1), Tunesia, Nefta south of Tozeur: ZSM 26/1951 (2), Tunesia, Nefta south of Tozeur; ZSM 4/1963, Tunesia, El Hamma; ZSM 472/79, Libya, Gharian/Tripolis; ZSM 510/1978, Libya, Wadi Bundindin. Uromastyx aegyptia Uromastyx aegyptia aegyptia Forsskal, 1775 BMNH 1908.6.9.6, Egypt, Tor / Sinai; BMNH 1951.1.2.55, Is- rael, Wadi Araba; BMNH 97.10.28.212, Egypt, Suez; BMNH 97.10.28.213, Egypt, Beltim Delta; MZUF 28899, Saudi Ara- bia, Sawawin; NMW 21182:1, Egypt, Cairo; NMW 21182:2, Egypt, Desert near Cairo; NMW 21183, Egypt, Suez; NMW 21187, Egypt, Cairo; NMW 21222, Egypt, Beltim; ZFMK 2703, Egypt, Lower Egypt; ZFMK 2704, Egypt, Lower Egypt: ZFMK 39073, Egypt, Suez; ZFMK 44216, Neotype of Uromastyx ae- gyptia aegyptia, Egypt, Suez; ZFMK 46502, Egypt; ZFMK 46504, Egypt; ZFMK 64405, Egypt, vicinity of Hurgharda; ZFMK 64406, Jordan, Wadi Araba. Uromastyx aegyptia microlepis Blanford, 1874 BMNH 1930.6.30.3, Yemen, Bin Khautar / Hadramaut; BMNH 1946.8.11.67, Iraq, Paralectotype of Uromastyx microlepis, Bas- rah; BMNH 1946.8.14.55, Lectotype of Uromastyx microlepis, Iraq, Basrah; BMNH 1950.1.4.71, Oman; BMNH 1950.1.5.4, Arabia, Miofa; BMNH 1952.1.3.51, Saudi Arabia, ElGaisuma- Turaif; BMNH 1953.1.8.50, Yemen, North of Jol / Hadramaut; BMNH 1970.2076; Saudi Arabia, Ruma; BMNH 1970.2481, Bonner zoologische Beitráge 56 (2007) 95 Bahrain, Ras Al Barr; BMNH 1970.2482, Bahrain, Ras Al Barr; BMNH 1971.748, Bahrain; BMNH 1972.1259, United Arab Emirates, Al Hamran / Abu Dhabi; BMNH 1972.833, United Arab Emirates, near Bada Zaid / Abu Dhabi; BMNH 1978.2072, Kuwait; BMNH 1980.569, Oman, Jiddat al Harasis; BMNH 1982.1327, Saudi Arabia, Dib Dibah; BMNH 1982.1328, Saudi Arabia, Dib Dibah; BMNH 1985.880, Saudi Arabia, 30 km SE Ronya; BMNH 1986.435, Saudi Arabia, Shigree; BMNH 1988.214, Saudi Arabia, 26°56’N, 38°59’E; BMNH 1988.93, Saudi Arabia, Al Rawdah, north of Khobar; BMNH 1996.207, United Arab Emirates, Jebel Gaddah near Jebel Dannah; BMNH 85.7.11.11, Iraq, Fao; BMNH 88.12.6.8, Iraq, Fao; ZFMK 20267, Iraq, Basrah; ZFMK 21091, Iraq, Basrah; ZFMK 42413, Oman, 100 km from Muscat; ZFMK 42414, Oman, 100 km from Muscat; ZFMK 43648, Saudi Arabia, 100 km NO Riyadh; ZFMK 43649, Saudi Arabia, 100 km NO Riyadh; ZFMK 44907, Iraq, Kirkuk; ZFMK 44908, Iraq, Kirkuk; ZFMK 44909, Iraq, Kirkuk; ZFMK 44910, Iraq, Kirkuk; ZFMK 44911, Iraq, Kirkuk. Uromastyx aegyptia leptieni Wilms € Bóhme, 2000 BMNH 1973.2039, United Arab Emirates, S Jebel Jayah; BMNH 1973.2040, United Arab Emirates, Jebel Ali SW Dubai; BMNH 1973.2041, United Arab Emirates, Tawi Bil Khabis 25km WSW Dayd; BMNH 1973.721, Oman, Munay; BMNH 1975.958, Oman, Rostaq 23°24’N 57°27’E; BMNH 85.11.7.4, Oman, Mus- cat; BMNH 85.11.7.5, Oman, Muscat; ZFMK 52398, Holotype Uromastyx leptieni, Wadi Sij1. Uromastyx alfredschmidti Wilms & Böhme, 2001 MHNG 1515.77, Paratype of Uromastyx alfredschmidti, Alge- ria, Tassili n’Ajjers; MHNP 1961.261, Paratype of Uromastyx alfredschmidti, Algeria, Hoggar; MHNP 9905, Paratype of Uro- mastyx alfredschmidti, Algeria, Hoggar; NMW 8224:3, Paratype of Uromastyx alfredschmidti, Sahara; ZFMK 24643, Holotype of Uromastyx alfredschmidti, Algeria, Tassili n’Ajjers /30 km NO Djanet. Uromastyx benti (Anderson, 1894) BMNH 1946.8.11.69, Paralectotype of Uromastyx benti, Yemen, Makulla, Hadramaut; BMNH 1946.8.11.70, Paralectotype of Uromastyx benti, Yemen, Makulla, Hadramaut; BMNH 1946.8.11.71, Paralectotype of Uromastyx benti, Yemen, Ma- kulla, Hadramaut; BMNH 1946.8.11.72, Lectotype of U. benti, Yemen, Makulla, Hadramaut; BMNH 1953.1.8.52, Yemen, Ha- dramaut; BMNH 1956.1.7.26, Yemen, Wadi Abr/ Hadramaut; MHNP 1895.43, Paralectotype of Uromastyx benti, Yemen, Ma- kulla, Hadramaut; MTKD 24589, Yemen, Makulla; NMW 16174, Yemen, Makulla; NMW 21213:1, Yemen, Makulla; NMW 21213:2, Yemen, Makulla; NMW 21213:3, Yemen, Ma- kulla; NMW 21213:4, Yemen, Makulla; NMW 21213:5, Yemen, Makulla; NMW 21213:6, Yemen, Makulla; NMW 21213:7, Ye- men, Makulla; NMW 21213:8, Yemen, Makulla; NMW 21213:9, Yemen, Makulla; NMW 21214:1, Syntype Uromastyx simonyi, Yemen, Assan; NMW 21214:2, Syntype Uromastyx si- monyi, Yemen, Assan; ZFMK 73680, Oman, Mirbat; ZFMK 73681, Oman, Mirbat; ZFMK 83347, Oman, Mirbat; ZFMK 83801, Oman, Mirbat; ZMH R04513, Yemen, Makulla/Hadra- maut. Uromastyx dispar Uromastyx dispar dispar Heyden, 1827 BMNH 1900.9.12.1, Sudan, Wadi Halfa; BMNH 1913.9.16.15, Sudan, Dongola Provinz; BMNH 1954.1.6.9, Tchad, Tibesti; BMNH 1956.1.1.6, Tchad, S of Zouar; BMNH 1958. 1.3.95, Tchad, Central Tibesti; BMNH 1962.286, Tchad, Bardai; BMNH 1962.287, Tchad, Bardai; BMNH 1973.3348, Tchad, Central Tibesti; BMNH 1986.721, Tchad, Ounianga 19°4’N/20°36’E; GMNH 1952-9100(1), Tchad, Tibesti; GMNH 1952-9100(2), Tchad, Tibesti; MHNP 1974.328, Egypt, Ismai- lia; MNHP 1993.0692, Tchad, Tibesti; BMNH 1900.9.12.1, Su- dan, Wadi Halfa; BMNH 1913.9.16.15, Sudan, Wadi Halfa; SMF10417, Lectotype of Uromastyx dispar, Desert near Am- bukol; ZFMK 2706, Tchad, Fada; ZFMK 39900, Sudan, SE Debba direction of Khartoum; ZFMK 65600, Tchad, Zouar; ZFMK 65601, Tchad, Zouar; ZFMK 65602, Tchad, Zouar. Uromastyx dispar flavifasciata Mertens, 1962 BMNH 1969.476, Mauritania, Ouadane; BMNH 1969.477, Mau- ritania, Ouadane; MHNG 1515.74, Mauritania, Guelta Zemour/Rio de Oro; MHNG 1515.75, Mauritania, Chingetti; MHNG 1515.76, Mauritania, Bir Moghrein/Fort Tringuet; MHNP 1981.178, Mauritania, Richat, Adrar; MHNP 1986.2012, Algeria, Tindouf; MHNP 1993.1501, Mauritania, Atar, Ar bou M’rait; MNHP 1993.5808, Mauritania, Matmata; SMF 58032, Holotype of Uromastyx acanthinurus flavifasciatus, Mauritania; ZFMK 17597, Mauritania, Hamdoun; ZFMK 17598, Maurita- nia, Atar. Uromastyx dispar maliensis Joger & Lambert, 1996 BMNH 1933.11.18.1, Mali, Gao; BMNH 1933.11.18.2, Mali, Gao; BMNH 1934.1.1.1, Mali, Taberreshat/17°40’N/0°10’E; GNHM 1930.32-5744 RE1772, Paratype Uromastyx maliensis, Mali Ti-N-Zaouatene; HLMD RA 1545, Holotype Uromastyx maliensis, Mali 40km SE Gao; MHNP 1965.0144, Paratype Uro- mastyx maliensis, Algeria, Taoudrart/Hoggar; NMW 21211, Al- geria, Tamanrasset; ZFMK 9232, Algeria, Ahaggar/Gara Dje- noun; ZMH-R04529, Algeria, Tassili du Hoggar; 11 Specimens from the trade (Mali). Uromastyx geyri Miller, 1922 BMNH 1961.417, Niger, near Abangharit/Air; BMNH 1970.1755, Niger, Iferouhane/Air; BMNH 1978.2093, Algeria, Hoggar; BMNH 1978.2094, Algeria, Hoggar; BMNH 1979.402, Algeria, 15km ENE Tamanrasset 22°57’N/05°47’E; BMNH 1986.733, Niger, N of Agadez; MZUF 21013, Algeria, Taman- rasset; MZUF 21014, Algeria, Tamanrasset; MZUF 21015, Al- geria, Tamanrasset; MZUF 21017, Algeria, In Ecker 180km N Tamanrasset; GNMH 1930.32-5744 Re1761; MHNG 1513.40, Niger, Tin Teloust/Air; MHNP 1932.128, Algeria, Tanezrouft; MHNP 1943.3, Algeria, Hoggar; MHNP 1974.1412, Algeria, Te- fedest; MHNP 1990.4665, Niger, El Meki/Air; MNHP 8971, Ni- ger, Telouess Tabelot; MNHP 8972, Niger, Telouess Tabelot; MTKD 25699, Niger, Agadez; MTKD 25700, Niger, Agadez; SMF 68765, Niger; SMF 68766, Niger; SMF 68767, Niger; ZFMK 9228, Algeria, Ahaggar/In Kelmet; NMW 21210, Alge- ria, Tamanrasset; NMW 21211, Algeria, Tamanrasset; NMW 22000, Algeria, Hoggar/Tamanrasset; NMW 23517:1, Algeria, 96 Thomas M. WiLMms et al.: On the Phylogeny and Taxonomy of the Genus Uromastyx Merrem, 1820 Ideles/Hoggar; NMW 23517:2, Algeria, Ideles/Hoggar; NMW 25481, Algeria, Tit near Tamanrasset; ZFMK 20042, Algeria, Hoggar/between In Eker and In Amguel; ZFMK 20043, Niger, 20 km S Arlit/Air; ZFMK 36627, Niger, 30-40 km N Gouga- ran; ZFMK 36628, Niger, 30-40km N Gougaran; ZFMK 40628, Niger, near Gougaran; ZFMK 9226, Paratype of Uromastyx geyri, Algeria, upper Tahihaout; ZFMK 9227, Paratype of Uro- mastyx geyri, Algeria, Oued Ouhat; ZFMK 9229, Algeria, Thar- emert-n-akh; ZFMK 9230, Neotype of Uromastyx geyri, Alge- ria, Gara Dienoum; ZFMK 9231, Algeria, Ahaggar/Gara Dje- noum; ZMH-R04523, Algeria, Oasis Abalessa E of Tamanras- set; ZSM 4451, Paratype of Uromastyx geyri, Algeria, Thar- emert-n-akh. Uromastyx macfadyeni Parker, 1932 BMNH 1946.8.14.52, Paratype of Uromastyx macfadyeni, So- malia, Dagah Shabell 24mls SE Berbera; BMNH 1946.8.14.54, Holotype of Uromastyx macfadyeni, Somalia, near Berbera; BMNH 1956.1.6.55, Somalia, Heis 20 mls W Mait. Uromastyx nigriventris Rothschild & Hartert, 1912 BMNH 1911.12.5.1, Paratype of Uromastyx acanthinurus nigri- ventris, Algeria, Ghardaia; BMNH 1911.12.5.2, Paratype of Uro- mastyx acanthinurus nigriventris, Algeria, Ghardaia; BMNH 1969.2073, Algeria, Laghouat; BMNH 1969.2074, Holotype of Uromastyx acanthinurus nigriventris, Algeria, Tilghempt bet- ween Laghouat and Ghardaia; BMNH 1969.2075; Algeria, La- ghouat; BMNH 1969.2080, Algeria, Ain Sefra; BMNH 1969.2085, Algeria, Laghouat; BMNH 1969.2086, Algeria, Ghardaia; BMNH 1969.2087, Algeria, Laghouat; BMNH 1969.2099, Algeria, Oued N’ca; BMNH 1969.2100, Algeria, Oued N’ca; BMNH 1969.2103, Algeria, (Oued N’ca)/Oued Mya; BMNH 1969.2109, Algeria, El Hadadra between El Golea and Ghardaia; BMNH 1970. 223, Morocco, Foum el Hassane; BMNH 1970. 224, Morocco, Foum el Hassane; BMNH 1970.220, Morocco, 3 km N Tuizgui-Remz, Tarfaya; BMNH 1970.221, Morocco, 5 km E Bou Izakarn; BMNH 1972.2280, Morocco, 3 km NNW Quarzazate; BMNH 1972.2281, Morocco, 3km NNW Quarzazate; BMNH 1972.2282, Morocco, 3km NNW Quarzazate; BMNH 1972.2283, Morocco, 2 km N Douar Zednagain, 3km NNW Quarzazate; BMNH 1972.2284, Mo- rocco, 2 km N Douar Zednagain, 3 km NNW Quarzazate; MZUF 21003, Algeria, Ghardaia; MZUF 21004, Algeria, Ghardaia; MZUF 21005, Algeria, Ghardaia; MZUF 21006, Algeria, Ghar- daia; MZUF 21007, Algeria, Ghardaia; MZUF 21008, Algeria, Ghardaia; MZUF 21009, Algeria, Ghardaia; MZUF 21010, Al- geria, Ghardaia; MZUF 21011, Algeria, Ghardaia; MZUF 21012, Algeria, Ghardaia; HLMD RA 1177, Morocco, SW Tizgui el Ha- ratuine; HLMD RA 1178, Morocco, 2 km N Rich; MHNP 1927.0094, Morocco, Colomb-Bechar; MHNP 1950.204, Alge- ria, Beni Ounif; MHNP 1953.18, Algeria, Zerhaura Indigene Beni Abbes; MHNP 1961.249, Morocco, Assa-Aouinet Torkos; MHNP 1961.250, Morocco, vicinity Doirat; MHNP 1961.251, Morocco, Foum el Hassane; MHNP 1961.252, Morocco, Foum el Hassane; MHNP 1961.253, Morocco, 10 km S Guercif; MHNP 1961.255, Morocco, Guercif; MHNP 1961.256, Mo- rocco, N Aouinet Torkos; MHNP 1961.257, Morocco, N Aoui- net Torkos; MHNP 1961.258; Morocco, Aoinet Torkos; MHNP 1961.259, Morocco, Zagora — Tagounite; MHNP 1961.260, Mo- rocco, Zagora — Tagounite; MHNP 1986.2013, Morocco, Bechar; MHNP 1986.2014, Morocco, Bechar; MHNP 1991.405, Mo- rocco, Quarzazate; MHNP 1993.800, Morocco: MHNP 1994.1199, Algeria, Beni Ounif; MHNP 1994.1205, Algeria, Beni Ounif; MHNP 1994.1207, Algeria, Beni Ounif; MHNP 1994.1208, Algeria, Beni Ounif; MHNP 1994.1209, Algeria, Beni Ounif; MTKD 18981, Algeria, Ain Sefra; MTKD 20205, Morocco; Quarzazate; MTKD 27995, Algeria, Ain Sefra; NMW 14895, Algeria, Ain Sefra; NMW 14896:1, Algeria, Ain Sefra; NMW 14896:10, Algeria, Ain Sefra; NMW 14896:2, Al- geria, Ain Sefra;, NMW 14896:3, Algeria, Ain Sefra; NMW 14896:4, Algeria, Ain Sefra; NMW 14896:5, Algeria, Ain Se- fra; NMW 14896:6, Algeria, Ain Sefra; NMW 14896:7, Alge- ria, Ain Sefra; NMW 14896:8, Algeria, Ain Sefra; NMW 14896:9, Algeria, Ain Sefra; NMW 21189:1, Algeria, Gardaia; NMW 21190:1, Morocco, Mazagan; NMW 21190:2, Morocco, Mazagan; NMW 21190:3, Morocco, Gus; NMW 21192:4, Al- geria, Beni Abbes; NMW 21192:5, Algeria, Beni Abbes; NMW 21195:2, Algeria, Beni Mzab; NMW 21199:1, Morocco, Co- lomb-Bechar; NMW 21199:2, Morocco, Colomb-Bechar; NU W 21200, Morocco, Aouinet-Torkoz; NMW 21204:2, Algeria, Beni Ounif/Figuig; NMW 21205, Morocco, Colomb-Bechar; NMW 21209:1, Algeria, Beni Abbes; NMW 21209:2, Algeria, Beni Ab- bes; NMW 34024, Morocco, Erfoud; SMF 58031, Morocco, Oued Moulonya/36 km E Guercif; SMF 69077, Morocco, Bou Afra near Figuig; SMNS 593:1, Algeria, Ain Sefra; SMNS 598, Algeria, Oued N’za; SMNS 599, Algeria, Beni Ounif/Ouuf; SMNS 601:1, Algeria, Ain Sefra; ZFMK 18021, Algeria, S Ghar- daia, Oued Sebseb; ZFMK 18022, Algeria, S Ghardaia, Oued Sebseb; ZFMK 2715, Algeria, Oued Mzab; ZFMK 2716, Alge- ria, Oued N’ca; ZFMK 2717, Algeria, Oued N’ca; ZFMK 2718, Algeria, Oued N’ca; ZFMK 2719, Algeria, Oued N’ca; ZFMK 2723, Morocco, Erfoud; ZFMK 2724, Morocco, Oujda; ZFMK 2725, Morocco, Oujda; ZFMK 2726, Morocco, Oujda; ZFMK 2727, Morocco, Oujda; ZFMK 2728, Morocco, Oujda; ZFMK 2729, Morocco, Oujda; ZFMK 30806, Morocco, 10 km W Ti- nerhir; ZFMK 41168, Algeria, Ain Sefra; ZFMK 41512, Mo- rocco, Goulmima; ZFMK 45946, Morocco, Quarzazate; ZFMK 49661, Algeria, El Homr; ZFMK 49742, Morocco, 10-15 km NW Quarzazate; ZFMK 51077, Algeria, Ain Sefra; ZFMK 51078, Algeria, Ain Sefra; ZFMK 52356, Morocco, south of Quarzazate, Ait-Saon; ZFMK 59062, Morocco, Quarzazate; ZFMK 59063, Morocco, Quarzazate; ZFMK 59064, Morocco, Quarzazate; ZFMK 60598, Morocco, 89 km E Guelmim; ZFMK 60600, Morocco, 52 km S Guelmim; ZFMK 60602, Morocco, Erfoud; ZFMK 60603, Morocco, Erfoud; ZFMK 60605, Mo- rocco, Erfoud; ZFMK 60606, Morocco, Erfoud; ZFMK 60607, Morocco, Erfoud; ZFMK 60608, Morocco; ZFMK 60609, Mo- rocco, Erfoud; ZFMK 60610, Morocco, Erfoud; ZFMK 7459, Morocco, Tinerhir; ZFMK 7462, Morocco, Tinerhir; ZMH R 04527, Morocco, Erfoud; ZMH R04517, Morocco, Rissani near Erfoud; ZSM 186/1983, Algeria, Ain el Hadjadi 24 km south of Ain Sefra; ZSM 31/1981, Morocco, Tinerhir; ZSM 34/1981, Al- geria, Ain Sefra; ZSM 44/1960, Morocco, Zagora; ZSM 58/1978 (1), Morocco, 25 km N Zagora; ZSM 58/1978 (2), Mo- rocco, 25km N Zagora; ZSM 689/1979, Morocco, Ksar es Souk west of Boudenib; ZSM 8/1994, Morocco, Meski. Uromastyx ornata Uromastyx ornata ornata Heyden, 1827 BMNH 97.10.28.199, Egypt, Tor/ Sinai; MHNP 1909.176, Egypt, Mt. Sinai; MHNP 1909.177, Egypt, Mt. Sinai; MHNP 6954, Egypt; MHNP 6970, Egypt; NMW 21217, Egypt, Dahab; NMW 21219:1, Egypt, Sherm Scheikh; NMW 21219:2, Egypt, Bonner zoologische Beitráge 56 (2007) 97 Sherm Scheikh; NMW 21219:3, Egypt, Sherm Scheikh; NMW 21219:4, Egypt, Sherm Scheikh; NMW 21220, Egypt, Tor; ZFMK 65174, Egypt, Wadi Feiran; ZFMK 65175, Egypt, Wa- di Feiran; ZFMK 65607, Egypt; ZFMK 65609, Egypt, ZFMK 8576, Israel, Elath; ZMH R 04525, Egypt, Sinai; ZMH R 04526, Egypt, Sinai; SMF 10403, Holotype of Uromastyx ornata, Sau- di Arabia, Mohila (Al Muwaylih). Uromastyx ornata philbyi Parker, 1938 BMNH 1946.8.11.60, Holotype of Uromastyx philbyi, Saudi Arabia, between Mecca and Shabwa; BMNH 1946.8.11.62, Pa- ratype of Uromastyx philbyi, Saudi Arabia, between Mecca and Shabwa; BMNH 1946.8.11.63, Paratype of Uromastyx philbyi, Saudi Arabia, between Mecca and Shabwa; BMNH 1946.8.11.64. Paratype of Uromastyx philbyi, between Mecca and Shabwa; BMNH 1946.8.11.65, Paratype of Uromastyx phil- byi, Saudi Arabia, between Mecca and Shabwa; BMNH 1946.8.11.66, Paratype of Uromastyx philbyi, Saudi Arabia, bet- ween Mecca and Shabwa; BMNH 1964.296, Saudi Arabia, Bu- rayman 21°40’N 39°10°E; BMNH 1975.518, Saudi Arabia, Baz- zah 22°00’N 39°30’E; BMNH 1975.519, Saudi Arabia, Buray- man 21°39’N 39°13’E; BMNH 1976.1748, Saudi Arabia, Wadi Fatma; BMNH 1979.960, Saudi Arabia, Burayman 21°45’N 39°15’E; BMNH 1985.882, Saudi Arabia, Mecca by pass km 91 /21°14’N 29°48’E; BMNH 1985.884, Saudi Arabia, Mecca by pass km115/21°15,5’N 39°55’E; BMNH 1986.434, Saudi Ara- bia, 21°14’N 39°55’E; BMNH 1986.436, Saudi Arabia, 16 km N of Jeddah; BMNH 1975.518, Saudi Arabia, Bazzah 22°N 39°30’E; BMNH 1980.55, Saudi Arabia, Jabal as Sinfa; MZUF 27884, Yemen, Ju Amlah 26 km NW Sa’dah; MZUF 27885, Ye- men, Ju Amlah 26 km NW Sa’dah; MZUF 27906, Yemen, Ju Amlah 26 km NW Sa’dah; MZUF 28187, Yemen, Ju Amlah 26 km NW Sa’dah; MHNG 2457.33, Saudi Arabia, Jebel Hababa; MHNG 2457.34, Saudi Arabia, Jebel Hababa; MHNG 2457.35, Saudi Arabia, Wadi Sawawin; MHNG 2536.49, Saudi Arabia, Makkah by pass km 56; MHNP 4318, Saudi Arabia, Jeddah; ZFMK 84442, Saudi Arabia, 19°05’N 41°50’E. Uromastyx ocellata Lichtenstein, 1823 BMNH 1914.5.14.13, Sudan, Sinkat; BMNH 1927.8.13.38, Su- dan, Merowe/Dongola; BMNH 1927.8.13.39, Sudan, Merowe/Dongola; BMNH 1937.12.5.117, Somalia, Borama Dis- trict; BMNH 1937.12.5.118, Somalia, Borama District; BMNH 1937.12.5.119, Somalia, Borama District; BMNH 1937.12.5.121, Somalia, Borama District; BMNH 1937.12.5.122, Somalia, Bo- rama District 42°45’N 10°45’E; BMNH 1937.12.5.123, Soma- lia, Borama District 42°45’N 10°45’E; BMNH 1937.12.5.124, Somalia, Borama District 42°55’E 10°55’N; BMNH 1937.12.5.125, Somalia, Borama District 42°55°E 10°55’N; BMNH 1937.12.5.127, Somalia, Borama District 43°E 11°N; BMNH 1937.12.5.128, Somalia, Borama District 43°E 11°N; BMNH 1937.12.5.130, Somalia, Borama District; BMNH 1953.17.63, Sudan, Tehamiyam; BMNH 1953.17.64, Sudan, Te- hamiyam; BMNH 1953.17.65, Sudan, Tehamiyam; BMNH 97.10.28.202, Sudan, Suakim; BMNH 97.10.28.203, Sudan, Sua- kim; BMNH 97.10.28.204, Sudan, Suakim; BMNH 97.10.28.205, Sudan, Suakim; BMNH 97.10.28.206, Sudan, Sua- kim; BMNH 97.10.28.207, Sudan, Suakim; BMNH 97.10.28.208, Sudan, Suakim; BMNH 97.10.28.209, Sudan, Sua- kim; MHNP 1897.348, Egypt; MHNP 1897.349, Egypt; NMW 21215, Oasis Harar, NMW 21216, Sudan, Suakim; ZFMK 20822, Sudan, Suakin; ZFMK 38396, Sudan, 40 km W Suakim direction Sinkat; ZMB 811, Holotype of Uromastyx ocellata, Su- dan, Nubia; ZSM 219/1976, Sudan, Dongola Province. Uromastyx princeps O’Shaughnessy 1880 BMNH 1931.7.20.270, Somalia, 11°5’N 49°0’E; BMNH 1931.7.20.272, Somalia, 8°54’N 48°54’E;, BMNH 1931.7.20.273, Somalia, Buran District 10°13’N 48°46’E; BMNH 1931.7.20.274, Somalia, 10°22’N 49°0’E; BMNH 1931.7.20.275, Somalia, 10°42’N 49°E; BMNH 1946.814.56, Holotype of Uromastyx princeps, Zanzibar (unreliable Locality, WILMS 2001); BMNH 1956.1.3.9, Somalia, Wachderria 45mls E Mait; BMNH 1961.1655, Somalia, Candala/Migiurtina; BMNH 1961.1656, Somalia, Candala/Migiurtina; BMNH 1983.735, Somalia, 5°56’N 48°55’E; MZUF 23691, Somalia, Scusciuban; MZUF 10536, Somalia, District of Alula; MZUF 23673, Somalia, Bur Dagner; MZUF 23674, Somalia, Bur Da- gner; MZUF 23675, Somalia, Meleden; MZUF 23676, Soma- lia, Sukorre; MZUF 23686, Somalia, Carin-Gié Bahaia; MZUF 23690, Somalia, Scusciuban; MZUF 23692, Somalia, Monti Car- car; MZUF 23693, Somalia, Scusciuban; MZUF 23694, Soma- lia, Scusciuban; MZUF 23695, Somalia, Scusciuban; MZUF 23696, Somalia, Scusciuban; MZUF 23782, Somalia, Passo del Didim S of Carin; MZUF 5497, Somalia, Valle di Run; MZUF 5623, Somalia, Valle di Run; MZUF 739, Somalia, Passo del Di- dim S of Carin; MHNP 1966.1071, Somalia, Candala; MHNP 1966.1072, Somalia, Candala; MHNP 5732, Yemen, Aden (un- reliable Locality); MHNP 5831, Somalia, Bender Meraya; MHNP 5832, Somalia, Bender Meraya ; SMF 22931, Somalia, Benden Cassim Migiurtinia; ZFMK 58048, Somalia, Bossasa; ZFMK 58985, Somalia, Bossasa. Uromastyx shobraki Wilms & Schmitz, 2007 BMNH 1938.2.1.47, Country not reliably traceable, southern He- jaz; BMNH 1987.854, Yemen, Tihama Taiz; BMNH 1988.54, Yemen, Mafraq-Al Mokka km 13,5; BMNH 1988.55, Yemen, Mafraq-Al Mokka km 13,6; MZUF 33614, Yemen, Mafraq-Al Mokka km 13,6; MZUF 33615, Yemen, Mafraq-Al Mokka km 13,5; MHNG 2455.100, Paratype of Uromastyx yemenensis sho- braki, Yemen, Mafraq-Al Mukha 13,5km; MHNG 2464.44, Ye- men, Mafraq-Al Mokka km 13,5; MHNG 2496.55, Paratype of Uromastyx yemenensis shobraki, Yemen, between Mocca and Wadi Zabid; MHNG 2496.56, Paratype of Uromastyx vemenen- sis shobraki, Yemen, between Mocca and Wadi Zabid; MHNG 2527.92, Yemen, Wadi Zabid; MHNG 2538.47, Yemen, Wadi Za- bid; MHNG 2542.13, Yemen, Wadi Zabid; MHNG 2542.14, Ye- men, Wadi Zabid; MHNG 2553.56, Paratype of Uromastyx ye- menensis shobraki, Yemen, Mafrag-Al Mokka km 1,5, MTKD 31624, Yemen, North Yemen; MTKD 32847, Yemen, North Ye- men; ZFMK 48680, Paratype of Uromastyx yemenensis sho- braki, Yemen, Mafraq-Al Mokka km 13,6; ZFMK 48681, Ho- lotype of Uromastyx yemenensis shobraki, Yemen, Mafraq-Al Mokka km 13,5; ZFMK 55651, Yemen, North Yemen; ZFMK 55652, Yemen, North Yemen; ZFMK 58047, Yemen, North Ye- men; ZFMK 60687, Yemen, North Yemen; ZFMK 73676, Ye- men, Mafraq - al Mocca; ZFMK 73677, Yemen, Mafraq - al Mocca. 98 Thomas M. WiLms et al.: On the Phylogeny and Taxonomy of the Genus Uromastyx Merrem, 1820 Uromastyx thomasi Parker, 1930 BMNH 1931.7.16.46, Oman, Wadi Hauf; BMNH 1946.8.14.43, Holotype of Uromastyx thomasi, Oman, BU’Juay; BMNH 1954.1.2.98, Oman, Ras Duggum; BMNH 1956.1.16.8, Bahrein (unreliable locality); BMNH 1971.1354, Oman, W of Bai; BMNH 1971.1355, Oman, W of Bai; BMNH 1973.2908, Oman, Masirah; BMNH 1975.1038, Oman, Masirah; BMNH 1977.335, Oman, Jiddat al Harasis 19°32’N 57°12’E; BMNH 1978.1322, Oman, Al Ajaız; BMNH 1978.2249, Oman, near Haq! 20°25’N 58°47’E; BMNH 1980.213, Oman, Thamarit 17°38’N 54°02’E; BMNH 1980.570, Oman, Jiddat al Harasıs 19°32’N 57°13’E; BMNH 1982.1221, Oman, Masirah. Uromastyx yemenensis Wilms & Schmitz, 2007 BMNH 1946.8.11.68, Paralectotype Uromastyx benti, Yemen, Makulla Hadramaut (doubtful record); BMNH 1963.755, Yemen, Wadi Tiban west of Aden; BMNH 95.11.27.6, Yemen, Hills 50 km from Aden; BMNH 95.11.27.7, Yemen, Hills 50 km from Aden; BMNH 99.12.13.106, Yemen, between Mount Manif and Jimil /N. of Lahej; BMNH 99.12.13.51, Yemen, Yabian moun- tains, MTKD 24554, Yemen, Zingibar, Abyan-Gouvernement; MTKD 25441, Yemen, Amran/Aden; MTKD 26951, Yemen, Lawdar, Abyan-Gouvernement, MTKD 26952, Yemen, Lawdar, Abyan-Gouvernement; MTKD 28873, Yemen, Lawdar, Abyan- Gouvernement; MTKD 29475, Paratype of Uromastyx vemenen- sis, Yemen, Zingibar, Abyan-Gouvernement; MTKD 34675, Ye- men, Abyan-Gouvernement; ZFMK 47860, Paratype of Uromas- tyx vemenensis, Yemen, Lawdar, Abyan-Gouvernement; ZFMK 47861, Holotype of Uromastyx yemenensis, Yemen, Lawdar, Abyan-Gouvernement; ZFMK 49036, Paratype of Uromastyx ye- menensis, Yemen, Lawdar, Abyan-Gouvernement. APPENDIX III. Morphological character coding (A) Number of tail worls (0) = 29-36 Whorls (1) = 15-28 Whorls (2) = 9-13 Whorls The taxon hardwickii has 29-36 primary tail whorls. In Leiolepis more than 100 scale rows are always present from the cloacal slit to the tip of the tail. Therefore the character state pres- ent in hardwickii is considered to be plesiomorphic (0). The ma- jority of the taxa (with the exception of princeps and thomasi) have 15—28 primary tail whorls (1). The short tailed taxa prin- ceps amd thomasi are considered to show a character state de- rived from state | (2). (B) Number of gular scales (0) => 29.7 (1) =< 25.9 With the exception of ornata and philbyi, all taxa have an av- erage count of more than 29.7 gular scales. In Leiolepis between 33-45 gular scales are present. Therefore high numbers of gu- lar scales are considered to be the plesiomorphic character state (0). In ornata and philbyi average gulare scale counts are 25.9 and 22.6 respectively (1). (C) Number of scales around midbody (1) = 265-319 With the exception of aegyptia, microlepis, leptieni and occiden- talis all taxa have an average scale count of less than 220 scales around midbody. Leiolepis has between 165-211 scales around midbody. Low scale counts are therefore considered to be the plesiomorphic character state (0). The high scale counts pres- ent in the taxa of the U. aegyptia group are considered to be apo- morphic (1). (D) Number of ventral scales between gular and inguinal fold (0) =< 121 (1)=> 121 With the exception of aegyptia, microlepis, leptieni and hard- wickii all taxa have an average number of ventral scales of less than 121. Leiolepis either has less than 100 ventral scales. Low numbers of ventral scales are therefore considered to be the ple- siomorphic character state (0), while high numbers of ventral scales are considered to by apomorphic (1). (E) Number of scales around Sth tail whorl (0) = 46.2 (1) =<37 In average hardwickii has 46.2 scales around the 5th tail whorl (Range: 40-52). Average values of all remaining taxa are con- siderably lower (between an average of 22.8 and 36.7). In Leiolepis the respective value is 71-125 scales. A high number of scales is therefore considered to be the plesiomorphic char- acter state (0). All remaining taxa have low numbers of scales and are therefore considered to show the apomorphe character state (1). Only in three taxa (microlepis, yemenensis and shobra- ki) single specimens might show values overlapping with the range of hardwickii (43 scales for microlepis, 40 scales for shobraki and yemenensis). (F) Number of scales between subocularia and supralabialia (0) => 3.6 (1) = < 3.2 Most taxa have 3-9 scales between the subocularia and supral- abialia. In Leiolepis 5-7 scales are present. A high scale count is therefore considered to be the plesiomorphic character state (0). Only two taxa (princeps and thomasi) show scale counts as low as two scales. In both taxa average scale counts are lower than in the remaining taxa (princeps 3.2 / thomasi 3.0). This char- acter state is considered to be apomorphic (1). From the remain- ing taxa only geyri and ornata have less than four scales between subocularia and supralabialia (ornata: 3.7; geyri: 3.6). Bonner zoologische Beitráge 56 (2007) 99 (G) Snout-vent-length (SVL) (0)=<28.0 cm (1)=> 35.5 cm Species ofthe genus Leiolepis can reach the following maximum snouth-vent-length: belliana: 16.6 cm; reevesii: 15.1cm; peguen- sis: 13.6 cm; triploida: 14.8 cm and guttata: 18.4 cm (PETERS 1971). Taxa of the genus Uromastyx reach a maximum SVL of: acanthinura: 25.3 cm; nigriventris: 24.0 cm; bei aegyptia: 40.0 cm; asmussi: 26.5 cm; alfredschmidti: 23.0 cm; benti: 19.6 cm; yemenensis: 18.5 cm; shobraki: 20.8 cm; dispar: 23.1 cm; flav- ifasciata: 28.0 cm; maliensis: 23.2 cm; microlepis: 35.5 cm; mac- fadyeni: 11.7 cm; geyri: 19.3 cm; hardwickii: 23.3 cm; lorica- ta: 27.7 cm; leptieni: 37,5 cm (WILMS & BÖHME 2007); ocella- ta: 17.4 cm, ornata: 19.6 cm; philbyi: 19.2 cm; princeps: 15.4 cm and thomasi: 18.2 cm. A maximum SVL of less than 28 cm is considered to be the plesiomorphic character state (0), while a maximum SVL of 35.5 cm or more is considered to be the apo- morphic character state (1). Because of the very limited knowl- edge on the maximum length of Uromastyx occidentalis and the supposed relationship of this species with the large growing taxa of Uromastyx aegyptia we assign apomorphic character state to this taxon. (H) Tail length (0) => 52.6 % of SVL (1) =< 42.9 % of SVL The character „short tailed“ in princeps and thomasi is consid- ered to be apomorphic (1) because this character state is excep- tional within the genus Uromastyx and all species of the outgroup possess long tails (0). (D Tail scalation (0) = last 2-8 whorls consisting of continuous scale rows (1) = last 12-21 whorls consisting of continuous scale rows The members of the Uromastyx ocellata group (ocellata, orna- ta, philbyi, benti, yemenensis, shobraki) as well as macfadyeni show a very unique scalation of the tail, with the last 12-21 whorls consisting of continuous scale rows in which one dorsal scale row corresponds with one row ventrally (1). In all other taxa the character ,,continous scale row on the tail” is restrict- ed only to the last 2-8 whorls. All other whorls in these taxa con- sist of one dorsal scale row with more than one corresponding scale rows ventrally. The latter character state is considered to be plesiomorphic (0), because it can easily be derived from the tail of a Leiolepis with its scalation consisting of small scales. (J) Value of the quotient of head width and head length (0) = 0.861-0.875 (1) = 0.897-0.952 (2) = 0.967-0.997 Within the genus Leiolepis this quotient values between 0.66-0.75. A low average value is considered to be a plesiomor- phy. Therefore princeps (value: 0.875), thomasi (value: 0.874) and yemenensis (value: 0.861) possess the plesiomorphic char- acter state (0), but which is not assigned to pemenensis for the phylogenetic analysis because of the proven relationship of yeme- nensis to the ocellata group within Uromastyx [vemenensis: (1)]. Most of the remaining taxa (exceptions hardwickii, asmussi, lor- icata) have average values for this quotient ranging between 0.897-0.952 which are considered to be the apomorphic char- acter state (1). The relatively broad headed taxa asmussi, lori- cata and hardwickii posses values for this quotient rangeing be- tween 0.967-0.997. This character state is considered to be dif- ferent from (1) and is therefore assigned to (2). (K) Value of the quotient of tail length and maximun with of the tail at the Sth whorl (0) = 4.4-5.2 (1) = 3.4-3.9 (2) = 1.2-2.1 The average value of this quotient within most taxa 1s between 4.4 bis 5.2. In the genus Leiolepis the respective values are be- tween 15 and 20, therefore higth values are considered to be ple- siomorphic (0). For acanthinura, nigriventris, dispar, flavifasciata, maliensis, macfadyeni and philbyi the average value is between 3.4 and 3.9. These character states are considered to be apomorphic (1). For the two short tailed taxa thomasi and princeps the values are 1.2 and 2.1 respectively. These character states are considered to be different to (1) and are therefore assigned to (2). (L) Enlarged tubercular scales on the dorsum (0) = tubercular scales absent (1) = tubercular scales present Leiolepis lacks enlarged tubercular scales on the dorsum. With- in the taxa in question only hardwickii, loricata and asmussi pos- sess enlarged tubercular scales. This character state is consid- ered to be apomorphic (1). (M) Intercalary scales between tail whorls dorsally (0) = intercalary scales present (1) = intercalary scales absent In Leiolepis tail scalation is not arranged in whorls. Therefore the presence of intercalary scales is considered to be the plesio- morphic character state, because their presence is easily derived from a tail witch 1s consisting of small scales (0). The apomor- phic character state shows a reduction and enlargement of sca- le rows on the tail (1). Bonner zoologische Beitráge Band 56 (2007) Heft | Seiten 101-106 Bonn, Marz 2009 Some New and Interesting “Microlepidoptera” from the Collection of the Zoologisches Forschungsmuseum Alexander Koenig (ZFMK), Bonn (Lepidoptera: Tineidae, Epermeniidae, Acrolepiidae, Douglasiidae) Reinhard GAEDIKE Bonn, Germany Abstract. The study of some of the so called “Microlepidoptera” families, the Tineidae, Epermeniidae, Acrolepiidae and Douglasiidae, at the collection of the ZFMK, brought some species of taxonomical and faunistical interest to light. Two species are described as new: Reisserita zouhari sp. n. (Tineidae), and Epermenia (Epermenia) dalianicola sp. n. (Eper- meniidae), and the type-material of the four families, represented in this collection, is listed and commented. Keywords. Tineidae, Epermeniidae, Acrolepiidae, Douglasiidae, new species, type material. The courtesy of my colleague Dr. Dieter Stüning, curator of the Lepidoptera collection at the Zoologisches Forschungs-museum Alexander Koenig, Bonn, enabled me to study the material of the four families Tineidae, Epermeniidae, Acrolepiidae and Douglasiidae, stored at this collection. The paper includes descriptions of two new species, a list of taxa represented by type-specimens and comments on some additional species which are of faunistical interest. TINEIDAE Scalidomia hoenei Petersen, 1991 Holotype d', [China, Jiangsu]. “Lungtan b. Nanking, Prov. Kiangsu, 25. IX. 1933, H. Hóne”; Paratype & same loca- tion and collector, but 22. VIII. 1933. Known from the type locality only. Rhodobates sinensis Petersen, 1987 Holotype 9, [China, Yunnan]. “Li-kiang, Prov. N. Yün- nan, 18. VII. 1934, H. Hone“. The collection contains numerous additional specimens from the type locality and from other locations in China: 12 7, 28 9, Prov. Nord-Yuennan, Li-kiang, between 28. V. and 17. VIII. 1934, leg. H. Höne; 2 9, Shanghai, 10. V. 1933, leg. H. Hone; 1 9, Kuling, 2. V. 1934, leg. H. Hone; 1 9, Prov. Shantung, Laushan bei Tsingtau, 6. VI. 1936, leg. H. Hone; 1 Y Batang (Tibet), Tal des Yangtze, ca. 2800 m, 9. VII. 1936, leg. H. Höne. The female genitalia were illustrated for the first time by Li & XIAO (2006). It is of interest, that the authors col- lected a large series of this species at the type locality, af- ter more than 70 years. Morophaga formosana Robinson, 1986 [ = kobella Robinson, 1986] 1 9, [China, Jiangsu]. “Shanghai, Provinz Kiangsu, 30. Y. 1999, H, Hone Distribution. First record from the Palaearctic region; pre- viously known only from the type locality (Taiwan) and from Fu-chou (China, Fujian). Nemapogon gliriella (Heyden, 1865) [ = cachetiellus Zagulajev, 1963; = cacheticus Zagulajev, 1964; = ibericus Zagulajev, 1968] 1 Y Düsseldorf-Eller, 6. VII. 1941, leg. Friedel. New record for Nordrhein-Westfalen. Distribution. The species is hitherto known from Germany, Austria, Czech Republic, Slovakia, Slovenia, Turkey and the Caucasus region (Georgia, Armenia). Every time, on- ly few specimens were collected. Biology. Larvae feed in fungi Stereum hirsutum (BETTAG 1995; BETTAG & Bastian 1996), Stereum rugosum (C. Vetter, Hamburg, 1996 in litt.). 102 Reinhard GAEDIKE: Some New and Interesting “Microlepidoptera” from the Collection of the ZFMK, Bonn Edosa spinosa (Gaedike, 1984) Holotype 9, [China, Shaanxi]. “Tapaishan im Tsinling, Prov. Súd-Shensi, 1700 m, 16. VI. 1936, H. Hóne”. Distribution. Known from the type locality only. Edosa sinica (Gaedike, 1984) Holotype 9, [China, Jiangsu]. “Prov. Kiangsu, Lungtan bei Nanking, 23. V. 1933, H. Hone”. Paratypes: 4 9, same location, but 20., 22., 28. V. 1933, leg. H. Hone. Distribution. Known from the type locality only. Reisserita zouhari sp. n. (Fig. 1) Holotype 9”, “Aegyptus Mariput El Agami, 9. 5. 1975, Ing. Vlad. Zouhar lgt.;” “Museum A. Koenig Ein[gang] Nr. 90/283 ex Coll. Zouhar;” “Gen. prap. [genitalia slide] Gaedike Nr. 5644;” “Holotypus & Reisserita zouhari sp. n. det. R. Gaedike 2007” Description. Wingspan 12 mm; head, palpi, thorax light ochre, basal area of tegulae darker; forewing light ochre, too, without any pattern, first half of costa darker. Male genitalia (Fig. 4). Uncus bilobate, each lobus with pointed end, between the lobi with deep incision; gnathus arms hooke-shaped, pointed, broadest in the middle; tegu- men and vinculum broad, saccus nearly triangular, apical- ly rounded; valva with long narrow transtilla, corpus di- vided into two processus, one short processus at begin- ning of the costa, the second one longer, hook-shaped, pointed, the view depending on the preparation; aedeagus as long as the whole genitalia, narrow, with two slightly curved cornuti. Female gentalia. Still unknown. Remarks. The new species 1s similar to Reisserita mau- ritanica (Baker, 1885), but the structure of uncus (deep incision, pointed lob1), and valva, the shorter and broad- er gnathus arms and the aedeagus with two cornuti make it distinguishable. The new species ıs named in honour of the collector, Mr. Vladimir Zouhar. Praeacedes atomosella (Walker, 1863) [= seminolella Beutenmüller, 1889; = thecophora Wal- singham, 1908; = despecta Meyrick, 1919; = deluccae Amsel, 1955; = malgassica Gozmány, 1970; = decui Ca- pu?e & Georgescu, 1977] 50,20, Egypt. Kahira-Garden city; Wadi El Natrun, be- tween 1970 and 1974, leg. V. Zouhar. Distribution. Pantropical (ROBINSON & NIELSEN, 1993). Palaearctic region including the Canary Islands; Azores; Malta; Cyprus; Egypt. Biology. The larvae feed from characteristic flattened, elongate oval cases composed of fibers and detritus and lined with silk. They are similar to those of Phaereoeca allutella (Rebel) but in average smaller and more round- ed at proximal end. The cases can be found at house walls. The feeding habit is not known with certainty, but prob- ably the larvae are chitinophagous like those of P. allutel- la (ROBINSON & NIELSEN 1993; O. Karsholt, Copenhagen, pers. observ. 2000). A detailed description of the larva is given by HINTON & BRADLEY (1956) under the name Titaenoses thecophora. Tinea subalbidella Stainton, 1867 [ = arcanella Zagulajev, 1960; = liberiella Zeller, 1879; = excavata Meyrick, 1914; = frugivora Meyrick, 1917; = insectivora Meyrick, 1932] 1 Y, Egypt. Kahira-Garden city, 25. IX. 1974, leg. V. Zouhar. New record for Egypt. Distribution & biology. According to ROBINSON (1989), who in detail described the distribution and the biology, the species is known from Palaearctic (Pakistan), Orien- tal (Thailand, India, Sri Lanka) and Afrotropical (Sierra Leone, Nigeria) regions. Specimens were reared from monkey skins, the pelt of a wild cat, horns and dead Or- thoptera. In Thailand, larvae shared nests with an infes- tation of Monopis longella. They behaved in a similar way to those of M. longella, tunnelling in the feathers and pu- pating just below the nest surface in a cylindrical cocoon of thin silk with adhering fragments of feathers. Tinea omichlopis Meyrick, 1928 [= nonimella Zagulajev, 1955] 2 9, [China, Shaanxi]. “Tapaishan im Tsinling, Süd-Shen- si, 28., 31. V. 1935, H. Höne”. Distribution. Hitherto known throughout the Palaearctic region from Italy, Balkan peninsula, Central Europe to Caucasus region, Middle to Far East. First record for Chi- na. Niditinea sinensis Petersen & Gaedike, 1993 Holotype 9, [China, Jiangsu]. “Prov. Kiangsu: Shanghai, 19. IX, 1943, H, Hone. Paratypes. 3 &, same location, but 19. VIII, 9., 17. IX. 1942, leg. H. Höne. Distribution. Known from the type locality only. Bonner zoologische Beitráge 56 (2007) 103 Crypsithyris japonica Petersen & Gaedike, 1993 Holotype 9, Japan, Unzen, 15. VU. 1937, leg. H. Höne. Distribution. Known from the type locality only. Crypsithyris hoenei Petersen & Gaedike, 1993 Holotype Y, [China, Yunnan]. ,,Li-kiang, Prov. Nord- Yuennan, 22. VI. 1934, H. Hone”. Paratype: 1 Y, same location, but 4. VII. 1934, leg. H. Hone. One additional gd” from the type location. Distribution. Known from the type locality only. Monopis megalodelta Meyrick, 1908 (Fig. 2) 1 9, Türkei, Bedirge, 4. V. 1968, leg. Mittendorf. Distribution. Previously known only from the Afrotropi- cal region, first record from the Palaearctics. The species is superficially similar to M. monachella (Húbner, 1796), but the shape of the white area on forewing is distinctly different, other differences are seen in the structure of the genitalia (Fig. 5). Monopis trapezoides Petersen & Gaedike, 1993 Holotype 9', [China, Zhejiang]. ,,Prov. Chekiang, West Tien-mu-shan, 15. IX. 1932, H. Hone”. Paratypes: 2 9, same location and collector, but 3., 5. IX. 1932. Distribution. Known from the type locality only. Gerontha hoenei Petersen, 1987 Holotype 9, [China, Yunnan]. ,,Li-kiang, Prov. N-Yuen- nan, 31. VII. 1935, H. Hóne”. Paratypes: 2 9', same lo- cality, but 26. VII., 2. VIII. 1935, leg. H. Höne; 1 Y Chi- na, West Tien-mu-shan, Prov. Chekiang, 2. VIII. 1932, leg. H. Hone. Additionally 3 specimens from the type locality. Distribution. Known from the type locality only. Tinissa insularia Robinson, 1976 Paratypes. 1 Y, 1 9, NO-Sumatra, Dolok Merangier, 180 m, September 1969, 9.1.-2.11.1970, leg. Diehl; 19 N- Sumatra, Ketambe, 40 km NW of Kutatjane, 300-500 m, 10.-18.V1.1972, leg. Roesler & Küppers. Distribution. Known from the type localities: Java, Malaya, Borneo, Celebes, Moluccas, Philippines, New Guinea, Solomon Islands, Sumatra. EPERMENIIDAE Epermenia (Epermenia) dalianicola sp. n. (Fig. 3) Holotype Y, “China. Luda, Dalian, 10. 5. 1959, Ing. VLAD. ZOUHAR lgt.;” “Museum A. Koenig Eing.[ang] Nr. 90/283 ex Coll. Zouhar;” “Gen. präp. [genitalia slide] Gaedike Nr. 5832;” “Holotypus Y Epermenia (Eperme- nia) dalianicola sp. n. det. R. Gaedike 2008;” Paratype. Without abdomen, same location, but 29. 4. 1959, “Museum A. Koenig Eing.[ang] Nr. 90/283 ex Coll. Zouhar;”“Paratypus Epermenia (Epermenia) dalianicola sp. n. det. R. Gaedike 2008;” Description. Wingspan 11 mm; head, palpi, thorax dark grey, tips of the scales whitish; scales on forewing whitish at base, distal half dark grey; on dorsum at 1/3 and 1/2 a tuft of raised black scales; fringe with two black lines in the middle and apically; on the cell, before and behind 1/2 and at 3/4, three light brown dots present, the apical dot is the largest; obliquely above the two tufts, indications of additional very small brown dots are present; hindwing light grey. The holotype is somewhat lighter, because the pattern is rubbed. Male genitalia (Fig. 6). Uncus long, narrow, slightly curved, with pointed tip; tegumen broad, apical edge and the middle stronger sclerotized than the other part; valva with small transtilla, ampulla broad, curved, with a blunt tip, basally with a strong sclerotized edge; cucullus broad, rounded; sacculus apically with a very small pointed tip below the sclerotized edge of ampulla; aedeagus shorter than valva, with a hooked cornutus. Female genitalia. Still unknown. Remarks. The new species is superficially similar to E. ochreomaculella asiatica Gaedike, 1979, but clearly dis- tinguishable by the structure of the genitalia, especially the shape of the ampulla. The species is named after the location of the types: Dalian. Epermenia (Cataplectica) sinica Gaedike, 1996 Holotype o’, [China, Yunnan]. ,,Li-Kiang, Prov. Nord-Yu- ennan, 19. VII. 1934, H. Hone“. Distribution. Known from the type locality only. Ochromolopis kaszabi Gaedike, 1973 1 9, [China, Hunan]. “Prov. Hunan, Hoengshan, 15. IV. 1933, H. Hóne”. New record for China. Distribution. Hitherto known from eastern parts of the Palaearctic region (Mongolia, Siberia, Russian Far East). 104 Reinhard GAEDIKE: Some New and Interesting “Microlepidoptera” from the Collection of the ZFMK, Bonn Figs 1-3. 1. Reisserita zouhari sp. n.; 2. Monopis megalodel- ta Meyrick, 1908; 3. Epermenia (Epermenia) dalianicola sp. n. Figs 4-6. 4. Reisserita zouhari sp. n., male genitalia (aedea- gus separated); 5. Monopis megalodelta Meyrick, 1908, male genitalia (left valva removed); 6. Epermenia (Epermenia) da- lianicola sp. n., male genitalia (left valva removed, aedeagus se- parated). Bonner zoologische Beiträge 56 (2007) 105 ACROLEPIIDAE Digitivalva (Digitivalva) asiatica Gaedike, 1971 Holotype 9, [China, Shaanxi]. “Tapaishan im Tsinling, Süd-Shensi, 1. VII. 1935, H. Hone”; 3 9, China, Li-kiang, Prov. Nord-Yuennan, 18. VI., 10. VII., 13. VII. 1934, leg. H. Höne First record besides the holotype. Distribution. China: Shaanxi, Yunnan. Digitivalva (Digitivalva) hoenei Gaedike, 1971 Holotype 9, [China, Yunnan]. ,,Li-kiang, Prov. Nord- Yuennan, 10. VII. 1934, H. Hone”. Paratype: 1 9, same location, but 8. VII. 1934, leg. H. Hóne; one additional specimen from the type locality. Distribution. Known from the type locality only. Acrolepiopsis sinense Gaedike, 1971 Holotype 9, [China, Zhejiang]. „Wenchow (Chekiang), 12. IV. 1934, H. Hóne”. Paratype. 1 9, same data. Distribution. Known from the type locality only. Acrolepiopsis deltoides Gaedike, 1971 Holotype 9, [China, Zhejiang]. ,, West Tien-mu-shan, Pro- vinz Chekiang, 4. IX. 1932, H. Hóne. Distribution. Known from the type locality only. DOUGLASIIDAE Tinagma anchusellum (Benander, 1936) 1 9, Jordanien, Amman, 25. III. 1968, leg. S. u. J. Klap- perich; 1 9, O-Jordanien, Zerkatal bei Romana, 15. III. 1966, leg. S. u. J. Klapperich: New record for the coun- try. Distribution. Known from North Europe (Sweden, Den- mark, Estonia, Latvia) through Central Europe and Bal- can peninsula (Greece) to Cyprus, Bulgaria, Ukraine, fur- ther from Turkey and the Caucasus region (Armenia, Azer- baidshan). Biology. Larvae are mining on Anchusa officinalis. Tinagma balteolellum (Fischer von Roeslerstamm, 1841) [= borkhauseniella Herrich-Schäffer, 1855] 1 9, Jordanien, Amman, 4. IV. 1968, leg. S. u. J. Klap- perich. New record for Jordania. Distribution. Known from nearly whole Europe, except northern part, and outside of Europe from Morocco. Biology. Larvae are mining in stems of Echium vulgare and E. biebersteini. Tinagma klimeschi Gaedike, 1987 32 O, Aegyptus, Alexandria, 25., 26. II. 1973; 2. III. 1974, leg. V. Zouhar; 1 9, Israel, Latroun bei Jerusalem, 25. II. 1968, leg. Bles- zynski. New record for the two countries. Distribution. Hitherto known only from Rhodos (type-lo- cality) and from Cyprus. Biology. Larvae are mining on Echium diffusum. Tinagma minutissimum (Staudinger, 1880) 1 7, Volgograd, 18.-24. V. 1967, leg. V. Zouhar Distribution. Previously known only from the type local- ity (Turkey: Amasia) and from Crimea. REFERENCES BETTAG, E. 1995. Zur Biologie und Verbreitung seltener Tinei- dae und der Eule Hypenodes turfosalis Wocke, 1850 in Rhein- hessen-Pfalz und an der Nahe (Lep., Tineidae et Noctuidae). Melanargia 7(4): 89-96. BETTAG, E. & BASTIAN, K. 1996. Verzeichnis der Klein-Schmet- terlinge (Insecta: Lepidoptera) von Rheinhessen-Pfalz. Teil VII: Tineidae (Echte Motten). Mitteilungen der Pollichia 83: 177-202. GAEDIKE, R. 1971. Die Acrolepiidae der China-Ausbeute H. Hö- ne. Beiträge zur Entomologie 21 (3/6): 273-277, 13 figs. GAEDIKE, R. 1996. Die Acrolepiidae und Epermeniidae der Ne- pal-Expeditionen der Zoologischen Staatssammlung München sowie eine neue Epermenia aus China (Insecta: Lepidoptera). Spixiana 19(2): 221-227, 18 figs. Hinton, H. E. & BRADLEY, J. D. 1956. Observations on species of Lepidoptera infesting stored products. XVI: Two new gen- era of clothes moths (Tineidae). The Entomologist 89(1113): 42-47, 4 figs. Li, H. & Xiao, Y. 2006. A study of the genus Rhodobates Ragonot (Lepidoptera: Tineidae) from China. Proceedings of the Entomological Society of Washington 108(2): 418-428, 30 figs. PETERSEN, G. 1987. A new Gerontha (Tineidae) from China. Ti- nea, Supplement 12: 152-154, 7 figs. PETERSEN, G. & GAEDIKE, R. 1993. Tineiden aus China und Ja- pan aus der Hóne-Sammlung des Museums Koenig (Lepido- ptera: Tineidae). Bonner zoologische Beitráge 44(3/4): 241— 250 14 figs. 106 Reinhard GAEDIKE: Some New and Interesting “Microlepidoptera” from the Collection of the ZFMK, Bonn ROBINSON, G. S. 1976. A taxonomic revision of the Tinissinae Author’s adress: Dr. Reinhard GAEDIKE, Florusstraße 5, of the World (Lepidoptera: Tineidae). Bulletin of the British 53225 Bonn, Germany; E-Mail: tinagma@msn.com. Museum (Natural History), Entomology 32(7): 253-300, 16 plates, 10 figs. ROBINSON, G. S. 1989. Tinea subalbidella Stainton (Lepidopte- . ra: Tineidae) — Nomenclature and Biology. Entomologist’s Ga- Received: 15.03.2008 zette 40: 207-210, 1 fig. Revised and accepted: 28.04.2008 ROBINSON, G. S. & NIELSEN, E. S. 1993: Tineid Genera of | Corresponding editor: D. Stüning Australia (Lepidoptera). Monographs on Australian Lepidop- tera. 2: XV + 344, 734 figs. Seiten 107-129 Bonn, Márz 2009 Bonner zoologische Beitráge Band 56 (2007) Heft 1/2 Islands Between the Realms: A Revised Checklist of the Herpetofauna of the Talaud Archipelago, Indonesia, with a Discussion About its Biogeographic Affinities André Kocu!)"), Evy AriDa2)3), Awal RıyanTo2) & Wolfgang Bónme!) DZoologisches Forschungsmuseum A. Koenig, Bonn, Germany; Museum Zoologicum Bogoriense, Cibi- nong, Indonesia, Current address: Zoologisches Forschungsmuseum A. Koenig, Bonn, Germany; “Corresponding author: andrepascalkoch@web.de Abstract. Since the last taxonomic account was published 80 years ago, we provide the first updated annotated check- list for the herpetofauna of the remote Talaud Archipelago, which lies between the biogeographic realms of Sulawesi, the Philippines, and the Moluccas. We report on a small collection from Pulau (= island) Salibabu, the second largest is- land within the Talaud Archipelago, which, until recently, was one of the least-known areas of the Sulawesi region. In total, 37 specimens representing three anuran species, 11 lizard species (four geckos, four skinks, two agamids, and one monitor lizard species), and two snake species were collected. Of these, six species, viz. one microhylid frog (Callulops cf. dubius), three lizard species (Nactus cf. pelagicus, Gehyra mutilata, and Eutropis cf. rudis), and two snake species (Typhlops sp. and Boiga irregularis), are recorded here for the first time for the Talaud Islands, while Hydrosaurus sp., Cyrtodactylus cf. jellesmae, Eutropis multicarinata and Emoia atrocostata have not been previously known from Sali- babu Island. Novel records for Talaud are completed by Hemidactylus frenatus and Eutropis multifasciata from the MCZ online database. A historical record of Gekko vitattus may be incorrect but seems possible given the biogeographic dis- tribution patterns of Australopapuan amphibian and reptile species discussed in this paper. Historical reports of Candoia reaching North Sulawesi appear dubious. Combined with previous records, the herpetofauna of Talaud currently com- prises 27 species of amphibians and reptiles that include three different species of frogs, 17 lizards (five geckos, eight skinks, three agamids, and one varanid), five snake species as well as one sea turtle and one crocodile. Two historical re- cords of Gekko vittatus and Bronchocela jubata, respectively, require verification. Finally, the zoogeographic affinities of the Talaud Islands and their implications for the course of past dispersal routes between the Philippines, North Sulawesi and the northern Moluccas are discussed in the light of herpetofaunistic distri- bution patterns compared with those shown by other groups of organisms. Keywords. Wallacea, Sulawesi region, amphibians, reptiles, biogeography, endemism. Abstrak (Bahasa Indonesia). Kami memperbaharui daftar jenis-jenis herpetofauna yang ditemukan di Kepulauan Ta- laud, yang secara biogeografi diapit oleh Sulawesi, Kepulauan Filipina, dan Kepulauan Maluku, setelah daftar yang per- tama dipublikasikan 80 tahun yang lalu. Laporan kami mencakup koleksi amfibi dan reptil yang berasal dari Pulau Sal- ibabu, yang merupakan pulau terbesar kedua di Kepulauan Talaud yang sampai saat ini kurang dikenal dibandingkan den- gan daerah lain di sekitar Sulawesi. Di antara koleksi kami yang berjumlah total 37 spesimen, termasuk di dalamnya adalah 3 jenis amfibi dari kelompok Anura, 4 jenis Cicak, 4 jenis Kadal, 2 jenis Londok, dan | jenis Biawak. Enam (6) jenis di antara koleksi kami ini baru pertama kalinya dilaporkan dari Kepulauan Talaud, yaitu 1 jenis katak Mikrohila (Callulops cf. dubius), 2 jenis Cicak (Nactus cf. pelagicus dan Gehyra mutilate), | jenis Kadal (Eutropis cf. rudis), dan 2 jenis Ular (Typhlops sp. dan Boiga irregularis). Catatan baru dari Pulau Salibabu adalah Hydrosaurus sp., Cyrtodacty- lus cf. jellesmae, Eutropis multicarinata and Emoia atrocostata. Kami juga menambahkan Hemidactylus frenatus dan Eutropis multifasciata ke dalam daftar kami, karena kedua jenis ini dilaporkan telah ditemukan di Kepulauan Talaud dalam daftar online MCZ. Laporan mengenai Gekko vitattus yang berasal dari Kepulauan Talaud kemungkinan kurang tepat, namun dapat dibenarkan bila didasarkan pada pola persebaran biografi amfibi dan reptil di daerah Australopapua yang kami ulas dalam tulisan ini. Selanjutnya, laporan tentang ditemukannya Candoia di bagian utara Sulawesi kurang dapat dibenarkan. Sampai saat ini, secara keseluruhan tercatat 27 jenis amfibi dan reptil dari Kepulauan Talaud, yang men- cakup 3 jenis Katak, 5 jenis Cicak, 8 jenis Kadal, 3 jenis Londok, satu jenis Biawak, dan 5 jenis Ular, di samping 1 je- nis Penyu dan 1 jenis Buaya. Seluruh catatan mengenai hewan-hewan ini kami susun dalam sebuah daftar beserta keteran- gan tambahannya. Kami juga memaparkan keterlibatan Kepulauan Talaud dalam jalur persebaran amfibi dan reptil yang telah terjadi di an- tara Pulau Sulawesi, Kepulauan Filipina, dan Kepulauan Maluku dan membandingkannya dengan jalur persebaran or- ganisme lain di daerah ini. Kata kunci. Wallacea, Sulawesi, amfibia, reptilia, biogeografi, jenis endemik. 108 André Koch et al.: A Revised Checklist of the Herpetofauna of the Talaud Archipelago, Indonesia 1. INTRODUCTION The Talaud Archipelago represents the northern most area within the Wallacea transition zone between the Oriental and Australian faunal regions. The remote island group lies in the northern Molucca Sea between North Sulawesi, Mindanao in the southern Philippines, and Halmahera in the northern Moluccas. The Talaud Archipelago compris- es three main islands: Karakelong, Salibabu and Kaburu- ang (Fig. 1). Geologically, the Molucca Sea Plate with its inverted U-shape dips east under Halmahera and west un- der the Sangihe Arc (MCCAFFREY et al. 1980). The Talaud Islands are situated on the Sangihe forearc separated by a deep ocean trench from the largely submarine Sangihe Arc with its few emergent volcanic islands like Sangihe and Siau (HaLL 2002). The Talaud Island group consists of a sedimentary sequence of Middle Miocene to Pleis- tocene age and includes probable mid-Miocene volcanic rocks and volcaniclustic turbidites (MOORE et al. 1981). Pleistocene coral limestone, the youngest rocks on Talaud, are exposed mostly along the coast lines (Fig. 2), but al- Talaud Islands Sangihe * so at 400-500 m elevation on Kaburuang, showing the sig- nificant amount of recent uplift (Moore et al. 1981). The highest elevation of only 650 m above sea level is found on Karakelong Island. The Talaud Islands together with Sulawesi and other is- lands of present day central Indonesia belong to a biogeo- graphic transition region called Wallacea (see summaries by Mayr [1944] and Sımpson [1977]). Australopapuan and Asian animals meet at the respective outer limits of their distribution. Consequently, in the light of biogeography, the Talaud Islands form the northernmost submerged area of this region. Traditionally, the Talaud Archipelago has been dealt with scientifically with Sulawesi to which it is indirectly connected via the Sangihe Island arch. Howev- er, the direct over-water distance to North Sulawesi of about 300 km is significantly more distant than Halma- hera in the Moluccas (ca. 210 km) or Mindanao in the Philippines (ca. 170 km). ear , =? Karakelong Fig. 1. Map of the Talaud Islands located between Sulawesi, Halmahera (Moluccas) and Mindanao (Philippines). Bonner zoologische Beitráge 56 (2007) 109 Fig. 2. Coast line with coral limestone deposits on Salibabu Island. Habitat of Varanus sp. and Emoia a. atrocostata. Photo by André Koch. The geographically isolated position between three bio- geographic subregions in the centre of the Indo-Australian Archipelago render the Talaud Islands an interesting field of research for biogeographers. Until recently, however, the Talaud Archipelago (together with Sangihe) has been one of the least-known areas of the Sulawesi region as de- fined by VANE-WRIGHT (1991). Hence, due to their minor geographic size together with their remote location, the Talaud Island group has been paid only little attention by herpetologists in the past and scientific literature about the herpetofauna of this small archipelago is scarce. VAN KAMPEN (1907; 1923) and DE Roow (1915; 1917) provid- ed the first herpetological data about the Talaud Islands based on material collected by M. WEBER during the Si- boga Expedition of 1899. DE Roow (1915; 1917) listed five different lizards species (Calotes jubatus, Varanus in- dicus, Lygosoma cyanurum, Lygosoma rufescens, and Mabuia multicarinata) and one snake species (Candoia carinata). Later, DE JONG (1928) published the results of another small collection by the Dutch botanist H. J. Lam who visited Karakelong and Salibabu as well as Miangas, a small islet south of Mindanao, in 1926 (Lam 1926; 1942). His Talaud collection comprised ten different species representing eight lizards and two snakes. Vouch- er specimens by M. WEBER and H. J. Lam from the Ta- laud Islands are mostly deposited in the Zoological Mu- seum of the University of Amsterdam. A few of their spec- imens are stored in the herpetological collection of the Mu- seum Zoologicum Bogoriense (MZB) on Java. Further material (e. g., collected by the Snellius Expedition in 1930) are located in the collections of the Natural Histo- ry Museum Naturalis in Leiden (RMNH), the Netherlands. The herpetology department of the Museum of Compar- ative Zoology (MCZ), Harvard University, houses a small number of vouchers (MCZ R-45768-775; MCZ A-24288) from Karakelong Island collected by the first FAIRCHILD Tropical Botanic Garden Expedition led by the late David FAIRCHILD (1869-1954). This expedition made a short stop-over at the Talaud Islands from 12 to 13 June 1940. The collection merely contains nine specimens re- presenting five lizard species (Bronchocela cristatella, Emoia caeruleocauda, Hemidactylus frenatus, Lipinia noctua, and Eutropis m. multifasciata) as well as one snake (Candoia paulsoni tasmai) and one hylid frog species (Litoria infrafrenata). To our knowledge, only McDow- ELL (1979), BROWN (1991) and SMITH et al. (2001) re- ferred, in part, to the FAIRCHILD collection. Therefore, these voucher specimens are included in our report. 110 André Koch et al.: A Revised Checklist of the Herpetofauna of the Talaud Archipelago, Indonesia This checklist of the Talaud Islands summarizes the pres- ent-day knowledge of the herpetofauna and discusses the biogeographic relations and implications of these data. It 1s the first updated account of the amphibians and reptiles of this remote oceanic archipelago since 80 years. 2. MATERIAL AND FIELD WORK Field studies on the east coast of Salibabu Island (ca. 95 km?) were conducted by AK and EA from 13 to 21 July 2005. Collections and observations were made in the vicinity of Lirung, a settlement with a ferry landing stage. Field numbers are AK039 to AK075. The surroundings near the coastline are dominated by agricultural vegeta- tion, especially coconut (Cocos nucifera) and nutmeg (Myristica fragrans) plantations (Fig. 3). The island’s hilly interior was not intensively surveyed. In the low central hills, however, primary habitats or at least areas of minor disturbance still exist (Fig. 4). As recently recognized by RILEY (2002), and unlike the neighboring island Salibabu, large areas of Karakelong Island (ca. 970 km?) are still Fig. 3. Coconut and nutmeg plantations dominate the cultivat- ed coastal landscape of Salibabu Island. Red arrow indicates Va- ranus sp. basking on a palm trunk. Photo by André Koch. Fig. 4. Undisturbed riverine habitat upcountry of Salibabu Is- land. Habitat of Limnonectes sp. and an unidentified snake spe- cies. Photo by André Koch. forested. Surveys were done throughout all hours of day and early night. Specimens were mainly collected manu- ally and with the assistance of a local villager. Voucher specimens were photographed prior to and after euthaniza- tion and preserved in 70 % ethanol. They are deposited in the Zoological Museum in Bogor (MZB). Tissue sam- ples of monitor lizards were taken for molecular investi- gations and preserved in 95 % ethanol. Photographic vouchers are deposited in the private photo collections of AK and EA. Species determination and distribution records follow BOETTGER (1895a, b; 1903), BOULENGER (1897), BARBOUR (1912), DE Roon (1913; 1915; 1917), VAN KAMPEN (1923), DE JONG (1928), MERTENS (1929), PARKER (1934), TAN- NER (1950), BROWN & ALCALA (1970), BROWN & ALCALA (1980), BROWN (1991), MONK et al. (1997), HALLERMANN (2005), DE LANG & VOGEL (2005), and ZIEGLER et al. (2007). All species known from the Talaud Islands are list- ed in Table 1. Abbreviations used are: SVL — snout vent length; TL — tail length; TiL — tibia length. Measurements are given in mm. Museum acronyms are as follows: CAS — California Academy of Science, San Francisco, USA (CAS-SU denotes the Stanford University collection [SU], which is also housed in the CAS); MCZ — Museum of Comparative Zoology, Cambridge, USA; MTD — Muse- um of Zoology (Museum fúr Tierkunde), Dresden, Ger- many; MZB — Museum Zoologicum Bogoriense, Cibi- nong, Indonesia; RMNH — Nationaal Natuurhistorisch Museum Naturalis, Leiden, Netherlands; ZFMK — Zoo- logisches Forschungsmuseum Alexander Koenig, Bonn, Germany; ZMA — Zoological Museum, University of Am- sterdam, Netherlands. Information about ZMA specimens was taken from the on- line database at http://1p30.eti.uva.nl/zmawebsite/search- specimens.php. Bonner zoologische Beitráge 56 (2007) 111 The website http://collections.oeb.harvard.edu/Herp/Rept- Search.htm provides information on Talaud specimens in the MCZ collections. 3. RESULTS ANNOTATED CHECKLIST OF THE AMPHIBIANS AND REP- TILES OF THE TALAUD ISLANDS AMPHIBIA Hylidae Litoria infrafrenata (Günther, 1867) (Fig. 5) Material examined: MZB Amph. 11480, 11490, 11492 (AK039-040, AK072) Additional material: MCZ A-24288, Karakelong, coll. FAIRCHILD Garden Expedition 1940; ZMA 8570 (3 spec.), Lirung, Salibabu, coll. M. WEBER 1899; ZMA 8576 (1 spec.), Beo, Karakelong, coll. M. WEBER 1899. Fig. 5. Litoria infrafrenata from Salibabu Island. Photo by An- dre Koch. Distribution: VAN KAMPEN (1907) was the first to record this species for the Talaud Islands (at that time as Hyla dolichopsis Cope, 1867, a junior synonym of L. infrafre- nata). According to BARBOUR (1912) it “(...) is the most widespread Hyla of this whole region [i.e. the Indo-Aus- tralian Archipelago]”. He listed four female specimens from Lirung and Beo, on Salibabu and Karakelong, re- spectively. Later however, VAN KAMPEN (1923) only states “Talaut Islands” without specifying the exact locality. Morphology: VAN KAMPEN (1907) mentioned that disk size varies in this species. Thus, in specimens from Lirung for instance, the disks are smaller than the tympanum (in one specimen three vs. four mm). The same relation ap- plies to our voucher specimens where the disks are also smaller than the eye diameter. SVL of largest specimen (MZB Amph. 11492) 115.5 mm; TiL: 62 mm. Males ex- hibit black rugosities on the inner side of the first finger. This is the case in our specimen MZB Amph. 11480. Microhylidae Callulops cf. dubius (Fig. 6) Material examined: MZB Amph. 11443 (AK047), MZB Amph. 11469 (AK054), MZB Amph. 11470 (AK053), MZB Amph. 11496 (AK074), MZB Amph. 11500 (AK052). Distribution: Our findings on Salibabu Island expand the known range of the genus by approximately two hundred kilometers to the north and represent a new family record for the Talaud Islands. Ecology: Subadult and adult voucher specimens were ei- ther found in small holes near the base of trees (Fig. 6), or after rainfall near a small stream in the early evening hours (7.30-8.30 pm). Morphology «€ Taxonomy: Characteristically short- snouted, small frogs with short limbs: SVL 23.0-39.6 mm, TiL 9.5-15.5 mm; dorsally uniform dark brown to gray- Fig. 6. Callulops cf. dubius from Salibabu Island. Photo by An- dré Koch. 112 André Koch et al.: A Revised Checklist of the Herpetofauna of the Talaud Archipelago, Indonesia ish black, ventral side either uniform light grey or dark grey with more or less light mottling, toes with weak light transverse bands. The Talaud population may represent a member of the Callulops (Phrynomantis) robustus group (R. GUNTHER, pers. comm.). The genus Callulops inhabits the Moluc- cas, New Guinea and surrounding island groups with 17 recognized species (FROST 2007), two of which (C. boettgeri and C. dubius) are found on Halmahera. Accord- ing to MÉHELY (1901) and PARKER (1934) C. boettgeri is characterized by large triangular discs at fingers and toes, while those of the fingers are larger than those of the toes. Having the finger tips pointed and those of the toes slight- ly enlarged, fingers and toes without webbings, no max- illary teeth, a smooth skin above and beneath, the nostril much closer to the tip of the snout, tympanum feebly dis- tinct and about half the diameter of the eye, presence of a supratympanic fold (but absent in MZB Amph. 11443 and 11500, possibly due to preservation), our voucher specimens resemble more the second Callulops species known from Halmahera, C. dubius (BOETTGER 1895b). However, in contrast to the latter species, the first finger is longer than the second in the Talaud specimens. It can- not be ruled out that the Talaud specimens represent two distinct species due to their differences in ventral color pat- tern and the presence or absence of a supratympanic fold. Ranidae Limnonectes cf. modestus (Boulenger, 1882) Material examined: MZB Amph. 11478 (AK073), 11493 (AK068), 11494 (AK050), MZB Amph. 11497 (AK051). Additional material: Probably ZMA 8870 (5 spec.), “Taloek, island North of Halmaheira” (= Talaud?!), coll. M. WEBER 1900 Distribution: Based on material collected by the Siboga Expedition of Max WEBER (six specimens according to van Kampen [1907], but only five according to the ZMA online catalogue; see “additional material”), VAN KAMPEN (1907) was the first to record L. modestus from the Ta- laud Islands without indicating the exact locality. This species 1s also known from North Sulawesi, the type lo- cality, and some islands in the Moluccas. The Philippine populations have been removed from the synonymy of L. modestus (INGER 1954). Ecology: While two adult voucher specimens were col- lected near a small stream in the vicinity of the village Lirung, the third (MZB Amph. 11493) was found as road kill next to a coconut plantation. Morphology & Taxonomy: All three adult specimens closely resemble in morphology and coloration. SVL 58.6-63.6 mm, TiL 32.7-33 mm, head as broad as long or broader than long; vomerine and maxially teeth pres- ent; nostril nearer to tip of snout than to eye; tympanum distinct, 3/4 the diameter of the eye; strong supratympan- ic fold; finger tips pointed, those of the toes with very small discs; first finger slightly longer than second; fin- gers without webbings, toes nearly fully webbed; dermal fringe along the outer side of fifth toe present; inner pal- mar tubercle swollen, outer missing; inner metatarsal tu- bercle present, outer missing; skin smooth without ridges or tubercles. Coloration dorsally olive-green to brownish with little dark spots; tympanum partly blackish; supratympanic fold above tympanum black; upper lips with dark vertical bars; limbs without dark cross bars; pos- terior part of thighs blackish with orange marbling; ven- tral side either unicolored whitish cream under head and chest, limbs and posterior half of venter orange, particu- larly towards vent (MZB Amph. 11494), or underside whitish with grey mottling under head and many grey spots and blotches on chest and venter. Probably three fe- males were collected without vocal sacs. Accordingly, the specimens lack bony processes in the lower jaw as typi- cal for males of L. modestus. A fourth subadult specimen of Limnonectes was collected in a narrow gorge with a small riverine (Fig. 4). SVL 27.3 mm, TiL 13.8 mm, head slightly longer than broad. Morphologically, this specimen resembles the three adult specimens, except for the indis- tinct supratympanic fold. However, it shows differences in color pattern. Thus, the dorsal side is brown with dis- tinct darker cross bars and blotches on the limbs; a dark cross bar between the eyes is present, too; the distinct mar- bling on the posterior part of the femurs is missing. MENZIES (1987) suggested that populations of Sulawesi and Ceram might represent different taxa. Further inves- tigations, morphologically and genetically, are needed to clarify the taxonomic identity of the Talaud populations. SAURIA Agamidae Bronchocela cristatella (Kuhl, 1820) (Fig. 7) Material examined: MZB Lac. 5080 (AK057). Additional material: ZMA 18832 (4 spec.), Gunung Doeata (= Mt. Duata), Karakelong, coll. H. J. Lam, 1926; ZMA 18833, Lirung, Salibabu, coll. H. J. Lam, 1926; MCZ R-45770, -771, Karakelong, coll. FAIRCHILD Garden Expedition, 1940 Bonner zoologische Beiträge 56 (2007) 113 Fig. 7. Bronchocela cristatella from Salibabu Island. Photo by André Koch. Distribution: DE JONG (1928) reported that H. J. Lam col- lected several specimens of B. cristatella on Karakelong and Salibabu Island. Two further specimens were collect- ed by the FAIRCHILD Expedition in 1940. As neither of these old voucher specimens has been included in a re- cent review of the genus Bronchocela (HALLERMANN 2005), their taxonomic identification requires verification. Nevertheless, our investigations confirm that B. cristatel- la also inhabits the Talaud Archipelago next to Sulawesi, the Philippines and the Moluccas. Ecology: The voucher specimen (MZB Lac. 5080) was found on a tree trunk near three small fish ponds where Hydrosaurus also occurred. Another subadult specimen of Bronchocela was discovered sitting on a thin branch above a small stream near Lirung village. Morphology: SVL 89 mm, TL 298 mm (TL/SVL: 3.34); eight supralabials; nine infralabials; 55 scales around mid- body, the first five to six upper scale rows of the lateral side keeled and pointing upwards; ventral scales larger than dorso-laterals; 35 strongly keeled scales under fourth toe; six to seven scales along canthus rostralis between nostril and anterior border of orbit; diameter of tympanum more than half diameter of orbit (0.83). Bronchocela jubata Duméril & Bibron, 1837 Material examined: none Additional material: ZMA 18866, Lirung, Salibabu, coll. M. WEBER 1899; ZMA 18869, Beo, Karakelang (= Karakelong), coll. M. WEBER 1899. Distribution & Taxonomy: Because HALLERMANN (2005) did not examine the ZMA specimens collected by WEBER, he stated that older records of B. jubata from the Talaud Islands by DE Roov (1915) may be correct. How- ever, 14 specimens from Sulawesi formerly deposited in the Leiden collection (RMNH 3021la-e, 3023a-d, and 7419a-e) as B. jubata, were redetermined by HALLERMANN (2005) as B. celebensis, which is endemic to Sulawesi. In turn, three older records of B. jubata from Sulawesi in the Berlin collection (ZMB) are correct according to this au- thor. From the entire Philippine Archipelago only one voucher specimen from Mindanao (ZMB 16305) was in- cluded in the review by HALLERMANN (2005). Four fur- ther specimens (ZMB 34117) allegedly from the Philip- pines lack specific locality data. Thus, 1t remains uncer- tain if two different species of Bronchocela live in sym- patry on the Talaud Islands. Pending molecular genetic studies and further field work will hopefully reveal the sys- tematics and the exact distributions of these morpholog- ically diverse agamids. Hydrosaurus sp. (Fig. 8) Material examined: MZB Lac. 5081 (AK049). Distribution: De JonG (1928) reported Hydrosaurus am- boinensis for Karakelong Island. This is the first record of this large agamid for Salibabu Island. Ecology: During recent field surveys on Salibabu Island three adult specimens were flushed out. One female spec- imen was observed sitting on a tree with remarkable thin branches. One juvenile voucher specimen was encoun- tered near a small fish pond. On Talaud, Hydrosaurus dis- plays a cryptic life style and seems to avoid the coast line. This agamid species prefers less anthropogenically influ- enced inland habitats with freshwater environment. Morphology: SVL 116.5 mm, TL 243 mm; tympanum di- ameter 3.1 mm; dorsal crest missing, nuchal crest indicat- ed by a row of 16 prominent scales; six to seven interocu- laria; 11 supralabials, sixth to eleventh under eye; 11 in- fralabials; mental large; one row of six and seven enlarged submaxillaries, respectively, separated from infralabials by one (first IL) or several small scales; lateral scales hetero- geneous, mainly small and strongly carinate, interspersed 114 André Koch et al.: A Revised Checklist of the Herpetofauna of the Talaud Archipelago, Indonesia Fig. 8. Hydrosaurus sp. from Salibabu Island. Photo by André Koch. with six groups of two to six tubercular, carinate scales; three further such enlarged scales along the side of the neck; towards the ventral side numerous enlarged, smooth scales arranged in more or less distinct transverse rows; ventral scales homogenous, smooth; subdigitals under fourth toe small at basis, followed by 35 relatively en- larged subdigitals after first phalanx; toes laterally with a row of extremely wide (up to 1.9 mm) scales, 36 at fourth toe forming a serrated edge. Ten and eleven femoral pores, respectively. Ground color green to brownish with en- larged scales of the dorsal and lateral side being lighter; ventral side whitish, throat darker. Taxonomy: Phenetically, the Talaud specimen resembles a juvenile A. pustulatus from Panay Island, central Philip- pines, in color pattern and scalation (photo courtesy of M. Gaulke). However, a systematic assignment of the Talaud population either to A. amboinensis from Sulawesi, H. pustulatus from the Philippines, A. weberi from Halma- hera or to another hitherto unrecognized taxon seems pre- mature at this time, due to the lack of diagnostic charac- ters (dorsal and nuchal crests, adult color pattern) in the juvenile voucher specimen and insufficient material for comparison of the three Hydrosaurus taxa currently re- cognized. Gekkonidae Cyrtodactylus cf. jellesmae (Boulenger, 1897) Material examined: MZB Lac. 5126 (AK056), MZB Lac. 5128 (AK055). Additional material: ZMA 15942 (1 spec.), Gunung Doeata (= Mt. Doata), Karakelong, coll. H. J. LAM 1926; ZMA 17884 (1 spec.), same data as previous specimen. Distribution: DE JonG (1928) reported one specimen (but see “additional material”) of C. marmoratus from Karake- long Island. However, BRONGERSMA (1934) showed that Sulawesi specimens of C. marmoratus were referable to C. fumosus. Accordingly, ISKANDAR & NIO (1996) doubt- ed the occurrence of C. marmoratus on Sulawesi as pre- viously mentioned (e.g., DE Roo 1915). This is the first record of C. cf. jellesmae for Salibabu Island and the northernmost occurrence of the entire range of this species. Ecology: Both voucher specimens, apparently a female and a male (the female carrying two well-developed eggs which are visible through the thin skin), were found at 20.30 p.m. at low elevations on the trunks of a tree and a coconut palm, respectively. Morphology: Both sexes lack preanal and femoral pores and the ear openings are oval as characteristic for C. jelles- mae (BOULENGER 1897; DE Root 1915). Lateral fold pres- ent between axilla and groin. However, in coloration and scalation the Salibabu population differs from those of Sangihe Island and north Sulawesi. The characteristic dark, V- and M-shaped dorsal markings are less distinct and the tubercles have light tips only laterally. In addition, the Talaud population shows less pronounced tubercles on dorsum, legs and tail. From the new described species, C. wallacei, the Talaud geckos are distinguished by the ab- sence of enlarged subcaudal scales (HAYDEN et al. 2008). Taxonomy: Probably DE JONG’s (1928) specimen from the Talaud Islands will prove to belong to C. cf. jellesmae which was frequently found on Sulawesi and adjacent is- lands compared to C. fumosus and C. wallacei (unpubl. data). Future molecular studies are required to enlighten the systematics and taxonomy of this morphologically cryptic complex of gecko species in the Sulawesi region. Gehyra mutilata (Wiegmann, 1834) Material examined: MZB Lac. 5124 (AK046). Distribution: Gehyra mutilata is herein reported for the first time for the Talaud Islands. This invasive gecko species inhabits Southeast Asia, Oceania, Madagascar, Mexico, California, Hawaii, and New Zealand. It was probably accidentally introduced from the Philippines or North Sulawesi by human transportation. Bonner zoologische Beitráge 56 (2007) 115 Ecology & Morphology: The voucher specimen, a female containing two eggs, was collected on the wall of a house in Lirung village where several specimens were observed. SVL 50 mm, TL 53 mm. Gekko vittatus Hottuyn, 1782 Material examined: none Distribution: In the distribution table of “East Indian” herptiles, BARBOUR (1912: 179) listed the Talaud Islands within the range of Gekko vittatus. This gecko species is known from the Moluccas, New Guinea and surrounding archipelagos. Thomas BARBOUR himself did not visit the Talaud Islands but he was aware of VAN KAMPEN’s (1907) account about amphibians based on Max WEBER 's collec- tions from that island group. The correctness of BAR- BOUR’s (1912) data cannot be proven as no information on voucher specimens was provided. Theoretically, how- ever, the occurrence of this gecko species on the Talaud Archipelago seems possible as other Australopapuan species also inhabit this island group. Future investigations in the herpetofauna of the Talaud Islands may answer this question. Hemidactylus frenatus Duméril & Bibron, 1836 Material examined: none Additional material: MCZ R-45768, and R-45769, Karakelong, coll. FAIRCHILD Expedition 1940. Distribution & Ecology: This circum-tropically distrib- uted gecko species was frequently seen on walls of hous- es. Our observations suggest that A. frenatus and G. mu- tilata are more common on Talaud than on neighboring Sangihe Island. Two voucher specimens (see ‘additional material’) of the FAIRCHILD Expedition are deposited in the MCZ collection. Nactus pelagicus complex (Fig. 9) Material examined: MZB Lac. 5086 (AK075). Ecology: During daytime one specimen was found under a rotten tree trunk in a hilly area near Lirung on Salibabu Island. Morphology: SVL 54 mm; tail regenerated, 55 mm long, fifth toe of left hind limb missing; probably a female, pre- anal pores missing; dorsum, limbs and tail (except for re- generated part) covered with granules and conical tuber- Fig. 9. Nactus cf. pelagicus from Salibabu Island. Photo by An- dré Koch. cles forming six regular longitudinal rows on the back, ir- regularly arranged along the lateral sides; rostral large; on- ly 5 supralabials (as compared to other populations); 7 in- fralabials; mental large, a pair of small chin-shields pos- terior to the mental; ventral side covered with small gran- ulous scales; 19 subdigits under fourth toe. Coloration of dorsal side dark brown with blackish blotches on the back, increasing in number towards the snout, labial sutures whitish; ventrally dark grayish-brown, regenerated tail with irregular light markings. Distribution & Taxonomy: This is the first record of the genus Nactus for the Talaud Islands representing the north- ern most population of this widespread Pacific species group which is known to consist of bisexual and partheno- genetic species (Moritz 1987; DONNELLAN & MORITZ 1995; ZuG 1998). The taxonomic status of most New Guinean population is still uncertain (KRAUS 2005; JACK- MAN et al. 2008). Traditionally, they have been assigned to N. pelagicus and N. vankampeni, but unisexual N. pelagicus have been shown to be restricted to southern Vanuatu, New Caledonia, and Oceania (ZUG & MOON 1995). TANNER (1950) reported a specimen from Moro- tai close to Halmahera which was obviously a male be- cause it “(...) has an angular series of 7 preanal pores”. Scincidae Emoia a. atrocostata (Lesson, 1826) (Fig. 10) Material examined: MZB Lac. 5130 (AK071). Additional material: RMNH 18659, Karakelong, coll. Snellius Expedition (Dr. H. BoscHma), 14-21 June 1930. 116 André Koch et al.: A Revised Checklist of the Herpetofauna of the Talaud Archipelago, Indonesia Fig. 10. Emoia a. atrocostata from Salibabu Island. Photo by André Koch. Distribution: According to BROWN (1991) the Talaud population of E. atrocostata belongs to the nominotypic subspecies. He examined one specimen from Karakelong (RMNH 18659). This coastal skink species is widespread ranging from the Philippines through the Indo-Australian Archipelago and many Pacific island groups such as the Carolines and Palau reaching North Australia. This is the first record of E. a. atrocostata for Salibabu Island. Ecology: The voucher (MZB Lac. 5130), a weak adult specimen infested with several red mites (Acari), was rest- ing on coral limestone with temporary ponds at the shore- line where this species was frequently found on Salibabu Island (Fig. 2). Early activity was observed at 7 a. m. al- though the sky was cloudy and temperatures were rela- tively low. Morphology: SVL 85.4 mm, tail short, laterally com- pressed, 73.6 mm long; TL/SVL 0.86; seven supralabials, fifth enlarged and below eye; tympanum small; lower eye- lid with a transparent disk; one pair of nuchals; 38 mid- body scale rows; 33 rounded lamellae under fourth toe. Emoia caeruleocauda (de Vis, 1892) Material examined: MZB Lac. 5114 (AK043), MZB Lac. 5115 (AK041), MZB Lac. 5118 (AK042), MZB Lac. 5127 (AK069). Additional material: MCZ 45774, coll. FAIRCHILD Expe- dition, 1940; ZMA 18384 (1 spec.), Liroeng (= Lirung), Salibabu, coll. M. WEBER 1899; ZMA 18391 (as Lygoso- ma triviale; 2 spec.), Beo, Karakelong, coll. H. J. LAM 1926; ZMA 18392 (as Lygosoma triviale; 5 spec.), Gu- nung Doeata (= Mt. Doata), Karakelong, coll. H. J. LAM 1926; ZMA 18395 (as Lygosoma triviale), Liroeng (= Lirung), Salibabu, coll. H. J. Lam 1926; ZMA 18409 (as Lygosoma triviale; 1 spec.), Beo, Karakelong, coll. M. WEBER 1899. Distribution & Taxonomy: The Talaud population was formerly recognized as E. cyanura (DE Root 1915; DE JONG 1928), which in contrast to E. caeruleocauda is widely distributed in Pacific islands from the Bismarck Archipelago eastwards (BROWN 1991). BROWN (1991) re- vised the genus and identified one specimen (MCZ 45774) from Karakelong as E. caeruleocauda. Morphology: According to Brown (1991), these phenet- ically similar species (E. cyanura and E. caeruleocauda) can be distinguished by the shape and number of lamel- lae under the fourth toe. Two specimens from Salibabu (MZB Lac. 5115, 5127) each had 44 lamellae under the fourth toe, thus confirming the occurrence of E. caeruleo- cauda on the Talaud Archipelago. Lower eyelid with a transparent disk; hind limb reaches the axilla. Ecology: On Salibabu Island £. caeruleocauda was fre- quently found on the ground and low vegetation. It is the most common lizard of the Talaud Archipelago. These conspicuous lizards use their metallic-blue tail fluttering from side to side to distract potential predators, mark male territories and attract potential mates. Eugongylus rufescens (Shaw, 1802) Material examined: none Additional material: ZMA 12491 (4 spec.), Gunung Doeata (= Mt. Duata), Karakelong, coll. H. J. Lam 1926; ZMA 12492 (1 spec.), Lirung, Salibabu, coll. M. WEBER 1899 Distribution: DE Root (1915) listed Eugongylus rufescens for Salibabu Island and DE JonG (1928) report- ed another specimen of this skink from Karakelong (ac- cording to the online ZMA database 4 specimens are sum- marized under collection number ZMA 12491). We could not find this species during our field work. The Talaud populations represent the most north-western distribution of this short-legged skink which inhabits New Guinea, the Solomon and Admirality Islands finally reaching north- ern Australia. Eutropis m. multicarinata (Gray, 1845) (Fig. 11) Material examined: MZB Lac. 5084 (AK063), MZB Lac. 5085 (AK062), MZB Lac. 5087 (AK061), MZB Lac. 5116 (AK060), MZB Lac. 5117 (AK058). Bonner zoologische Beitráge 56 (2007) 117 Fig. 11. Eutropis m. multicarinata from Salibabu Island. Pho- to by André Koch. Additional material: ZMA 18454 (1 spec.), Beo, Karakelong, coll. M. WEBER 1899. Distribution: DE Roo1 (1915) recorded Eutropis multi- carinata (at that time still in the genus Mabuya, but tax- onomically partitioned by MAUSFELD et al. [2002]) from Karakelong Island. Our findings represent the first record of this species for Salibabu Island. Thus, on the Talaud Islands E. multicarinata has its most south-eastern distri- bution. Ecology: Five specimens were collected in the vicinity of Lirung (partly at night, 8.30 p.m.) while resting at the base of a tree. During daytime, brown skinks, though undeter- mined, climbed up tree trunks when chased. These obser- vations suggest that E. multicarinata is adapted to a se- mi-arboreal life style. Morphology: According to DE Roo (1915), E. multicar- inata can be distinguished from the phenetically similar species E. rudis and E. multifasciata by the absence of postnasal scales and five or seven strong keels on nuchals, dorsals and laterals. According to BROWN & ALCALA (1980) E. multicarinata has 22 to 29 lamellae (usually 24- 28) beneath the fourth toe, while £. multifasciata has on- ly 16 to 21, and E. rudis 18 to 22. Furthermore, these au- thors provide the following scalation data. E. multicari- nata has 28 to 32 scales around midbody, while E. mul- tifasciata has 30 to 34, and E. rudis 28 to 32. All speci- mens match the definitions by DE Roon (1915) and BROWN & ALCALA (1980) in having six or seven keels on dorsals (fused nuchals with up to 11 keels!), 22 to 25 lamellae be- neath fourth toe, and 32 to 33 scales around midbody. At least in one specimen (MZB Lac. 5087) the postnasal is absent. However, in MZB Lac. 5084 and MZB Lac. 5085 a postnasal is present. The prefrontals are separated by the frontal. The dorsal side is brown, sometimes interspersed with few darker spots, laterally with a dark brown streak from behind the eyes along the body, sometimes bordered above by a light brown streak; ventral side light yellow- ish to grey, throat in three specimens bluish-grey; supral- abials also grayish. The Talaud population of E. multicar- inata can be assigned to the nominal subspecies because the parietals are separated by the interparietal. All vouch- er specimens, however, lack blackish spots or blotches be- low the head and throat as characteristic for E. m. bore- alis (BROWN & ALCALA 1980). Eutropis multifasciata (Gray, 1845) Material examined: none Additional material: MCZ R-45772 and MCZ R-45773, Karakelong, coll. FAIRCHILD Garden Expedition 1940. Distribution: Eutropis multifasciata, a widespread species known also from Sulawesi, the Philippines and Halmahera in the Moluccas, was not found during this biodiversity survey. However, more than 60 years ago, the FAIRCHILD Garden Expedition collected two specimens (see “addi- tional material”) of this skink on Karakelong which have not yet been reported. We provide here the first record of this skink from the Talaud Island. In addition, H. J. LAM collected this species on Miangas north of the Talaud Is- lands (DE JONG 1928). Taxonomy: Although £. multifasciata in known to occur sympatrically with 1ts congeners E. multicarinata and E. rudis on Mindanao in the Philippines (BROWN & ALCALA 1980), the identification of the FAIRCHILD voucher speci- mens should be re-confirmed due to the observed differ- ences in scalation characters in E. multicarinata and E. cf. rudis (see below). Eutropis cf. rudis (Boulenger, 1887) (Fig. 12) Material examined: MZB Lac. 5142 (AK044), MZB Lac. 5113 (AK070). Ecology: Two subadults (MZB Lac. 5142: SVL 36.8 mm, TL 50.0 mm; see Fig. 12) were collected near Lirung on Salibabu Island in a coconut and nutmeg plantation and on the beach, respectively. Morphology & Taxonomy: To distinguish E. rudis from E. multifasciata, BROWN & ALCALA (1980) mention that the prefrontals are in broad contact in the latter species while they are usually narrowly separated (or rarely in con- tact) in E. rudis. Aside this scalation character, both vouch- er specimens largely agree with the definition by BROWN 118 André Koch et al.: A Revised Checklist of the Herpetofauna of the Talaud Archipelago, Indonesia Fig. 12. Eutropis cf. rudis from Salibabu Island. Photo by André Koch. & ALCALA (1980) in having 20 and 22 keeled lamellae un- der fourth toe, 32 and 30 midbody scale rows (usually 28-30, rarely 31 or 32; 28-31 in E. rudis from Sulawesi according to HOWARD et al. [2007]), respectively. A post- nasal is present. The hind limbs of MZB Lac. 5142 reach the axilla as postulated by DE Roon (1915), while MZB Lac. 5113 exhibits tricarinate dorsals as typical for £. rud- is. The second specimen’s dorsal scales have up to five keels (very rarely in E. rudis according to BROWN & AL- CALA [1980]), of which the three medians are strongly pro- nounced and flanked by two feeble outer keels. The Ta- laud specimens share this character with the newly de- scribed E. grandis from Sulawesi (HOWARD et al. 2007), from which, however, they are clearly distinguishable by higher midbody scale counts (30-32 vs. 25-27 in E. gran- dis). In contrast to BROWN & ALCALA (1980) and HOWARD et al. (2007) both vouchers have 40 paravertebral rows (vs. 34-38 in Philippine £. rudis and 34-35 in Sulawesian E. rudis, respectively) between parietals and base of tail. The dorsal side is brown, laterally with a dark brown streak starting behind the nostrils, bordered above by a light brown streak; ventral side light grey, supralabials also grayish. In specimen MZB Lac. 5113 the ventral side is light gray only on the first half of the body; the rest in- cluding hind limbs and tail is brownish. Because both voucher specimens are not fully grown, the identification as either E. rudis or E. multifasciata is not unequivocal. Deviations in vertebral scale counts from those provided by BROWN & ALCALA (1980) may be ex- plained by geographic variation. Therefore, we here pre- liminarily allocate them to E. cf. rudis. Further investiga- tions will reveal if E. rudis really inhabits the Talaud Is- lands. Lamprolepis smaragdina ssp. (Lesson, 1830) (Fig. 13) Material examined: none Additional material: ZMA 11304 (1 spec.), Karakelang (= Karakelong), coll. H. J. Lam 1926; ZMA 18231 (1 spec.), Liroeng (= Lirung), Salibabu, coll. H. J. LAM 1926. E LATA Fig. 13. Lamprolepis smaragdina ssp. from Salibabu Island. Photo by André Koch. Bonner zoologische Beitráge 56 (2007) 119 Distribution & Taxonomy: DE JONG (1928) reported two specimens of L. smaragdina from the Talaud Islands. MERTENS (1929), who examined DE JONG’s voucher spec- imens mentioned that despite their minor geographic dis- tance to the Philippines the Talaud specimens (and even those of the Miangas further north to Mindanao) resem- ble L. smaragdina viridipuncta inhabiting the Pacific arch- ipelagos of Palau, the Carolines, and eastward to the Mar- shall Islands. Nevertheless, he allocated the Talaud pop- ulation to the Philippine subspecies L. s. philippinica. MERTENS (1929) remarked that one specimen from Lirung, Salibabu, was the largest of more than 100 vouch- er specimens he examined from all over the species” range; 1ts SVL measured 114 mm, and the black markings typi- cal of Philippine populations were largely reduced. This pattern of L. s. viridipuncta was also found in the only specimen from Mindanao available to MERTENS (1929). He finally designated the Talaud and Miangas populations as a hybrid form of the Philippine and Pacific taxa. Morphology: Observations and photographs of a few adult specimens active on trees and coconut palms near the coast of Lirung match MERTENS’ (1929) description of the Salibabu specimen. The uniformly green coloration without any markings is different from populations of L. smaragdina on both neighboring islands of Sangihe and north Sulawesi. No specimens of the viridipuncta sub- species were examined. Lipinia noctua (Lesson, 1826) Material examined: none Additional material: ZMA 18348, Gunung Doeala (= Mt. Doala), Karakelong, coll. H. J. LAM 1926; MCZ R-45775, Karakelong, coll. FAIRCHILD Garden Expedition 1940 Distribution: The record for Talaud by DE JonG (1928) was not mentioned by ZWEIFEL (1979) who examined the variation in the moth skink, Lipinia noctua. A second spec- imen (MCZ R-45775) was collected by the FAIRCHILD Ex- pedition. The Talaud population represents the most north- westerly distribution for this cosmopolitan Pacific species. During our survey we could neither record this species, nor could we examine DE JONG’s voucher specimen. Varanidae Varanus sp. (aff. indicus Daudin, 1802) (Figs 3, 14) Material examined: MZB Lac. 5176 (AK067), MZB Lac. 5177 (AK064), MZB Lac. 5178 (AK059), MZB Lac. 5179 (AK066), MZB Lac. 5180 (AK065); MZB Lac. 581 (954), \ > SC Es \ 4 ee Fig. 14. Varanus sp. from Salibabu Island. Photo by André Koch. Karakelong Island, coll. H. J. LAM 1926; MZB Lac. 4195, Beo, Karakelong Island, coll. J. RILEY of WCS (World Conservation Society) 9 June 1999; ZMA 15411b, Lirung, Salibabu, coll. M. WEBER and Siboga Expedition 25-27 July 1899; ZFMK 87587 (formerly ZMA 1541 1a), same data as previous specimen. Distribution: The Talaud population represents the most north-westerly distribution of a member of the Varanus indicus species group (ZIEGLER et al. 2007). Interesting- ly, AUFFENBERG (1980: 109) listed two specimens of Y sal- vator from Liroeng (= Lirung) on Salibaboe (= Salibabu) Island in the collection of the Museum Zoologicum Bo- goriense (MZB Lac. 947 and 954, respectively). These vouchers, however, could not be located in the MZB col- lection. According to the MZB herpetology catalogue, the collection numbers to which AUFFENBERG (1980) referred belong to a specimen of V. cf. nebulosus and V. indicus, respectively (I. SIDIK pers. comm.). The latter was collect- ed by H. J. Lam on Salibabu Island in 1926. According- ly, we could not confirm a potential co-existence of Pa- cific monitor lizards with the water monitor, V. salvator, on Salibabu Island. Although local people reliably stated that there are two giant reptile species found on the island, the description of the second species fits exactly with the large sailfin lizard, Hydrosaurus. Ecology: On Salibabu Island monitor lizards are frequent- ly found in disturbed and cultivated areas like coconut plantations near the coast line. Thus, a preference for salt- water influenced habitats as reported for Y. indicus on New Guinea (PHILIPP 1999) could also be confirmed for the Ta- laud monitor lizards. Five mature specimens were found during day time on palm or tree trunks while basking or mating (Fig. 3). Only once, a juvenile specimen was flushed out near a runnel but escaped in the dense vege- tation. These observations suggest an ecological separa- 120 André Koch et al.: A Revised Checklist of the Herpetofauna of the Talaud Archipelago, Indonesia Fig. 15. Defecated prey remains from specimen MZB Lac. 5178 of Varanus sp. representing a large orthopteran (Sexava sp., Tet- tigoniidae) and fragments of a crab. tion of different age groups in the Talaud monitor lizards. While juvenile specimens seem to hide in thicket on the ground, adults obviously prefer a semi-arboreal life. The avoidance behaviour of juveniles seems reasonable be- cause cannibalism is common in many monitor lizard species. Although monitor lizards are regularly hunted for food supply by local people, no direct threat of the Salibabu population could be recorded. Natural predators are scarce as no large carnivorous mammals are known from these islands (RILEY 2002). However, crocodiles (Crocodylus sp.) and the reticulated python (Python reticulatus) should be considered natural predators. Diet: After being caught specimen MZB Lac. 5178 defe- cated the undigested remains of a small crab and a large green orthopteran (Sexava sp., Tettigoniidae) (Fig. 15). Furthermore, the stomach of ZFMK 87587 contained a well-preserved specimen of a large spider (“Olios” coc- cineiventris group, Deleninae) and a fragmented crab. These findings are in congruence with our observations of an arboreal-aquatic life habit in the Talaud monitor lizards. Parasites: Some specimens exhibited several ticks (prob- ably Amblyomma sp.) on the ventral side mostly near the umbilical region and at the base of the tail. Reproduction: Mating behaviour was observed near the coast only two meters alongside a dirt road at 10.30 a. m., 19 July 2005. Two monitor lizards (MZB Lac. 5177 and MZB Lac. 5180) in mating position were discovered at a palm trunk about six meters above ground. When a local collector climbed up the coconut palm, the pair uncoupled and both specimens tried to hide in the crown. One of the specimens even jumped down the palm to escape. This be- haviour was observed twice while chasing monitor lizards on Salibabu Island. In addition, two voucher specimens (ZFMK 87587 and MZB Lac. 5177) contained two and three eggs, respec- tively, suggesting that the reproductive period of Talaud monitor lizards occurs throughout May, June and July at least. The two small eggs are ca. 8 x 17 mm in size, while the three well-developed eggs measure 29 x 63 mm, 28 x 63 mm, and 27 x 61 mm, respectively. Bonner zoologische Beiträge 56 (2007) 121 Taxonomy: DE Roon (1915), DE JONG (1928) and BRAN- DENBURG (1983) referred the Talaud population to the Pa- cific monitor lizard, V. indicus. However, due to particu- lar characteristics in color pattern and pholidosis togeth- er with molecular genetic evidence (ZIEGLER et al. 2007), the Talaud monitor lizards represent a new member of the V. indicus species group (KOCH et al. subm.). SERPENTES Typhlopidae Typhlops sp. Material examined: MZB Serp. 3227 (AK045). Distribution: First record of a blind snake species for the Talaud Islands. Ecology: Two specimens were found under a coconut shell in a plantation near the coast of which one specimen was preserved. Morphology & Taxonomy: ToL 130 mm; body width 2.2 mm; 255 ventrals, 13 or 14 subcaudals. The voucher spec- imen resembles the drawing of the holotype of 7 hedraeus (CAS-SU 12346) depicted by IN DEN BOSCH & INEICH (1994) in having the eye restricted to the ocular scale not reaching the suture to the preocular; a subocu- lar, in contrast to 7. ater from Sulawesi, the Moluccas and New Guinea, is absent. However, the voucher differs from T. hedraeus in having two instead of one preocular and the latter is not smaller than the ocular. The upper jaw is not visible laterally. 7. hedraeus is found on several Philip- pine Islands including Mindanao, Luzon and Negros, the type locality. Due to differences in morphology, the Ta- laud population may represent a distinct species. Boidae Candoia paulsoni tasmai Smith & Tepedelen in Smith et al. 2001 (Fig. 16) Material examined: MZB Serp. 2949, Niampak, Karake- long Island, 21 March 1999; MZB Serp. 2950, Essang, Karakelong Island, coll. WCS 5 April 1999. Additional material: MCZ 45767 (paratype), Karakelong, coll. FAIRCHILD Garden Expedition 1940; ZMA 16937, Gunung Doeata (= Mt. Duata), Karakelong, coll. H. J. LAM 1926; ZMA 16943 (4 spec.), Liroeng (= Lirung), Salibabu, coll. M. WEBER 1899 Distribution: Beside Talaud, C. paulsoni tasmai inhab- its Halmahera and surrounding islands as well as the tip of North Sulawesi (SMITH et al. 2001; DE LANG & VOGEL 2005). Consequently, the Talaud Islands are the most north-western population of this subspecies which is sep- arated by about 800 km from the remaining subspecies of C. paulsoni on New Guinea and the Solomon Islands (SMITH et al. 2001). In contrast, C. carinata is found on Sangihe Island enclosed between C. paulsoni on the Ta- laud Islands in the north and the northern peninsula of Su- lawesi in the south. However, historical records of Can- doia from the “Minahassa” peninsula, North Sulawesi, by MEYER (1887) and independently by BOETTGER (1898) as adopted by BOULENGER (1897), DE Roo (1917), STERN- FELD (1920), MCDOWELL (1979), IN DEN BOSCH (1985), ISKANDAR & NIO (1996), SMITH et al. (2001), and DE LANG & VOGEL (2005) may be incorrect as was repeatedly as- sumed for some others of MEYER 's vertebrate collections (see HICKSON [1889: 158] and STRESEMANN [1939: 305] for birds and FEILER [1990] for mammals, respectively). Confusion of locality data may have easily occurred since MEYER did not exclusively undertake his collection on the Minahassa peninsula of Sulawesi, but also purchased many voucher specimens from Charles W. CURSHAM, a trader in Manado, who in turn sent out native hunters (MEYER & WIGLESWORTH 1898). Intense fieldwork in North Sulawesi in recent years has revealed no voucher specimens confirming the historical records (J. MCGUIRE pers. comm., KOCH 8 ARIDA unpubl. data). It ıs, therefore, reasonable to suggest that the “Minahassa” specimen (not present in the MTD collection and most probably lost dur- Fig. 16. Candoia paulsoni tasmai (MZB Serp. 2950) from Ka- rakelong Island. Photo by André Koch. 122 André Koch et al.: A Revised Checklist of the Herpetofauna of the Talaud Archipelago, Indonesia ing World War II, U. Fritz & E. LEHR pers. comm.) ac- tually originated from Talaud or Halmahera with its satel- lite islands, the actual range of C. paulsoni tasmai. The same may apply to BOETTGER’s (1898; 1903) specimen (collected by W. KÜKENTHAL, presumably in the Senck- enberg Museum collection, Frankfurt), which also lacks exact locality data. Morphology: Although we did not find this snake species during our survey, two specimens (see “additional mate- rial”) collected on Karakelong Island by a team of WCS lead by Jon RILEY, were examined in the MZB collection. These specimens (SVL 524/650 mm, TL 78/80 mm; TL/SVL+TL: 0.13/0.11) fit the definition by SMITH & TE- PEDELEN in SMITH et al. (2001) in having 30/31-37/38-26/26 scales around the body (anterior-mid- body-posterior), 179/183 ventrals, 36/36 subcaudals, on- ly one enlarged supraocular reaching the orbit, 11/11 cir- cumorbitals, 10/10 supralabials (6-7/5-7 contacting orbit), 10/13 infralabials, dorsals keeled except for the second to fifth lateral rows; a distinctive white postanal spot is miss- ing. The color pattern of the specimens varies. Specimen MZB Serp. 2949 shows a reddish-brown background col- or with darker blotches indicating a faded zigzag band dor- sally, the head becoming dark brown towards the tip; ven- tral side beige with brown mottling or blotches. In con- trast, specimen MZB Serp. 2950 (Fig. 16), like the Talaud specimen examined by SMITH et al. (2001), has a relative- ly light beige background color with a distinctive dark red- brown zigzag band dorsally and indistinctive brown blotches laterally; markings of the tail are dark brown. The ventral side shows nearly the same coloration as the dor- sal with darker mottling and blotches. Taxonomy: Earlier authors (e.g., DE ROO 1917, DE JONG 1928, DE Haas 1950, MCDOWELL 1979) referred speci- mens from the Talaud Islands to Candoia carinata (for- merly known as Enygrus carinatus), which was later split into three different species, 1.e., Candoia carinata, C. su- perciliosa, and C. paulsoni with several subspecies (SMITH et al. 2001). Based on only one voucher specimen (MCZ 45767 from Karakelong), these authors allocated the Talaud population of bevelnosed boas to a new sub- species, C. paulsoni tasmai. Pythonidae Python reticulatus (Schneider, 1801) Material examined: none Distribution: This widespread snake species inhabits con- tinental South and Southeast Asia, the Philippines and In- donesia, east to the Moiuccas and the Tanimbar Islands. The reticulated python was reported from Bitunuris on Salibabu Island (DE LANG & VOGEL 2005). Ecology: From Salibabu Island it was reported that a python approximately 5 m long had killed a child (Kop- STEIN 1927, 1930). DE LANG & VOGEL (2005) recounted several cases of persons devoured by pythons. They ex- trapolated that every year at least one human is eaten in Sulawesi by a reticulated python. Taxonomy: As was shown by AULIYA et al. (2002), even small island populations like those from Tanahjampea and Selayar, located between the south-western peninsula of Sulawesi and Flores in the Lesser Sunda Islands, show subspecific differentiations. The Sangihe Island popula- tion also represents a distinct lineage (AULTYA et al. 2002). Therefore, the systematic status of the Talaud population compared to those of the Philippines, Sulawesi and the Moluccas is unclear and deserves taxonomic revision. Colubridae Remarks: One medium-sized brown snake disappeared under stones when encountered in a narrow gorge with a small stream (Fig. 4). Another small brown snake was en- countered near the root of a tree, indicating that, apart from the following species, further unknown snakes inhabit the Talaud Islands. Boiga irregularis (Bechstein, 1802) Material examined: MZB Serp. 3238 (AK048). Distribution & Ecology: The brown tree snake Boiga ir- regularis is a widespread species known from most islands east of Sulawesi, and reaching Australia. In some Pacif- ic islands where B. irregularis was accidentally intro- duced, it has been responsible for a dramatic decline in the native vertebrate faunas (e.g., RODDA & FrITTS 1992; RoDDA et al. 1997). One road kill was found in the vicin- ity of Lirung, Salibabu Island. This is the first record of B. irregularis for the Talaud Archipelago. Morphology: One preocular, two postoculars, one lore- al; eight supralabials, the fourth to sixth in contact with orbit; ten and eleven infralabials, respectively; 19 scales around body, about 247 ventrals and 115 subcaudals; the vertebrals are slightly enlarged. The specimen shows a nar- row pattern of indistinct light and dark grey cross-bands particularly on the anterior half of body as is typical for some B. irregularis. Bonner zoologische Beitráge 56 (2007) 123 Cerberus r. rynchops (Schneider, 1799) Material examined: none Additional material: RMNH 6262, Karakelong, coll. Dr. H. BoscHma 14-21 June 1930. Distribution: This widespread homalopsine snake is known to occur from continental Southeast Asıa through the Philippines and Indonesia reaching New Guinea (GyY1 1970; ALFARO et al. 2004). DE JonG (1928) reported the species for Karakelong as did BRONGERSMA (1934) based on a specimen (see “additional material”) collected in 1930. While DE Haas (1950) listed only the Talaud Islands in the species account of C. rynchops, Salebabu (= Salibabu) Island is mentioned in the distribution appendix. Taxonomy: Gy! (1970) recognized two subspecies: the widely distributed C. r. rynchops and C. r. novaeguineae from New Guinea. According to DE LANG & VOGEL (2005) the Talaud population belongs to the nominotypic sub- species. ALFARO et al. (2004), however, demonstrated that C. r. rynchops is split into four to five phylogenetic line- ages (India and Myanmar, Philippines, Greater Sunda Is- lands and Sulawesi, and the Thai-Malay Peninsula and Gulf of Thailand). Thus, the taxonomic status of the Ta- laud population remains unclear. CROCODYLIA Crocodylidae Crocodylus sp. Material examined: none Taxonomy: In 1997 two unidentified crocodiles were ob- served on Karakelong Island (J. RILEY, pers. comm. in PLATT & LEE 2000). Although currently the taxonomic sta- tus of the Sulawesi populations and hence that of the Ta- laud Archipelago is still unsolved, it is reasonable to sug- gest that the widespread Crocodylus porosus or another species, C. mindorensis from the Philippines, respective- ly, inhabits the Talaud island group (PLATT & LEE 2000; PLATT et al. 2007). Ecology: Reportedly, these crocodiles live in mangrove swamps which are threatened by destruction. However, ap- parently due to religious taboos villagers do not kill croc- odiles (J. WARDILL, pers. comm. in PLATT & LEE 2000). TESTUDINES Cheloniidae Chelonia mydas (Linnaeus, 1758) Material examined: none Ecology: Chelonia mydas is known to have nesting sites on the Talaud Islands (WHITTEN et al. 2002). Although no trade with sea turtles and their products was noticed on Salibabu Island, repeated observations were made in the markets of Manado and elsewhere on North Sulawesi that sea turtles and their eggs were sold for human consump- tion. 4. DISCUSSION Species diversity, island endemicity and nature conser- vation Some authors have pointed out the depauperate but high- ly endemic character of the nearby Sangihe and Talaud Archipelagos (e.g., LAZELL 1992; RILEY 2002). This state- ment partly contradicts the geological assumption that the Talaud block did not emerge above sea level until the Pleistocene (MOORE et al. 1981). Repeated drops in glob- al sea levels during the Pleistocene did probably not con- nect the Talaud Archipelago with the neighboring Sangi- he Islands, from which it is separated by a deep ocean trench, or with any other surrounding landmasses (VORIS 2000). Accordingly, LAZELL (1992) hypothesized that am- phibians and reptiles must have colonized Talaud by over- water dispersal. Faunal and floral colonizers of the Talaud Archipelago had presumably less than two million years to succeed or even to evolve endemic lineages. Thus, the degree of endemism of Talaud’s herpetofauna is relative- ly low (only 4 %). Currently, only Varanus sp. seems to be restricted to the island group but the number of Talaud endemics may increase by up to 27 % in the future, when the taxonomic status of some problematic specimens be- longing to widespread species or species groups has been clarified (e.g., Nactus cf. pelagicus, Cyrtodactylus cf. jellesmae, Hydrosaurus sp., Lamprolepis smaragdina ssp., Callulops cf. dubius, and Typhlops sp.). Detailed system- atic investigations including modern molecular techniques are needed to solve their taxonomic identity. As expected for small oceanic islands (STERNFELD 1920; BROWN 1957; ADLER et al. 1995; ALLISON 1996; CROM- BIE & PREGILL 1999), skinks (next to geckos) are the most diverse reptile group on the Talaud Islands, while amphib- ians are very rare due to their limited ability to cross salt- water barriers. However, no Sphenomorphus skinks were 124 found during our biodiversity survey on Salibabu Island although this taxonomically difficult and diverse genus is well represented on the adjacent islands of Sulawesi, Halmahera and Mindanao (BROWN & ALCALA 1980; ISKANDAR & NIO 1996; MONK et al. 1997). In addition, fly- ing lizards of the genus Draco were not observed during our field work, nor during previous expeditions to Salibabu Island (DE JONG 1928; McGuire et al. 2007). In contrast, LAZELL (1992) predicted the sympatric occurrence of two Draco species for Talaud, one of which he assumed to be a giant, based on biogeographic knowledge of the Lesser Antilles Anolis lizards. Therefore, future field surveys con- centrating on the forested areas of the larger island of Karakelong are needed to verify the presence of Draco spp. on the Talaud Islands. The same applies to Gekko vit- tatus and Bronchocela jubata, which have been reported from Talaud in the past (see above). Further unrecognized amphibian and reptile species may be anticipated to inhab- it the Talaud Archipelago. As in many Pacific island biotas and beyond, Boiga irreg- ularis, Gehyra mutilata, and Hemidactylus frenatus are most probably non-native, naturalized species of the Ta- laud Archipelago. Because these reptiles are not utilized by man, they have likely reached this island group via ac- cidental human-mediated transportation. Nevertheless, the ability to “use” boats and ferries for over-water dispersal renders these three species as candidates to cross far dis- tances on natural rafts as well. A determination between accidental human-mediated displacement on the one hand and natural colonization on the other seems virtually im- possible. In any case, the new record of the invasive brown tree snake B. irregularis for Talaud may have severe conse- quences for the native vertebrate fauna of this island group. As was demonstrated by several studies (e.g., SAVIDGE 1987; RoDDA & Fritts 1992; RODDA et al. 1997), this snake species is responsible for the dramatic decline in bird and lizard populations on Guam. This could be particu- larly fatal for endemic lizard species like the Talaud mon- itor lizards which have a very restricted distribution range (KOCH et al. subm.). SAVIDGE 1988 and McCorb et al. (1994) reported from Guam that B. irregularis even preys on juveniles of Varanus indicus. Therefore, the develop- ment of the Talaud population of this invasive snake species deserves special attention in the future. Deforestation and destruction of natural habitats represent another major threat to the amphibians and reptiles of the Talaud Archipelago as was pointed out by RILEY (2002) for the mammals of this island group. Coconut plantations replace natural vegetation in most areas. Therefore, a wildlife reserve has been established on the larger island of Karakelong to protect endemic faunas (RILEY 1997). André Koch et al.: A Revised Checklist of the Herpetofauna of the Talaud Archipelago, Indonesia Biogeographic relations and past dispersal routes Although it may seem premature, some conclusions can be drawn about the biogeographic origin and affinities of the Talaud Islands’ herpetofauna based on current distri- bution patterns of amphibians and reptiles of the Indo-Aus- tralian Archipelago. Firstly, several typical Australopapuan species or species groups (viz. Litoria infrafrenata, Can- doia paulsoni, Nactus cf. pelagicus, Eugongylus rufescens, Lipinia noctua, and Varanus sp. [aff. indicus]), reach their most north-western distributions on the Talaud Islands. These Australopapuan faunal elements probably arrived at this island group via Halmahera and adjacent islands in the northern Moluccas. Some species with Oriental origin, such as Eutropis multifasciata and Eutropis cf. rudis, have their most easterly distribution on or around the Talaud Is- lands. Consequently, the Talaud Archipelago’s biogeo- graphic position within the Wallacea transition zone is clearly warranted (see Cox 2001 for a recent review of the Wallacea region). Interestingly, few widespread Oriental species like Varanus salvator did not reach the Talaud Archipelago despite its abundance on Sulawesi and small- er off-shore islands. An explanation for this may be the flow of Pacific water into the Indian Ocean via the Indone- sian seas. This throughflow enters the Indonesian Islands in north-eastern to south-western direction between Min- danao and the Talaud group passing the Celebes Sea while another sea current from the Pacific circulates between Ta- laud and Halmahera (GORDON & FINE 1996, GORDON 2005), thus hampering over-water dispersal on natural rafts. Secondly, the herpetofauna of Talaud shows clear biogeo- graphic relations to Halmahera in the northern Moluccas and, to a lesser extent, to Sulawesi and to the Philippines. With the northern Moluccas the Talaud Archipelago has 20 amphibian and non-marine reptile species in common, while it shares only 14 amphibians and non-marine rep- tiles with Sulawesi and the Philippines, respectively (see Tab. 1). With regard to amphibians, two out of three Ta- laud species also occur on Halmahera and, at least in one case (Litoria infrafrenata), farther southeast on New Guinea and adjacent islands. In contrast, no frog species from the Philippines seem to have reached the Talaud Is- lands. Limnonectes modestus which inhabits Sulawesi is also found on the Talaud Islands and some islands of the Moluccas. For the terrestrial reptiles, four species (respec- tively five when Gekko vitattus is included) are exclusive- ly shared by the Talaud Archipelago and the northern Moluccas including areas further southeast. These are Can- doia paulsoni tasmai, Nactus cf. pelagicus, Eugongylus rufescens, and Lipinia noctua. Due to their central position in the Molucca Sea, there are no native species exclusively common to the Talaud Is- Bonner zoologische Beitráge 56 (2007) 125 Table 1. Distribution records of amphibians and reptiles from the Talaud Islands as compared to the surrounding islands of Sula- wesi, Halmahera and the Philippines. Probable non-native, i.e. invasive species of the Talaud Islands are indicated by a hash key (++). New records for the Talaud Island are marked with an exclamation mark (!); new records for Salibabu Island are denoted by an asterisk (*). An “(x)” denotes that different subspecies or closely related taxa of a species complex, respectively, are involved. l indicates that Nactus cf. pelagicus has only been recorded from Morotai but due to close proximity most probably also occurs on Halmahera. Locality records for surrounding islands are taken from BOETTGER (1895a, b; 1903), BOULENGER (1897), BARBOUR (1912), DE Roow (1915; 1917), DE JONG (1928), TANNER (1950), BROWN & ALCALA (1970), BROWN & ALCALA (1980), BROWN (1991), Monk et al. (1997), HALLERMANN (2005), and ZIEGLER et al. (2007). Locality Taxon Sulawesi Talaud Halmahera Philippines ANURA Hylidae (1 sp.) Litoria infrafrenata x x Microhylidae (1 sp.) Callulops cf. dubius x! x Ranidae (1 sp.) Limnonectes cf. modestus x Xx (x) (x) SAURIA Agamidae (3 spp.) Bronchocela cristatella x X x X Bronchocela jubata X x fa Hydrosaurus sp. (x) x” (x) (x) Gekkonidae (5 spp.) Cyrtodactylus cf. jellesmae X x (x) Gehyra mutilata # x x! x X Gekko vitattus 2 x Hemidactylus frenatus # x x x X Nactus cf. pelagicus x! x! Scincidae (8 spp.) Emoia a. atrocostata x > x x Emoia caeruleocauda x x X x Eugongylus rufescens x x Eutropis m. multicarinata x E Eutropis multifasciata x x! x x Eutropis cf. rudis x x! X Lamprolepis smaragdina ssp. (x) x (x) (x) Lipinia noctua x X Varanidae (1 sp.) Varanus sp. (aff. indicus) x (x) SERPENTES Typhlopidae (1 sp.) Typhlops sp. x! (x) Boidae (1 sp.) Candoia paulsoni tasmai 2 x x Pythonidae (1 sp.) Python reticulatus x x X x Colubridae (2 spp.) Boiga irregularis # x x! Xx Cerberus r. rynchops x x x A CROCODYLIA Crocodylidae (1 sp.) Crocodylus sp. x x x x TESTUDINES Cheloniidae (1 sp.) Chelonia mydas y j ee = a EB a: oe 126 André Koch et al.: A Revised Checklist of the Herpetofauna of the Talaud Archipelago, Indonesia lands and the Philippines and Halmahera, and only one species, Eutropis cf. rudis, in common with the Philippines and Sulawesi. However, some widespread non-invasive species like Cerberus rynchops, Python reticulatus, Bronchocela cristatella, Emoia atrocostata, E. caeruleo- cauda, Eutropis multifasciata, and Lamprolepis smarag- dina are found at all four localities. It seems plausible that such widespread species were exchanged between the Philippines and the Moluccas by using Sulawesi as a stopover rather than the Talaud Islands (but see below). Hence, the Philippines may have been colonized via the Sangihe Island chain or arrived at the Moluccas via the Banggai-Sula Islands finally reaching the Talaud Archi- pelago in the north. Nevertheless, it remains doubtful whether the Talaud Islands served as natural stepping stones (particularly during periods of lowered Pleistocene sea levels) on the dispersal route between the Philippines and North Sulawesi. INGER (1954) argued that the pres- ent distribution of the genus Oreophryne Boettger, 1895, which also occurs on Sulawesi and some islands of the Lesser Sundas may account in favor of the Moluccan-Ta- laud corridor. Oreophryne, however, was never recorded from the Talaud Islands. The same applies to the genus Cornufer Tschudi, 1838 (currently a synonym of Platy- mantis Gunther, 1858), which ranges from the Philippines through the Moluccas to New Guinea and the Solomons, but is not known from Sulawesi. According to INGER (1954), this taxon constitutes more evidence for the Ta- laud avenue between the Philippines and the Moluccas which also was later identified as major route of faunal dispersal by HOLLOWAY & JARDINE (1968). In either case, the extinction or replacement of former Talaud populations by later invaders may explain their absence from the is- land group. As noted by WHITTEN et al. (1987) biological evidence for a stepping stone hypothesis between the Philippines and the Moluccas via Talaud is still insubstan- tial. Only further field work and detailed morphological comparisons of sufficient material together with molecu- lar investigations including individuals from all areas in- volved could trace the unique history for each single species and the region. Comparison with other organism groups Distribution patterns of some other organism groups vary from the herpetological results presented here. VANE- WRIGHT & DE JONG (2003) traditionally included the Ta- laud Islands as part of the Sulawesi region in their review of Sulawesi butterflies. They predicted that colonization from the Philippines into North Sulawesi happened via Sangihe and Talaud. Likewise, NATUs (2005) in her analy- sis of endemic centers of Indonesian vertebrates defined the Sulawesi region as the mainland of Sulawesi plus sur- rounding island groups like Sangihe and Talaud in the north. Interestingly, our herpetological results contradict the findings in butterflies by VANE-WRIGHT & DE JONG (2003) who eventually concluded that the fauna of the Ta- laud Archipelago most probably “belongs” to the Min- danao fauna. With regard to mammals, the Talaud and Sangihe Islands are home to an endemic marsupial, the Talaud bear cus- cus (Ailurops ursinus melanotus). These populations rep- resent the northernmost subspecies of this Sulawesian pha- langer (FEILER 1977; 1990). At the same time, A. ursinus is the only example of a non-volant and non-invasive mammal that inhabits both neighboring archipelagos (RI- LEY 2002). Human translocation could be responsible for the exceptional distribution pattern of this species which is frequently hunted as food. On the other hand, both is- land groups harbor their own endemics, e.g., Tarsius san- girensis on Sangihe and Melomys caurinus on Talaud (RI- LEY 2002), which have their respective closest relatives on North Sulawesi and the northern Moluccas (FLANNERY 1995; MENZIES 1996; SHEKELLE 2003). There is another population of T. sangirensis on the nearby Siau Island, which may warrant a separate taxonomic status (BRAN- DON-JONES et al. 2004). As a result, these mammal taxa confirm the minor faunal similarity between Sangihe and Talaud, and at the same time, supply evidence for their close biogeographic relationships with Sulawesi and New Guinea, respectively. Consequently, the tradition to incorporate the Talaud Arch- ipelago into the Sulawesi subregion within the transition- al zone of Wallacea seems only conditionally justified from a herpetofaunistic point of view. Rather, this island group should be recognized as part of the northern Moluc- can subregion with closer zoogeographical links to Halmahera compared with Sulawesi or the Philippines. The transitional character of the herpetofauna of the Ta- laud Islands composed of both typical Oriental and Aus- tralopapuan faunal elements clearly reinforces the justi- fication of the Wallacea region between Southeast Asian and Australopapuan biotas. Acknowledgements. We would like to thank the Indonesian In- stitute of Sciences (LIPI) for the permission (2767/SU.3/KS/2005) to conduct research in Indonesia. The kind support of Pak Dedy Darnaedy (Research Center for Biology, LIPI) Pak Mulyadi and Pak Irvan Sidik (both MZB) is acknowl- edged, too. Further we are grateful to Ibu Mumpuni (MZB) for the permission to examine specimens under her care. In addi- tion, we are greatly indebted to Pak Henry, teacher in Lirung (Salibabu Island), who substantially supported our field work as well as to Sahid Kampi (Tahuna, Sangihe Island) and his broth- er in Lirung (Salibabu Island) for logistic support. We thank Rainer Gúnther (Berlin, Germany), Jakob Hallermann (Ham- burg, Germany), Maren Gaulke (Munich, Germany), and An- Bonner zoologische Beiträge 56 (2007) 127 dreas Schmitz (Geneva, Switzerland) for discussions about the identification of single specimens. Bernhard Huber (ZFMK), Pe- ter Jáger (Naturmuseum Senckenberg, Frankfurt, Germany) and Siegfried Ingrisch (Bad Karlshafen, Germany) kindly provided identifications of the arthropod remains of Varanus specimens. Kai Philipp (Baden-Baden, Germany) kindly provided the pho- tograph of a Talaud monitor lizard and Uwe Fritz, Edgar Lehr (both MTDK) and Eulalia Gasso-Miracle (RMHN) information about Sulawesi specimens in their collections. We also appre- ciate the shared information of Jimmy McGuire (Berkeley, USA) about the herpetofauna of Sulawesi. Finally, Jon Riley (Great Britain) and Ruud de Lang (Rotterdam, the Netherlands) kind- ly made available relevant literature sources about the Talaud Islands. Robert Neal (Brisbane, Australia), Mark Auliya (Selan- gor, Malaysia), and an anonymous reviewer made many help- ful comments on an earlier version of the present paper. The com- panies M & S Reptilien (VS Weigheim, Germany) and Peter Hoch (Waldkirch, Germany) provided equipment for the expe- dition. Financial support for the ongoing project about the her- petofaunal diversity, endemism, and biogeography of Sulawesi and adjacent islands was provided by a PhD scholarship (AK) from the Evangelisches Studienwerk e.V. Villigst. This is publication No. 7 of the co-operation project about the herpetofauna of the Sulawesian region between the herpetolog- ical sections of the Zoologisches Forschungsmuseum A. 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Koenig, Adenauerallee 160, 53113 Bonn, Germany; E-Mail: andrepascalkoch@web.de; Evy ARIDA, Museum Zoologicum Bogoriense, Jl. Raya Bogor km 46, 16911 Cibinong, Indonesia. Current address: Zoo- logisches Forschungsmuseum A. Koenig, Adenauerallee 160, 33113 Bonn, Germany; E-Mail: evyarıda@lycos.com; Awal RIYANTO, Museum Zoolog- icum Bogoriense, Jl. Raya Bogor km 46, 16911 Cibinong, Indonesia; E-Mail: awal_lizards@yahoo.com; Wolfgang BÖHME, Zool. Forschungsmuseum A. Koenig, Adenauer- allee 160, 53113 Bonn, Germany; E-Mail: w.boehme.zfmk@uni-bonn.de. Received: 13.08.2008 Revised: 15.01.2009 Accepted: 20.01.2009 Corresponding editor: R. van den Elzen AFRICA Africa calls Africa - Birds 8: Birding is the continent's premier birding journal, bringing you in-depth insights into birds and their behaviour with articles aimed at birdlovers of every level, from academics and dedicated twitchers to nature enthusiasts. 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/ + 20:00 / E p Fianarantsoa / + 21:00 € N EN isa A / + 22:00 N e + 23:00 q \ a + 24:00 We ~ 22 N Ñ NX ‘ > PN Bemaraha Ye 4 j | a ee wae 0 100 200km + 25:00 SS \ | \ J [__ ] Parks and reserves Collection localities: « Other localities 4 + Eliurus antsingy 7 * Eliurus carletoni Sd 2 E A o Eliurus cf.antsingy > a Eliurus danieli Fig. 1. Map of Madagascar showing localities mentioned in the text. The complete map of the island to the right shows the is- land wide delineation of limestone deposits based on Bésairie’s (1964) classification of the geology of Madagascar. These are ex- tracted from his categories 14 and 18 “Marnes & calcaires” [=marls and chalks]. The three circular areas delineated on the full map are presented in finer detail in the left hand circular insets and in which the distribution of different Eliurus spp. are given. 136 Steven M. GOODMAN et al.: A new species of Eliurus from the Réserve Spéciale d' Ankarana, northern Madagascar rithm. For ML analyses, we first identified the best-fit model of nucleotide substitution using the Akaike infor- mation criterion (AIC) as employed in ModelTest ver. 3.6 (POSADA & CRANDALL 1998). We subsequently evaluat- ed whether a molecular clock fit our data using a hierar- chical log-likelihood ratio test. Parameters for the result- ing best-fit model were fixed in a heuristic search using 10 replicates of random taxon addition and TBR branch swapping. Nodal support was calculated for both MP and ML analyses using non-parametric bootstrapping (FELSEN- STEIN 1985). All bootstrap analyses employed 1000 pseudoreplicates analyzed with heuristic searches as above. Polymorphism and divergence statistics were cal- culated using DNAsp ver. 4.10 (ROZAS et al. 2003); in ad- dition, we report ML-corrected divergence values as cal- culated in PAUP*. 3. RESULTS After comparisons of the specimens from the Réserve Spé- ciale (RS) d’Ankarana, previously identified as Eliurus antsingy, usmg external and craniodental measurements and characters, as well as a molecular analysis, these an- imals represent a previously unknown species of Mala- gasy rodent falling within the £. antsingy group (sensu CARLETON & GOODMAN 2007). This new taxon is de- scribed below. Superfamily Muroidea Illiger, 1811 Family Nesomyidae Major, 1897 Subfamily Nesomyinae Major, 1897 Eliurus carletoni, new species Figs. 2-3; Tables 1-3 Holotype. An adult female specimen in the Field Muse- um of Natural History (FMNH 173105) prepared as skin, skull, and partial skeleton, collected 9 April 2002 by Steven M. Goodman (original number SMG 12832). The round skin and skull are in fine condition. Recorded ex- ternal data include: TOTL, 335 mm; HBL, 143 mm; TL, 183 mm; HFL, 29 mm; EL, 25 mm; and WT, 99 gm. The basisphenoid suture is fused and third molar slightly worn. The animal was noted as having large mammae, no em- bryos or placental scars. The mammae formula was one axial pair and one inguinal pair. Muscle tissue samples were preserved in lysis buffer. Type locality. Madagascar, Province d'Antsiranana, Réserve Spéciale d'Ankarana, Campement des Anglais (Anilotra), 7.5 km NW Mahamasina, 12°54.4’S, 49°06.6°E; 125 m above sea-level (Fig. 1). Diagnosis. A species of Eliurus falling within the E. ants- ingy species group (sensu CARLETON & GOODMAN 2007) Fig. 2. Photograph of live Eliurus carletoni (FMNH 169718), a young adult male from the forest near the Andrafiabe Cave in the Reserve Spéciale d’Ankarana. The light colored ventral por- tion of the tail is slightly exaggerated by the photographic flas- hes. Photograph taken by Harald Schütz. characterized by a dark brown dorsal pelage, contrasting grayish-white venter, unicolor dark brown tail (including terminal tuft), and short hind foot (HFL 28-29 mm) and ears (EL 23-25 mm) (Fig. 2). Cranial size moderately large (ONL 40-41 mm), rostrum relatively short for the genus (LR/ONL ca. 31%), molar rows moderately long (LM1-3 5.4-5.7 mm), incisive foramina notably short and wide (LIF/LD ca. 47-49%), palatine process short and stout, and auditory bullae relatively small in comparison to other members of the E. antsingy species group. Giv- en present taxon and character sampling, the species is fur- ther diagnosed by the following 18 unreversed synapomor- phies in the cyt-b gene (the first nucleotide given is the ancestral state, followed by the nucleotide position in the cyt-b gene, followed by the derived state. Changes that result in an amino acid replacement are shown in bold; Bonner zoologische Beitráge 56 137 Fig. 3. Different views of Eliurus carletoni (holotype FMNH 173105): upper left-dorsal view of cranium, upper righi-ventral view of cranium, and lower-lateral view of cranium and mandible. Photograph by John Weinstein (FMNH Z94480_07d). 138 Steven M. GOODMAN et al.: A new species of Eliurus from the Reserve Spéciale d’Ankarana, northern Madagascar Table. 1. Summary statistics (mean + standard deviation, observed range, and sample size) for external morphological measure- ments of Eliurus antsingy from Bemaraha, E. antsingy from Namoroka, E. carletoni from Ankarana, E. danieli from Isalo, and E. majori from Montagne d'Ambre. See Materials and Methods for definitions of acronyms. Variable E. antsingy Bemaraha E. antsingy Namoroka 329.2 + 21.90 305-364, n=5 146.8 + 5.19 142-153, n=4 170.2 + 15.42 153-195, n=5 29.8 + 1.26 28-31, n=4 24.8 + 1.26 24-25, n=5 92.8 + 6.55 87-101, n=4 354 HBL 160 TL 186 HFL 32 EL 26 WT 131 all other changes are silent): C189T, C201A, C270T, C312T, C321T,C322T, C585A,;T598C, T678C, T710C, C722T, T768C, T837C, T849C, C879T, A945C, A978G, C1069T. Referred specimens. Other specimens of Eliurus carletoni have been examined in this study, all from the RS d’Ankarana and collected by S. M. Goodman (except as noted). The following three specimens were obtained from the type locality: FMNH 173104 (SMG 12728), a young adult female, with partially fused basisphenoid suture, on 6 April 2002; FMNH 173106 (SMG 12833), a young adult male, with abdominal testes, with unfused basisphenoid suture, on 9 April 2002; FMNH 173109 (SMG 12848), an adult female, with largely fused basisphenoid suture, on 11 April 2002. An additional three specimens were tak- en from 2.6 km E Andrafiabe, in forest near Andrafiabe Cave, 12°55.9’°S, 49°03.4’E; ca. 50 m above sea-level: FMNH 169718 (SMG 11928), a young adult male, with unfused basisphenoid suture, obtained on 22 January 2001; FMNH 169719 (SMG 11946), a young adult female, with unfused basisphenoid suture, on 24 January 2001; FMNH 169720 (SMG 11994), an adult scrotal male, with fused basisphenoid suture, on 26 January 2001. Two addition- al specimens were collected by Achille P. Raselimanana near the trail junction of routes to Matsaborimanga, E. danieli E. carletoni E. majori Ankarana Isalo Montagne d’Ambre 328.3. = 9.07 335,837 336.0 + 14.62 318-335, 322-365, n=3 n=7 147.7 + 4.04 150; 152 146.6 + 4.18 143-150, 141-154, n=3 n=6 174.3 + 9.61 179; 195 179.1 + 13.66 164-183, 165-208, n=3 n=7 28.8 + 0.50 30,32 2891.51 28-29, 2731, n=4 n=8 24.3 + 0.96 26, 28 19.8 + 1.49 23-25, 17-21, n=4 n=8 94.8 + 4.72 91, 100 19 = 558 88-99, 70-89, n=8 n=4 Campement des Anglais, and Andafiabe, 10.0 km NW Ma- hamasina, ca. 12°53.2’S, 49°06.6’E; 100 m above sea-lev- el: FMNH 173107 (SMG 12846), an adult female, with largely fused basisphenoid suture, on 11 April 2002; FMNH 173108 (SMG 12847), a young adult male, with unfused basisphenoid suture, on 11 April 2002. In addi- tion, a number of incomplete cranial specimens referable to E. carletoni have been recovered pellets of the Mada- gascar Red Owl (Tyto soumagnei) collected at the RS d’Ankarana, Perte des Rivieres (12°57.283’S, 49°7.627’E). Etymology. This new species is named in honor of Dr. Michael D. Carleton of the National Museum of Natural History, Smithsonian Institution, Washington, D.C. in recognition of his immense contributions to the field of rodent systematics. In particular, Dr. Carleton has worked extensively on the systematics and morphological evolu- tion of Madagascar’s native rodents. Distribution. This species is currently only known from the RS d'Ankarana in the extreme northern portion of Madagascar (Fig. 1). Specimens obtained in the RS d’Analamerana and the Réserve Forestiére de Loky-Man- ambato (Daraina region) may be referable to this taxon; future studies will address this question. Bonner zoologische Beiträge 56 139 Fig. 4. Comparison of incisive foramen and molar toothrows — (right) in the holotype of Eliurus carletoni (FMNH 173105) from the Reserve Spéciale d’Ankarana and (left) £. antsingy from the Parc National de Bemaraha (FMNH 172271). Note that this struc- ture is reduced in length and width in the former species, as well as the palatine process, which is thick and short. Photograph by John Weinstein (FMNH Z94480_08d). Description. Overall body hair texture soft and relative- ly fine. Dorsum cover hairs 7-9 mm in length over mid- dle rump and 5-6 mm at the level of the nape. Cover hairs of dorsum largely bi-colored with proximal two-thirds dark brown and distal one-third medium buff, although some hairs tipped with faint dusky brown. Guard hairs dark brown to blackish. The overall coloration of the up- perparts, including forehead and face, is a dark agouti with a diffused mid-dorsal line that is notably darker than the surrounding fur. In several individuals, this band broad- ens towards the rump. Fur coloration of lower legs ap- proaches a lighter tannish-brown. In a few individuals (FMNH 173107, 173108), the dorsal pelage coloration is slightly lighter, with the forehead, face, and lower legs be- ing tannish-brown. Along mid-portion of body, the limit between the dorsal and ventral pelage is relatively well demarcated. The coloration of the ventrum is variable and ranges from being nearly entirely white (FMNH 169719, 173107, 173108), being largely white with a diffusion of gray towards the throat and base of limbs (FMNH 173109), to being broadly diffused with light gray and having dusky brown at the base of the limbs (as in holo- type FMNH 173105). A diffuse dusky-brown band extends over the mid-portion of the dorsal tarsus surface and prox- 140 Steven M. GOODMAN et al.: A new species of Eliurus from the Réserve Spéciale d’Ankarana, northern Madagascar Table. 2. Summary statistics (mean + standard deviation, observed range, and sample size) for craniodental measurements of adult Eliurus antsingy from Bemaraha, E. antsingy from Namoroka, E. carletoni from Ankarana, E. danieli from Isalo, and E. majori from Montagne d'Ambre. See Materials and methods for explanation of acronyms and age classes. Measurements for two of the E. antsingy specimens (MNHN 1966.2220, 1966.2222) were taken from CARLETON & GOODMAN (2007). Variable E. antsingy E. antsingy E. carletoni E. danieli E. majori Bemaraha Namoroka Ankarana Isalo Montagne d’Ambre ONL 43.2 +1.14 41.0 + 1.40 40.2 + 0.48 39.1, 40.6 36.9 + 0.68 41.9-44.1, 39.3-42.5, 39.5-40.6, 36.1-38.0, n=3 n=5 n=4 n=7 ZB 21.2=#0;51 19.320,62 19.7 +0.31 18.9, 19.1 18.8 + 0.42 20.8-21.8, 18.4-20.3, 19.4-20.0, 18.3-19.4, n=3 n=6 n=3 n=7 BBC 16,1 0.26 15,1 # 0.20 15.1 20,25 14.6, 15.1 14.4 # 0.18 15.8-16.3, 14.9-15.4, 14.8-15.4, 14.1-14.7, n=3 n=6 n=4 n=8 IOB 6.1 0.15 5.9 + 0.14 5.6 + 0.08 3.3298 9.225.021 5.9-6.2, 5.7-6.1, 5.5-5.7, 4.9-5.5, n=3 n=6 n=4 n=8 LR 14.9 + 0.21 13.2:+:0.55 12.6 + 0.24 122,133 1128251037 14.7-15.1, 12.7-14.1, 12.2-12.7, 11.5-12.6, n=3 n=5 n=4 n=7 BR 7.4 + 0.06 6.8 + 0.23 7.0+0.17 7.0, 7.4 6.9 + 0.07 7.4-71.5, 6.4-7.0, 6.7-7.1, 6.8-7.2, n=3 n=6 n=4 n=8 PPL 15.2 + 0.61 14.3 + 0.61 14.3: 0.25 13.8, 13.8 12.8 + 0.60 14.7-15.9, 13.7-15.0, 14.0-14.6, 12.0-13.6, n=3 n=6 n=4 n=8 LBP 7.8 + 0.10 8.1 + 1.06 7.6+0.21 1:15.78 7.4 + 0.43 71-79, 7.3-10.2, 7.3-7.9, 6.8-8.0, n=3 n=6 n=4 n=8 PPB 6.1 + 0.06 5.520.413 5520.22 Dd 050.15 6.0-6.1, 5.3-5.6, 5.2-5.7, 5.4-5.8, n=3 n=6 n=4 n=8 LIF 6.7 + 0.10 5.7 BOS 2 Se 022 5.3, 5.4 5.1.=10.32 6.6-6.8, 4.7-6.0, 4.9-5.4, 4.6-5.6, n=3 n=6 n=4 n=8 BIF 2.9 +0.21 2.4 + 0.22 2.6 0.10 2.1521 2.2.0214 2.7-3.1, 2.0-2.6 2.42.6, 2.0-2.5, n=3 n=6 n=4 n=8 LD 11.8 + 0.56 10.9 + 0.89 10.7 + 0.24 9.8, 10.1 9.8 + 0.21 Bonner zoologische Beitráge 56 141 Table. 2. (pursuit). Variable E. antsingy E. antsingy Bemaraha Namoroka 11.3-12.4, 10.1-12.5, n=3 n=6 BMils 8.2+0.15 7.5+0.14 8.0-8.3, 1377, n=3 n=6 DAB 6.3 + 0.26 57+0.21 6.0-6.5, 5.5-6.0, n=3 n=6 BZP SS 3.9+ 0.34 3.2-4.3, 3.54.5, n=3 n=6 BOC 9.6+0.10 8.8 + 0.20 9.5-9.7, 8.5-9.1, n=3 n=6 LMI-3 5.8+0.15 5.6 + 0.23 5.7-6.0, 5.339; n=3 n=6 WMI 1.6 + 0.10 1.6 + 0.08 ¡PSA 1.4-1.6, n=3 n=6 E. carletoni E. danieli E. majori Ankarana Isalo Montagne d’Ambre 10.4-10.9, 9.4-10.0, n=4 n=7 755026 Eds AS 1323025 7.3-1,8, 7.0--7.7, n=4 n=7 5.6 20.17 5.6, 5.8 5.3 0,18 5.45.8, 5.0-5.5, n=4 n=8 3.650,15 3.8; 3.8 3.2+0.18 3.4-3.7, 3.0-3.5, n=4 n=8 8.9 + 0.17 8.6, 8.7 8.3 + 0.20 8.7-9.1, 8.1-8.6, n=4 n=8 5.6 +0.14 o 3920.20 543.7, 5.6-6.3, n=4 n=8 1.6 + 0.06 1:6, 1.7 1.7 20.07 1.5-1;6; 1.6-1.8, n=4 n=8 imal metatarsus; the balance of these structures is domi- nated by light-colored fur intermixed with light brown. Relatively well-developed white ungual tufts at base of claws. Caudal tail tuft prominent over distal half of length, tuft hairs becoming gradually longer toward the tip and meas- uring about 10-13 mm long near the terminal portion. Tail fur notably mono-colored, dark blackish-brown along complete length. The only exception is FMNH 173108, which has a 7 mm wide largely white-haired section start- ing about 30 mm from the terminal tip. Proximal half of tail covered with short black hairs that sparsely cover the caudal scales. Ventral tail scales distinctly rectangular- shaped and on the dorsal surface dark brownish and on the ventral surface light gray at base merging to dark brownish distally. One animal with a short bobbed tail (FMNH 169720) has a distorted dark blackish-brown ter- minal tuft, but there is no evidence of white replacement hair. Hind feet short and broad; absolutely and relatively small- er than in £. antsingy from Bemaraha (FMNH 172721). Plantar pads six and arranged as characteristic of the genus (CARLETON 1994, his fig. 2). Pinnae relatively short and not as long as E. antsingy from Bemaraha (Table 1); col- or dark brown externally and sparsely clothed with fine light-colored hairs. Cranium the size of other moderately large members of the genus (ONL = 39.5-40.6 mm), but more diminutive than E. antsingy from Bemaraha (ONL = 41.9-44.1 mm) and smaller on average than £. antsingy (ONL = 39.3-42.5 mm) from Namoroka. Similar in overall pro- file shape to E. antsingy, but skull less sharply arched to- wards occiput (Fig. 3). Rostrum proportionately shorter (LR/ONL = 30.9-31.3%), as compared to E. antsingy from Bemaraha (LR/ONL = 34.9%) and all other members of the genus. Braincase not notably rounded as in, for exam- ple, E. danieli or other members of the E. majori group. Subsquamosal fenestra notably reduced, similar to E. ants- 142 Steven M. GOODMAN et al.: A new species of Eliurus from the Réserve Spéciale d’Ankarana, northern Madagascar Table. 3. Factor loadings from principal component analyses of log transformed craniodental measurements of specimens of adult Eliurus antsingy (Bemaraha and Namoroka), E. carleto- ni, E. danieli, and E. majori (Montagne d’Ambre). For this ana- lysis, a graphical representation of the first two factors is pre- sented in Fig. 5. On the basis of ANOVA analyses, two cranio- dental variables, LBP and WMI, showed no significant diffe- rences across taxa and were not included in the PCA analysis. Factor 2 Factor 1 Factor 3 ONL -0.944 0.254 0.018 ZB -0.869 -0.002 -0.048 BBC -0.939 0.027 0.114 IOB -0.883 0.215 0.172 ER -0.944 -0.005 -0.186 BR -0.457 -0.543 -0.420 PPL -0.901 0.211 0.119 PPB -0.602 -0.639 0.0343 LIF -0.730 -0.472 0.197 BIF -0.639 -0.421 0.386 LD -0.831 0.386 -0.214 BMI -0.618 -0.617 -0.032 DAB -0.931 0.071 -0.083 BZP -0.545 0.447 -0.439 BOC -0.923 -0.125 0.247 LMI-3 0.108 -0.847 -0.254 Eigenvalues 9.842 3.519 1.554 Proportion of total variation 54.7% 74.2% 82.9% explained ingy from Bemaraha, without exposing the interior brain- case. Hamular process of the squamosal reduced and stout, similar to that in E. antsingy from Bemaraha. Zygomatic arches not notably heavy, although the zygomatic plates are proportionately stouter as compared to other members ofthe genus of sımilar ONL length. Nasolacrımal capsule more inflated and nasolacrımal foramen closer to zygo- matic notch than in £. antsingy from Bemaraha. Incisive foramina bluntly pointed on their posterior and anterior ends; notably short (LIF/LD = 47.1-49.5%) for members of the genus; in £. antsingy from Bemaraha the LIF/LD = 56% (Fig. 4). Posterior palatine foramina ob- long ovals, situated within the maxillary-palatine suture at the level ofthe M1-M2. Palatine process stout and short. Supernumerary palatal foramina occurring in the palatine bones, but minute in size and irregularly formed: in the holotype these occur on the animal's left side at the lev- el of M2-M3 and on the right side in the central portion of M3. No posterolateral palatal foramen. Posterior mar- gin of the bony palate terminates at the posterior ends of the third molars. The anterior portion of the mesoptery- goid fossa is broad, U-shaped, and extending distally with- out a constriction in the central winged portions of the pterygoid process. Auditory bullae relatively small for the genus; from ventral view the ectotympanic covering on- ly a portion of the petrosal part of auditory bullae and a small portion of the periotic. Anterior enamel surface of upper and lower incisors dull yellowish-orange. Alveolus of lower incisor terminating at the level of the coronoid process and below the sigmoid notch. The base of the incisor root forming slight rise on mandibular ramus but not a capsular process. Toothrows not particularly elongated (LM1-3 = 5.4-5.7 mm) and rel- ative to cranial size, in proportion to those of E. myoxi- nus and E. tanala Major, 1896 (CARLETON 1994, Appen- dix 2). Molars robust with trilaminar configuration on pla- nar surface, typical of other members of this genus; up- per and lower first and second molars about equal in length and both notably longer than third molars. Morphological comparisons. On the basis of numerous morphological traits, £. carletoni is placed along with E. antsingy from Namoroka and Bemaraha in the same species assemblage (CARLETON & GOODMAN, 2007). Giv- en the morphological uniqueness of this species complex, as outline by CARLETON et al. (2001) and CARLETON & GOODMAN (2007), our comparisons here are largely con- fined to other populations within the group. The northern population of E. majori, occurring on Montagne d’Am- bre, is also included, because this is geographically the closest known moderately sized Eliurus. Further compar- isons are also made to £. danieli from transitional dry/hu- mid forests of the PN de I’Isalo in the central west. Both E. majori and E. danieli are included within the £. ma- jori group, which CARLETON & GOODMAN (2007) allied with the E. antsingy group (see below). In general, the dorsal pelage coloration of E. carletoni is similar to E. antsingy; however, the dorsum cover hairs towards the middle rump tend to be slightly longer in £. antsingy (9-11 mm) as compared to E. carletoni (7-9 mm). Specimens of E. majori from Montagne d’Ambre have distinctly dark slate gray dorsums and a denser and more svelte dorsal pelage and those of E. danieli have bright plumbeous gray cover hairs. The ventral pelage of E. carletoni varies from only being slightly tinged with white in FMNH 173105 (the holotype), to being largely white-bellied. However, in all cases E. carletoni has more Bonner zoologische Beitráge 56 143 whitish-colored fur on the belly than £. antsingy from Be- maraha (FMNH 172721), whereas all eight of the skin preparations of E. antsingy from Namoroka have white venters. Both E. carletoni and E. antsingy have mono-col- ored tails, generally with dark-blackish brown fur along the complete length, and the caudal tuft occupies the dis- tal half of the tail. Some individuals of E. antsingy from Namoroka have notably light brown tails and reduced cau- dal tufts; these include both young adults (FMNH 178591, 178593) and full adults (FMNH 178592) and this difference does not appear to be age related. Amongst the specimens of E. carletoni there is one anomalous case (FMNH 169720), which has a small white band along its length. Other species of E/iurus with largely mono-col- ored dark-blackish brown tails include £. myoxinus and E. webbi Ellerman, 1949, which can be distinguished from E. carletoni by the tail length and pilosity, as well as nu- merous craniodental characters (CARLETON 1994, 2003; CARLETON et al. 2001). Eliurus carletoni is distinctly smaller and lighter than £. antsingy from Bemaraha (FMNH 172271), but overlaps in all external measurements with specimens of E. ants- ingy from Namoroka (Table 1). The cranium of E. car- letoni (ONL = 39.5-40.6 mm) approximates the size of E. carletoni holotype PE 2 -2.0 -1.0 0.0 PC 1 other moderately large members of the genus (CARLETON 1994), including E. danieli and E. majori, but it is notably smaller than £. antsingy from Bemaraha (ONL = 41.9-44.1 mm) and smaller on average than E. antsingy (ONL = 39.2-42.5 mm) from Namoroka (Table 2). The rostrum is proportionately shorter in £. carletoni (LR/ONL = 30.9-31.3%) as compared to E. antsingy from Bemara- ha (LR/ONL = 34.2-35.1%) and from Namoroka (LR/ONL = 32.3-33.2%), as well as most other members of the genus (calculated from descriptive statistics in CAR- LETON, 1994, Appendix 2): E. minor (LR/ONL = 33.3%), E. myoxinus (LR/ONL = 34.1%), E. majori (LR/ONL = 34.4%), E. tanala (LR/ONL = 36.0%), E. webbi (LR/ONL = 35.3%), and E. petteri Carleton, 1994 (LR/ONL = 35.3%). The hamular process of the squamosal notably reduced within the £. antsingy group; in contrast to E. danieli and E. majori (from Montagne d’ Ambre) in which the hamu- lar process 1s elongated and associated with a more promi- nent subsquamosal foramen. Eliurus carletoni has a no- tably short (LIF/LD = 47.1-49.5%) and bluntly pointed incisive foramen, as compared to E. antsingy from Be- maraha (LIF/LD = 56%; Fig. 4). The palatine process in E. carletoni is generally thick and short, particularly in A E. carletoni @ E antsingy (Namoroka) O E. antsingy (Bemaraha) A E. majori (Montagne d'Ambre) mM ECE. daniel 1.0 2.0 Fig. 5. Projections of factor 1 (x-axis) and factor 2 (y-axis) in principal component analysis on 16 In-transformed craniodental variables of Eliurus spp. Loadings of variables on each axis are shown in Table 3. 144 - Ankarana 173106 © 0.02 Ankarana 169718 © substitutions/site Ankarana 169719 © m Ankarana 173107 © | 8 = Ankarana 173109 0 S = _ 190 | Ankarana 173108 © | = 100 Ankarana 173104 Ankarana 173105 100 Bu ng [| 100 5.8% [ens 175913 + Namoroka 175909 " | M Ni | a 700 Namoroka 175910 E | 3 10) — 100 | Namoroka 175912 = | > 97 14.6% Q Bemaraha 172721 w a — E. danieli UADBA 10483 100 100 _——— E majori 154610 100 100 E. majori 178686 Steven M. GOODMAN et al.: A new species of Eliurus from the Réserve Spéciale d'Ankarana, northern Madagascar 10 “AE 0 oo km o 100 -12 Ankarana o -14 6 Namoroka 8 -18 Bemaraha ” 20 —— = > 42 44 45 48 50 52 Fig. 6. The maximum-likelihood phylogeny inferred from analysis under the best-fit model of nucleotide substitution for the cyt- b data (GTR + I, no clock; -InL = 3137.31). The tree is rooted with sequences of Eliurus danieli and two E. majori. The ingroup consists of specimens referred to E. antsingy or E. carletoni. Terminal branches within the ingroup are labeled with collecting lo- calities shown in the map (right), followed by the museum (FMNH, unless otherwise noted) voucher number. Numbers above and below branches refer to maximum-likelihood and parsimony bootstrap support, respectively. The divergence between populations (ML-corrected) is shown next to circles at each interpopulation node. comparison to the narrow and relatively long structure in E. antsingy and members of the E. majori group. The pos- terior palatine foramina are consistently the same size and are positioned within the maxillary-palatine suture at the level of the M1-M2 for all members of the E. majori and E. antsingy groups. The external portion of the auditory bullae of E. carletoni is notably smaller than in £. ants- ingy from Bemaraha (DAB 5.4—5.8 mm vs. 6.0-6.5 mm), whereas anımals from Namoroka largely overlap E. car- letoni in this measurement (DAB 5.5—6.0 mm). In all specimens of E. antsingy and E. carletoni, the mo- lars have a robust trilaminar configuration and the ante- rior enamel surface of upper and lower incisors are dull yellowish-orange. The position of the lower incisor root with respect to the coronoid process is likewise consis- tent across members of this group. The toothrows in £. carletoni are shorter than E. antsingy from Bemaraha (LM 1-3 = 5.4-5.7 mm vs. 5.7-6.0 mm), but overlap with those in specimens from Namoroka (LMI-3 = 5.3-5.9 mm). Members of the E. antsingv group have proportion- ately shorter molar rows relative to cranial size than mem- bers of the E. majori group: E. carletoni LM1-3/0NL = 13.7-14.0%, E. antsingy from Bemaraha LM1-3/ONL = 13.6%, E. danieli LM1-3/ONL = 15.1-15.3%, and E. ma- jori from Montagne d’Ambre LM1-3/ONL = 15.5-16.6%. Members of the E. antsingy species assemblage have the upper and lower third molars subequal in length to the first and second molars. The same condition is found in £. danieli, while the three molars are largely the same size in E. majori from Montagne d'Ambre. Principal Component Analysis of craniodental measure- ments (Fig. 5) shows complete separation of E. antsingy taken in Bemaraha from E. carletoni and E. majori from Montagne d’Ambre; although there is broad overlap be- tween E. carletoni and E. antsingy from Namoroka. The factor loadings for this analyses (Table 3) indicates that the majority of variables for factor 1 and coronal length of maxillary toothrow (LM 1-3) for factor 2 showed heavy negative loadings. Factor 1 explained 54.7% of the total variation, factor 2 an additional 19.5%, and factor 3 an- Bonner zoologische Beiträge 56 dsyeusy I DEV gell 195-USY Cc DEV rcell Nod I IL 2601 JUSTIS € Le) COL AUL+-EIV I VD 8LOl JUSTTS € IL S601 JUSJIS € Ie L 1€8 JUSTIS € Le) 8901 jUSTIS I Ie) VCL JUSTIS € Leo 666 JUSTIS € Le) L89 JUSTIS € Le) 0€9 JUSTIS € Le ps9 JUETIS € Le) 1c9 JUSTIS € Le) $85 JUSTIS € DEV c8s JUSTIS € Le) 875 JUSTIS € DEV 68? 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Whereas it is the case that E. carletoni and E. antsingy from Namoroka showed broad morphologi- cal overlap in the PCA, based on the molecular phyloge- ny (see next section) these two populations are not sister taxa within the £. antsingy species group. Molecular variation and phylogeny. There were 6 unique cyt-b haplotypes among the 8 specimens from Ankarana, and nucleotide diversity (7) among them was 0.003. Each of the specimens from the Namoroka population had a unique haplotype, and nucleotide diversity within this pop- ulation was also 0.003. Average sequence divergence (cor- rected for within-population polymorphism; NEI 1987, eq. 10-21) was 4.4% (ML-corrected divergence = 4.6%) be- tween the Namoroka and Bemaraha populations and 4.9% (5.8% ML-corrected) between Ankarana and the two southern populations. Phylogenetic analysis of the cyt-b dataset using parsimo- ny resulted in three minimum-length trees (L = 350; CI = 0.80; RI = 0.86). The strict-consensus of these trees is less resolved than but entirely consistent with the topology re- sulting from a maximum-likelihood analysis of this dataset under its best-fit model (GTR+I, no clock; Fig. 6). Two well-supported clades are apparent within E. antsingy when this tree is rooted with £. danieli and two individ- uals of E. majori. The first includes the specimens from Ankarana, the second comprises specimens from Namoro- ka and Bemaraha. When character state changes are op- timized on this phylogeny, 18 unreversed synapomorphies can be mapped to the branch subtending the Ankarana specimens (E. carletoni); two of these result in amino acid substitutions. An additional 16 unreversed synapomor- phies (all silent changes) can be mapped to the branch sub- tending E. antsingy from Namoroka and Bemaraha (Table 4). Ecological notes. All of the specimens of E. carletoni used in this study were collected in the forests of the Ankarana Massif, specifically the RS d’Ankarana in northern Madagascar (Fig. 1). This region lies on a block of karstic limestone that was formed during the Jurassic and subse- quently uplifted (CARDIFF & BEFOUROUACK 2003). The weathering of the exposed limestone surface has formed the pinnacle karst known in Malagasy as tsingy, which is characteristic of the Ankarana Massif, as well as the 1s- ingy habitat at Bemaraha and Namoroka. The zone surrounding the Ankarana Massif receives slight- ly less than 2000 mm of rainfall annually, of which most falls between the months of December to April, resulting in a prominent seven-month dry season (FOWLER et al. 1989; HAWKINS et al. 1990). The deep canyons, which pro- vide protection from the sun and desiccating wind, and un- derground streams passing close to or resurging at ground level, result in relatively mesic forest conditions within these formations. It is in such areas that E. carletoni has been found during biological inventories. On the basis of trap captures set both on the ground and along lianas, E. carletoni can be characterized as a pri- marily scansorial forest-dwelling inhabitant of the dry de- ciduous habitats of the Ankarana Massif. It has been trapped in pristine and disturbed forests, generally asso- ciated with tsingy rock formations, and on the ground, branches and vines. Some notes associated with captured animals help to characterize the different microhabitats it occupies: “...at base of tsingy. Trap on ground between rocks and just above water source” (FMNH 169718); “at base of tsingy. Trap 1.75 m off ground on 5 cm diam[eter] liana arching over ground to mid-canopy” (FMNH 169719); “Trap on ground in narrow passage between two rocks — resting on tsingy” (FMNH 169720); “Trap on ground along large fallen and rotten tree trunk next to 1s- ingy wall” (FMNH 173104); “Trap about 1 m off the ground on 20 cm diam[eter] fallen tree trunk” (FMNH 173106); and “In disturbed deciduous forest. Not within 500 m of tsingy formation” (FMNH 173107, 173108). On the basis of rapid inventory information it is difficult to infer many details on the reproductive behavior and sea- sonality of E. carletoni, but a few observations can be pre- sented. Individuals collected in late January 2001 includ- ed a young adult male and a female (FMNH 169718, 169719) that were not in reproductive condition, as well as an adult male (FMNH 169720) with large scrotal testes and convoluted epididymides. A series of animals captured during early April 2002 included one adult male that was not in reproductive condition (FMNH 173106); three fe- males each of which had large mammae but no embryos, and one of which had discernable placental scars (one in each the left and right uterine horns) (FMNH 173104, 173107, 173109); and a male with partially scrotal testes, (measuring 5 x 5 mm) and slightly convoluted epi- didymides (FMNH 173108). A recent study on the dietary habits of the owl Tyro sou- magnei within the RS d’Ankarana, found that this raptor feeds extensively on E. carletoni (which was identified as E. antsingy in that study). Across several different seasons and years, E. carletoni comprised 22.1% of the minimum number of individuals and 49.8% of the biomass con- sumed by this owl (CARDIFF & GOODMAN 2008). At Ankarana, other than the bat fauna (14 species; GOODMAN et al. 2005), the diversity of mammals weighing less than 100 gm is rather limited. No other species of native ro- dent has been captured at this site, but two species of in- troduced rodent (Rattus rattus [Linnaeus, 1758] and Mus musculus Linnaeus, 1758) occur here. In addition, a shrew (Suncus madagascariensis [Coquerel, 1848]), a small Bonner zoologische Beitráge 56 147 shrew-tenrec (Microgale brevicaudata G. Grandidier, 1899), and a mouse lemur (Microcebus tavaratra Ra- soloarison, Goodman & Ganzhorn, 2000) are known from the reserve. Previous reports of E. myoxinus ın Ankarana (NICOLL & LANGRAND 1989) have not been substantiated during recent biological inventories (CARLETON et al. 2001; GOODMAN, unpublished data). 4. DISCUSSION CARLETON & GOODMAN (2007) examined morphological variation in described members of the genus Eliurus and divided these animals into five different groups. Two of the species assemblages important for the current discus- sion include the Eliurus majori group (comprising E. ma- jori, E. penicillatus Thomas, 1908, and E. danieli) and the Eliurus antsingy group (comprising E. antsingy, and the newly described E. carletoni presented herein). On the ba- sis Of several external morphological and craniodental characteristics, these authors suggested that these two species groups might be more closely related to each oth- er than either is to any other species of Eliurus. This hy- pothesis remains to be critically tested with molecular da- ta, but preliminary results based on mitochondrial cyt-b data (JANSA, unpublished data) lend credence to this sug- gestion. As currently understood, members of the E. antsingy group have a broad but disjunct geographic distribution across northern Madagascar, and occur in zones of low- land deciduous forest resting on limestone outcrops (Fig. 1). The type locality of E. antsingy is the Bemaraha Plateau in central west Madagascar, a limestone forma- tion dating from the mid-Jurassic. In their description of this taxon, CARLETON et al. (2001) tentatively assigned Eli- urus specimens from the Namoroka Massif — another 1s- ingy limestone area 300 km further to the north — to £. antsingy. However, they noted that the animals from Namoroka were consistently smaller and had whiter ven- tral pelage than those from Bemaraha; these differences are upheld in the larger sample from Namoroka present- ed herein. Based on our molecular phylogeny of the E. antsingy species group (Fig. 6), the Namoroka animals are the sister taxon of the animal from Bemaraha, but the two populations differ by an average 0f 4.7% (ML-corrected) sequence divergence, and there are several fixed changes in the cyt-b gene that uniquely characterize each (Table 4). Therefore, these two populations could be considered two distinct species, or they may simply represent ex- tremes of variation in a contiguous population. Our Be- maraha samples of E. antsingy come from the southern end of this approximately 100 km limestone formation and additional material from the unsampled ¿singy habitat in the northern portion of this formation could harbor pop- ulations that will be critical for assessing the species sta- tus of the Bemaraha and Namoroka populations, which are here conservatively retain as the single species E. ants- ingy. The Eliurus specimens from the Ankarana Massif — de- scribed here as E. carletoni — as well as a large-bodied Eli- urus from the Forét d’ Ambilondambo (Daraina) were col- lected after CARLETON et al. (2001) published their descrip- tion of E. antsingy. CARLETON & GOODMAN (2007) ten- tatively assigned these specimens to E. antsingy based on morphometric comparisons, but they called for addition- al collections from intermediate localities as well as ge- netic data to test this conclusion. Although we lack criti- cal specimens from new locales, the application of mo- lecular data to existing specimens clearly shows that the Ankarana population is reciprocally monophyletic relative to E. antsingy from Namoroka and Bemaraha, and that the two clades differ by 5.8% (ML-corrected) sequence diver- gence. In addition, the two species are diagnosable by sev- eral fixed changes in the cyt-b gene (Table 4). Addition- al collections from limestone areas — such as Ankara and Kelifely — between Namoroka and the northern portion of the island will help to resolve whether these two popula- tions are genetically isolated. However, the lack of gene flow and high degree of molecular divergence between them apparent in this study, leads us to recognize the Ankarana population as a new species, E. carletoni. We note that specimens from Daraina are morphologically very similar to E. carletoni and may be assignable to this species as well. Ongoing morphological and molecular studies will help resolve this as well as other pressing questions in Eliurus systematics. Acknowledgements. For access to specimen in their care, we very much appreciate the helpfulness of the following museum staff: P. D. Jenkins (BMNH); C. Denys (MNHN); and D. Rako- tondravony and V. Soarimalala (UADBA). Field surveys in lime- stone areas of western and northern Madagascar were financed by grants from the Volkswagen Foundation and the National Ge- ographic Society (6637-99 and 7402-03). Permission to conduct the fieldwork was granted by the Association Nationale pour la Gestion des Aires Protégées and the Direction des Eaux et Forets. We are most grateful to Lucienne Wilmé for producing the map in Fig. 1 and Harald Schütz for making available the photograph of the living animal shown in Fig. 2. Molecular work was fund- ed in part by funds from the University of Minnesota. We are grateful to V. Soarimalala for comments on an earlier version of this manuscript. Zusammenfassung. Die Nager-Art Eliurus antsingy (Nesomy- inae) ist aktuell aus einigen disjunkten Kalkstein-Regionen in trockenen Tiefland-Waldgebieten im nórdlichen und westlichen Madagaskar bekannt. Frúhere Untersuchungen haben morphol- ogische Unterschiede zwischen Individuen aus verschiedenen isolierten Populationen festgestellt, sahen aber alle als Ange- hórige derselben Art an. In der vorliegenden Untersuchung úber- 148 Steven M. GOODMAN et al.: A new species of Eliurus from the Réserve Spéciale d' Ankarana, northern Madagascar prúfen wir die morphologische und genetische Variabilitát in- nerhalb von und zwischen Populationen von E. antsingy, mit Schwerpunkt auf dem Status der Population der Reserve Spé- ciale d’Ankarana im äußersten Norden der Insel. Während eine Unterscheidung der Population in Ankarana von der ihr nächst vorkommenden -500 km südlich (Namoroka) nicht móglich ist, unterschieden sich beide Populationen in morphologischen Merkmalen von der der Typuslokalität (Bemaraha) der Art. Im Gegensatz dazu bestehen zwischen allen drei Populationen deut- liche genetische Unterschiede. Die phylogenetische Analyse dieser Daten deutet darauf hin, dass die beiden morphologisch ähnlichen Formen keine Schwester-Taxa sind. Auf der Grund- lage dieser Ergebnisse sehen wir die Tiere der Reserve Spéciale d’Ankarana als neue Art an, fassen aber diejenigen aus den bei- den anderen untersuchten Populationen wegen nicht ausreichend großer Stichproben vorläufig weiterhin als E. antsingy auf. REFERENCES BESAIRIE, H. 1964. Carte Geologique de Madagascar, au 1:1,000,000e, trois feuilles en couleur. Service Géologique, Antananarivo. CARDIFF, S. G. & BEFOUROUACK, J. 2003. The Réserve Spéciale d'Ankarana. Pp. 1501-1507 in: GOODMAN, S. M. & BENSTEAD, J. P. (eds.) The natural history of Madagascar. The Universi- ty of Chicago Press, Chicago. CARDIFF, S. G. & GOODMAN, S. M. 2008. Natural history of red owls (Tyto soumagnei) in dry deciduous tropical forest in Madagascar. The Wilson Journal of Ornithology 20: 892-898. CARLETON, M. D. 1994. Systematic studies of Madagascar’s en- demic rodents (Muroidea: Nesomyinae): revision of the genus Eliurus. American Museum Novitates 3087: 1-55. CARLETON, M. D. 2003. Eliurus, tufted-tailed rats. Pp. 1373-1380 in: GOODMAN, S. M. & BENSTEAD, J. P. (eds.) The natural history of Madagascar. The University of Chicago Press, Chicago. CARLETON, M. D. & GOODMAN, S. M. 2007. A new species of the Eliurus majori complex (Rodentia: Muroidea: Nesomyi- dae) from south-central Madagascar, with remarks on emer- gent species groupings in the genus E/iurus. American Mu- seum Novitates 3547: 1-21. CARLETON, M. D., GOODMAN, S. M. & RAKOTONDRAVONY, D. 2001. A new species of tufted-tailed rat, genus Eliurus (Muri- dae: Nesomyinae), from western Madagascar, with notes on the distribution of E. myoxinus. Proceedings of the Biologi- cal Society of Washington 114: 972-987. FELSENSTEIN, J. 1985. Confidence limits on phylogenies: an ap- proach using the bootstrap. Evolution 39: 783-791. FOWLER, S. V., CHAPMAN, P., CHECKLEY, D., HURD, S., MCHALE, M., RAMANGASON, G.-S., RANDRIAMASY, J.-E., STEWART, P., WALTERS, R. & WILSON, J. M. 1989. Survey and management proposals for a tropical deciduous forest reserve at Ankarana in northern Madagascar. Biological Conservation 47: 297313. GOODMAN, S. M., ANDRIAFIDISON, D., ANDRIANAIVOARIVELO, R., CARDIFF, S. G., IFTICENE, E., JENKINS, R. B., KOFOKY, A., MBO- HOAHY, T., RAKOTONDRAVONY, D., RANIVO, J., RATRIMOMA- NARIVO, F., RAZAFIMANAHAKA, J. & RACEY, P. A. 2005. The distribution and conservation of bats ın the dry regions of Madagascar. Anımal Conservation 8: 153-165. GREENE, E. C. 1963. Anatomy of the rat. Hafner Publishing Com- pany, New York. HAWKINS, F. A., CHAPMAN, P., GANZHORN, J. U., BLOXAM, Q.M. C., BARLOW, S. C. & TONGE, S. J. 1990. Vertebrate conserva- tion in Ankarana Special Reserve, northern Madagascar. Bi- ological Conservation 54: 83-110. JANSA, S. A., GOODMAN, S .M. & TUCKER, P. K. 1999. Molecu- lar phylogeny and biogeography of the native rodents of Mada- gascar (Muridae: Nesomyinae): a test of the single-origin hy- pothesis. Cladistics 15: 253-270. JANSA, S. A., BARKER, F. K. & HEANEY, L. R. 2006. The pattern and timing of diversification of Philippine endemic rodents: evidence from mitochondrial and nuclear gene sequences. Sys- tematic Biology 55: 73-88. Musser, G. G. & CARLETON, M. D. 2005. Superfamily Muroidea. Pp. 894-1531 in: WILSON, D. E. & REEDER, D. M. (eds) Mammal species of the world. A taxonomic and geo- graphical reference, 3rd ed. The John Hopkins University Press, Baltimore. Nel, M. 1987. Molecular evolutionary genetics. Columbia Uni- versity Press, New York. NICOLL, M. E. & LANGRAND, O. 1989. Madagascar: Revue de la conservation et des aires protégées. WWE, Gland. PosaDA, D. & CRANDALL, K. A. 1998. ModelTest: testing the model of DNA substitution. Bioinformatics 14: 817-818. Rozas, J., SANCHEZ-DELBARRIO, J. C., MESSEGUER, X. & ROZAS, R. 2003. DnaSP, DNA polymorphism analysis by the coales- cent and other methods. Bioinformatics 19: 2496-2497. SOARIMALALA, V. & GOODMAN, S. M. 2003. Diversité biologique des micromammiferes non volants (Lipotyphla et Rodentia) dans le complexe Marojejy-Anjanaharibe-Sud. Pp. 231-278 in: GOODMAN, S. M. & WILME, L. (eds.) Dans Nouveaux ré- sultats d'inventaires biologiques faisant reference a l’altitude dans la région des massifs montagneux de Marojejy et d'An- janaharibe-Sud. Centre d’Information et de Documentation Scientifique et Technique, Antananarivo, Recherches pour le Développement, Série Sciences biologiques, No. 19. SWOFFORD, D. L. 2002. PAUP*: phylogenetic analysis using par- simony (*and other methods), ver. 4. Sinauer Associates, Inc., Sunderland, MA. Voss, R. S. 1988. Systematics and ecology of ichthyomyine ro- dents (Muroidea): patterns of morphological evolution in a small adaptive radiation. Bulletin of the American Museum of Natural History 188: 259-493. Authors’ addresses: Steven M. GOODMAN, Field Muse- um of Natural History, 1400 South Lake Shore Drive, Chicago, IL 60605, USA and Vahatra, BP 3972, Antana- narivo (101), Madagascar; E-Mail: sgoodman@fieldmu- seum.org; Martin RAHERIARISENA, Département de Biolo- gie Animale, BP 906, Université d’ Antananarivo, Antana- narivo (101), Madagascar and Vahatra, BP 3972, Antana- narivo (101), Madagascar; E-Mail: mraheriarisena@ya- hoo.fr; Sharon A. JANSA, Bell Museum of Natural Histo- ry, 1987 Upper Buford Circle, University of Minnesota, St. Paul, MN 55108, USA; E-Mail: jansa003@umn.edu. Received: 18.04.2008 Accepted: 05.06.2008 pene” Bonner zoologische Beiträge 56 149 APPENDIX SPECIMENS EXAMINED OF ELIURUS Listed below are specimens that formed the basis for the study’s morphological comparisons. FMNH numbers in bold indicate specimens also used in the molecular study. Abbreviations for protected areas in Madagascar are: PN, Parc National; RNI, Reserve Naturelle Integrale; RS, Reserve Spéciale. ELIURUS ANTSINGY: Mahajanga Province: Antsingy forest near Bekopaka, ca. 19°07.5°S, 44°49.0’E (MNHN 1966.2220, 1966.2222); PN de Bemaraha, 3.5 km E Bekopaka, near Tombeau Vazimba, 100 m, 19°08.4’S, 44%49.7”E (FMNH 172721); RNI [now PN] de Namoro- ka (FMNH 167563-167566; UADBA 16169); RNI [now PN] de Namoroka, 26 km NW Andranomavo, Forét d’Ambovonomby, 200 m, 16°28.2°S, 45°20.9?E (FMNH 175909, 175910, 175911); RNI [now PN] de Namoroka, 31 km NW Andranomavo, Forét de Mahabo, 100 m, 16°23.4°S, 45°20.9°E (FMNH 175912, 175913); RNI [now PN] de Namoroka, Site Andriabe, 2.0 km SE Namoroka (village), 16%24.4”S, 45°18.4’E, 110 m (FMNH 178587, 178591-178593). ELIURUS CARLETONT: Antsiranana Province: RS d’Ankarana, 2.6 km E Andrafiabe, forest near Andrafiabe Cave, 50 m, 12°55.9°S, 49°03.4’E (FMNH 169718, 169719, 169720); RS d’Ankarana, Campement des Anglais (Anilotra), 7.5 km NW Mahamasina, 125 m, 12°54.5°S, 49°06.6’E (FMNH 173104, 173105, 173106, 173109); RS d’Ankarana, 10 km NW Mahamasina, 100 m, 12°53.2’S, 49°06.6°E (FMNH 173107, 173108). ELIURUS DANIELI: Fianarantsoa Province: PN de l’Isalo, 28 km SE Berenty-Betsileo, along Sahanafa River near foot of Bevato Mountain, 22°19.0°S, 45°17.6’E, 650 m (FMNH 175934, UADBA 10483); PN de l’Isalo, 24 km W Ranohira bas, Andranohavo (Canyon des Rats), 22°28.9°S, 45°22.9°E, 700 m (FMNH 175933). ELIURUS MAJORT. Antsiranana Province: PN de Montagne d’Ambre, 12 km SW Joffreville, Grand Lac, 1325 m, 12°35.8°S, 49°09.6°E (FMNH 154345); PN de Montagne d’Ambre, 5.5 km SW Joffreville, 1000 m, 12°31.6°S, 49°10.3°E (FMNH 156341-156344, 154603-154609, 154610, 154611-154616). Province de Fianarantsoa : PN de Midongy-Sud, NE slope Mt Papango, 3.5 km SW Be- fotaka, 1100-1450 m, 23°50.3’S, 46°57.5’E (FMNH 178686). Bonner zoologische Beiträge | Band 56 Fn 3 ii Seiten 151-157 Bonn, September 2009 A tiny new species of Sylvisorex (Mammalia: Soricidae) from the Bamenda Highlands, Cameroon Rainer HUTTERER!, Jan RIEGERT? & Ondfej SEDLÁCEK3 ! Zoologisches Forschungsmuseum Alexander Koenig, Bonn, Germany; 2 University of South Bohemia, Ceské Budéjovice, Czech Republic; 3 Charles University, Praha, Czech Republic Abstract. A new species of shrew, Sy/visorex silvanorum n. sp., is described from specimens collected near Lake Bam- bili in the Bamenda Highlands of North West Cameroon. It is one of the smallest species of the genus occurring in the highlands of Cameroon and may be the sister taxon to 5. vulcanorum from eastern Africa. Together with the rodents Colomys eisentrauti and Lophuromys eisentrauti, the new species forms a set of species currently known only from the Lake Bam- bili area. Keywords. Africa, Cameroon, Lake Bambili, forest, shrew, Sy/visorex, new species. 1. INTRODUCTION Shrews of the genus Sylvisorex Thomas, 1904 are restrict- ed to lowland and montane forests of central Africa, from Cameroon/Nigeria in the northwest to Tanzania in the southeast. Currently 12 extant species are recognized (HUTTERER 2005), but recent fieldwork has yielded fur- ther species that will be described in the near future. Oth- er species new to science still rest in museum collections and await formal description, such as the one named and described below. It was recognized as new by the senior author more than 30 years ago but known only from a sin- gle incomplete specimen since until additional material be- came available from owl pellets collected in 2003 by the junior authors which finally enabled us to define the new species. 2. MATERIAL AND METHODS The holotype of the new taxon was collected during a Mu- seum Koenig expedition to Cameroon from winter 1973 to spring 1974; see BÖHME (1975) for a brief description of the expedition. The participants W. BÓHME and W. HARTWIG visited the Bafut Ngemba Forest Reserve and the adjacent Bambili craters during March 1974. Shrews were collected with snap traps set along creeks and in un- dergrowth. The few shrews collected were briefly dis- cussed by HEIM DE BALSAC (1975) but never properly do- cumented. Pellets of the African grass owl Tyto capensis were col- lected during November 2001 in the lower of the two Bambili volcano craters, about 8 km SE from Bamenda city, Bamenda Highlands, Cameroon (5° 55” N, 10° 14” E, 2 400 m a.s.1.) (RIEGERT et al. 2008). Both volcano calderas were flooded by lakes in the past (ANONYMOUS 1972). The lake persists only in the lower crater, the up- per one has been gradually covered by wet grasslands. Skull measurements were taken with an electronic calliper. All measurements are given in millimetres (mm), and body mass in grams (g). The terminology of cranial and dental structures follows MEESTER (1963) and JENKINS (1984); external and cranial measurements are the same as previously defined by STANLEY et al. (2005) and KER- BIS PETERHANS et al. (2008). Voucher specimens men- tioned in the text are deposited in the Zoologisches Forschungsmuseum Alexander Koenig, Bonn (ZFMK) and in The Natural History Museum, London (BMNH). The taxonomy follows HUTTERER (2005). 3. THE NEW AFRICAN PYGMY SHREW Sylvisorex silvanorum n. Sp. Holotype and type locality. Skin and partial skull of a young adult female (ZFMK 74.430), collected by W. 152 Rainer HUTTERER et al.: A tiny new species of Sylvisorex from the Bamenda Highlands, Cameroon Fig. 1. Sylvisorex silvanorum n. sp., skin of holotype (ZFMK 74.430) in dorsal and ventral view. Total length 1s 99 mm. Paratypes. Two incomplete skulls (ZFMK 2003.1103, 2003.1104) removed from pellets of Tyto capensis, col- lected November 2001 by J. Riegert and O. Sedlácek on a slope of the lower Bambili volcano craters (5° 55’ N, 10° 14° E, 2 400 m a.s.l.), about 8 km SE of the city of Bamenda, Bamenda Highlands, NW Province, Camty of Bamenda, Bamenda Highlands, NW Province, Cameroon (RIEGERT et al. 2008). Diagnosis. A very small and dark species of Sylvisorex with a total length below 100 mm, tail 85 % of head and body length; no long bristle hairs on tail. Description. Sy/visorex silvanorum n. sp. is a small ter- restrial shrew with dark fur. The hairs on the dorsum are dark brown, between Fuscous (RIDGwAY 1912, plate XLVI) and Chaetura Drab. Ventrally the hairs gradually turn into Hair Brown. Hairs on dorsum are about 3.4 mm in length; ventrally they are slightly shorter. The basal por- tion of individual hairs is Dark Plumbeous (Plate LII) with tips Drab to Hair Brown (Plate XLVI). The entire body, including head, ears, limbs, and tail have the same dark colour, only the central part of the belly is a bit lighter (Fig. 1). Facial vibrissae reach 14 mm in length. Ear conch is small, round and pocketed, and covered by very short hairs, except for the inner fold which carries a row of long hairs. Hind foot narrow, with elongate digits and short claws. Ventral inner surface of hind foot covered by small granule-like bumps, heel covered by short hairs. Tail of Table 1. External measurements of Sy/visorex silvanorum and other small species of the genus with no long bristle hairs on the tail. Species TE HB Tail S. johnstoni! 64 39 25 S. silvanorum! 99 55 44 S. vulcanorum? 95.9 49 46.9 S. camerunensis | 122 65 57 S. grant? 114 58 55.9 S. isabellae! 119 60 59 S. morio3 119 71 51 HF Ear Wt Tail/HB 9 = - 64% 10.0 4.8 = 85% 10.4 6.4 3:3 96% 12 9 87.6% 11.6 1:3 = 96.4% 13 6 8 98.3% 13 10 8 72% ! holotype; 2 means of measurements taken from HUTTERER & VERHEYEN (1985); 3 ZFMK 69.370. Bohme and W. Hartwig on 07.03.1974 (field no. 98) near Lake Bambili, Mt. Lefo, Bafut Ngemba F. R. (5° 55” N, 10° 14” E), 1 800-1 900 m a.s.l., Bamenda Highlands, North West Province, Cameroon. Measurements of holotype. Head and body length 52 mm, tail 44 mm, hind foot length 11.5 mm, ear conch 5.0 mm. medium length (85% of head and body length); colour uni- form, no long bristle hairs present. Skull (Figs. 2, 3) short with a rounded braincase. Dorsal profile (Fig. 3) inflated, rostrum short but slender. Interor- bital constriction relatively broad (24.9% of estimated condylo-incisive length). Infra-orbital bridge narrow, lachrymal foramen large. Bonner zoologische Beiträge 56 153 Fig. 2. Sylvisorex silvanorum n. sp., skull of holotype; cranium and upper dentition in occlusal view, and left mandible in labial view. Scale 1 mm. Upper teeth: First upper incisor small, with a sharp ante- rior tip and a well-developed talon. First upper unicuspid large, second and third unicuspids half and fourth unicus- pid a quarter of the length of the first unicuspid. P4 with a small parastyle and a large paracone; the protocone is well expressed. M1-2 square-shaped in occlusal view, pro- tocone strongly developed. M3 relatively large (0.69 x 1.14 mm in the holotype). Mandible: Slender with thin ramus and weak coronoid process. Foramen mentale below posterior root of lower p4. Condyle small, oblique, and about as wide as high. Lower teeth: Tip of first incisor curved upwards, with two denticulations on cutting blade. Lower p4 as long as wide (occlusal view). Lower molars long and slender; third low- er molar (1.11 x 0.54 mm) with a deep talonid basin. Comparisons. The species is not easily recognized in the field. HEIM DE BALSAC (1975) identified all three speci- mens of Sylvisorex collected in 1974 near the Bambili crater as belonging to “Sy/visorex granti Thomas, 1907”, but a subsequent inspection of the same material by the senior author revealed that one (field no. 98) represented the new S. silvanorum, another (field no. 106) S. camerunensis, and a third (field no. 113) a new species (not yet described) related to S. morio, a species restrict- ed to Mount Cameroon. S. silvanorum is smaller in ex- ternal and cranial measurements (Tables 1, 2) than most other species of Sylvisorex, with the exception of S. john- 154 Rainer HUTTERER et al.: A tiny new species of Sylvisorex from the Bamenda Highlands, Cameroon Table 2. Cranio-dental measurements of Sylvisorex silvanorum and allies. Species El PL UTR IO MB GW PGL HCE LTR COR S. johnstoni! 15.0 6.1 6.4 3.8 4.7 7.1 - 3.8 5.9 355 S. silvanorum? - 5.80 6.36 3:99 4.61 - - - 5292 3.45 S. silvanorum? c.15.6 5.90 6.72 3.82 3.13 7.94 5.57 5.20 - - S. silvanorum* = z 7.20 3.83 - - - . 5.80 3.74 S. vuleanorum> 15.8 5.90 6.6 4.0 5.0 7.8 5.28 4.90 6.1 3.6 S. camerunensis® 17.14 6.85 7.46 4.12 3.13 8.50 3.19 4.77 6.81 3.89 S. granti! 16.60 6.68 7.19 3.99 5.28 8.12 5.42 4.41 6.48 3.96 S. isabellae® 18.81 7.55 8.14 4.43 iz 9.30 6.08 5.41 7.48 4.42 S. morio? 19.63 8.22 8.55 4.50 5.56 9.38 6.22 5.44 7.87 4.30 ! holotype BMNH 87.11.26.1; 2 holotype ZFMK 74.430; 3 paratype ZFMK 2003.1103; * paratype ZFMK 2003.1104; 5 average measurements of type series (n=6) from HUTTERER & VERHEYEN (1985); © holotype ZFMK 69.358; 7 ZFMK 68.538; $ holotype ZFMK 69.365; ? ZFMK 69.370; all measurements taken by RH. Fig. 3. Sylvisorex silvanorum n. sp. (upper figure, combined drawing taken from holotype and paratype), lateral view of the skull compared to Sylvisorex vulcanorum (lower figure, holotype from HUTTERER & VERHEYEN 1985); scale 1 mm. Bonner zoologische Beitráge 56 155 Table 3. Species of small mammals recorded from the Bambili area in 1974 and 2001. 2001 ** Bubo cinerascens 2001** Tyto capensis 1974* Traps, shotgun Species Species endemic to Bamenda Highlands Species endemic to Cameroon Mountains ai rrr ee a Sylvisorex silvanorum n. sp. Sylvisorex cf. morio Sylvisorex camerunensis Myosorex okuensis SE We ee an lon Crocidura olivieri Crocidura virgata : 5 Crocidura attila Paraxerus cooperi 11 Graphiurus lorraineus NW Lophuromys eisentrauti Lophuromys sikapusi Hybomys eisentrauti 4 Colomys eisentrauti 3 Dasymys rufulus 1 Hylomyscus aeta 1 Lemniscomys striatus Praomys hartwigi 1 Otomys occidentalis Mastomys sp. Mus musculoides Mus setulosus Muridae indet. Totals 35 vr + WN N o ON ies) KH wr x KK ~~ KK nS Rune x 33 8 10 * HEIM DE BALSAC (1975), DIETERLEN (1979, 1983), HUTTERER et al. (1992), VAN DER STRAETEN & HUTTERER (1984), HUTTERER (unpublished); ** RIEGERT et al. (2008). stoni (Dobson, 1888). The latter has a much shorter tail, shorter hind feet, and a shorter skull (HUTTERER & VER- HEYEN 1985). There is only one species which is similar in size and form: Sylvisorex vulcanorum Hutterer & Ver- heyen, 1985, a species known only from high elevations (1900-3000 m) of the Albertine Rift, separated from the Bamenda Highlands by some 2000 km of lowland forest and savannah. S. silvanorum n. sp. differs from S. vulcano- rum by a shorter tail (44 versus 46.9 mm), and by cranial details. In S. silvanorum n. sp., the lachrymal foramen is large (small in S. vulcanorum), the first upper incisor is pointed (short and rounded in $. vulcanorum), and the sec- ond and third upper unicuspids are about equal in size (sec- ond smaller than third in $. vulcanorum). The few avail- able measurements may also indicate that the upper tooth row is slightly shorter in S. vulcanorum. It is probable that both species represent sister taxa, a hypothesis which will have to be tested with genetic data in the future. Some other small species of Sy/visorex, such as S. kon- ganensis Ray & Hutterer, 1996, S. pluvialis Hutterer & Schlitter, 1996, or S. akaibei Mukinz1, Hutterer & Barriere, 2009, were not considered because they are easily distin- guished, even in the field, by their hairy tails; they also have very different skulls (RAY & HUTTERER 1996, HUT- TERER & SCHLITTER 1996, MUKINZI et al. 2009). Small and dark-furred species of Suncus which occur in Central African lowland and mountain forest, e.g. S. infinitesimus (Heller, 1912), S. remyi Brosset, Dubost & Heim de Bal- sac, 1965, S. hututsi Kerbis Peterhans & Hutterer, 2009, are all smaller in external and cranial dimensions, and have shorter tails covered with long and numerous bristle hairs (KERBIS PETERHANS & HUTTERER 2009). Habitat. The holotype specimen was collected in dense undergrowth along a stream running down from the relict montane forest of the reserve towards the Bambili crater 156 Rainer HUTTERER et al.: A tiny new species of Sylvisorex from the Bamenda Highlands, Cameroon Fig. 4. Lower Bambili crater showing relict montane forest on the slopes and montane grassland along the shore of the lake. Photograph taken in 2001. (W. BÖHME, field notes). The slopes of the lower Bambili crater are covered by a mosaic of montane forest, La- siosiphon and bamboo woods and grassland. The slopes are partly rocky (Fig. 4). The surroundings of the craters are covered mainly by extensive pastures, Pteridium fern growth and open Lasiosiphon woods. It is not known where the owls picked up the shrews but the slope forest and the wet grassland bordering the lake may constitute suitable habitats. Etymology. Derived from si/vanus (Latin, meaning for- est god). We dedicate this cryptic pygmy shrew to the syl- vans, small and invisible forest creatures occurring in the beliefs, mythologies, and fairy tales of Africans, Euro- peans, and many other peoples on earth. As a common name we suggest “Bamenda pygmy shrew”. DISCUSSION The forests around Mt. Lefo are still full of mysteries. Dur- ing their field work in 1974, BOHME and HARTWIG collect- ed 35 specimens, representing 12 species of shrews and rodents (Table 3). Four of the species were subsequently described as new taxa: Colomys (goslingi) eisentrauti Di- eterlen, 1983, Lophuromys (sikapusi) eisentrauti Dieterlen, 1979, Hybomys eisentrauti Van der Straeten & Hutterer, 1986 (tentatively assigned to Hybomys cf. badius by Musser & CARLETON 2005), and Sylvisorex silvanorum n. sp. (see DIETERLEN 1979, 1983, VAN DER STRAETEN & HUTTERER 1986). A further species of Sylvisorex still awaits description. HUTTERER et al. (1992) proposed full species rank for Lophuromys eisentrauti, and further but yet unpublished research suggests that Colomys eisentrauti and Hybomys eisentrauti may deserve species status as well. The forests around Mount Lefo (including the Bam- bili craters) apparently house a number of local endemic mammals, some of which may be threatened with extinc- tion. The local Colomys eisentrauti and Lophuromys eisen- trauti have not been found again since their discovery in 1974. At least four mammal species are only known from the Lefo Mountains, 8 from the Bamenda Plateau, and more than 10 are endemic to the Cameroon Mountains chain (HUTTERER et al. 1992). STAGER & ANFANG-SUTTER (1999) provided evidence from lake sediment cores of pro- nounced climatic changes in the West Cameroon High- lands since 24,000 years BP, particularly in the Lake Bam- bili area. It is probable that such climatic fluctuations trig- gered speciation in the Bambili area and other highlands in NW Cameroon (EISENTRAUT 1973). Acknowledgements. We wish to thank GACR (206/03/H034), MSM (LC 06073, 6007665801 and 0021620828) and GAAV (IAA601410709 and KJB601110703) for financial support of the research of JR and OS in Cameroon. We further thank A. Riegertová, D. Sedlácková, S. Janeéek and P. Janeckovä for as- sistance in the field. RH thanks Paulina D. Jenkins for access to the collections of the British Museum. Karin Ulmen assisted with the SEM graphs, and Uwe Vaartjes mounted figure 2. Zusammenfassung. Eine neue Spitzmausart, Sylvisorex silva- norum n. sp., wird nach Material, das am Bambili-Kratersee im Bamenda Hochland Nordwest-Kameruns gesammelt wurde, be- schrieben. Es handelt sich um eine der kleinsten Arten der Gat- tung, die vermutlich die Schwesterart zu S. vulcanorum aus Ost- afrıka ist. Wie die Nagerarten Colomys eisentrauti und Lophu- romys eisentrauti ist die neue Sylvisorex-Art bisher nur aus der Umgebung des Bambili-Kraters bekannt. REFERENCES ANONYMOUS (1972): Baffousam NB-32-XI, Carte du Cameroun. L Institut Géographique National, Yaounde. BÖHME, W. (1975): Zur Herpetofaunistik Kameruns, mit Be- schreibung eines neuen Scinciden. Bonner zoologische Bei- träge 26: 2-48. DIETERLEN, F. (1979): Zur Kenntnis der Gattung Lophuromys (Muridae; Rodentia) in Kamerun und Gabun. Bonner zoolo- gische Beiträge 29: 287-299. DIETERLEN, F. (1983): Zur Systematik, Verbreitung und Ökolo- gie von Colomys goslingi Thomas & Wroughton, 1907 (Mu- ridae; Rodentia). Bonner zoologische Beiträge 34: 73-106. EISENTRAUT, M. (1973): Die Wirbeltierfauna von Fernando Poo und Westkamerun unter besonderer Bedeutung der pleistozá- nen Klimaschwankungen für die heutige Faunenverteilung. Bonner zoologische Monographien 3: 1-428. HEIM DE BALsAc, H. (1975): Nouvelles données sur la fauna so- ricidienne du Cameroun. Bonner zoologische Beitráge 26: 94-99, HUTTERER, R. (2005): Order Soricomorpha. In: WiLson, D. E. & REEDER, D. A., eds, Mammal species of the world: A taxonomic and geographic reference. Third edition. Johns Hopkins University Press, Baltimore. HUTTERER, R., DIETERLEN, F. & NIKOLAUS, G. (1992): Small mammals from forest islands of eastern Nigeria and adjacent Cameroon, with systematical and biogeographical notes. Bon- ner zoologische Beiträge 43: 393-414. Bonner zoologische Beiträge 56 157 HUTTERER, R. € VERHEYEN, W. (1985): A new species of shrew, genus Sylvisorex, from Rwanda and Zaire (Insectivora: Sori- cidae). Zeitschrift für Säugetierkunde 50: 266-271. HUTTERER, R. & SCHLITTER, D. A. (1996): Shrews of Korup Na- tional Park, Cameroon, with the description of a new Sylvi- sorex (Mammalia: Soricidae). Pp. 57-66, In: Contributions ın Mammalogy: A Memorial Volume Honoring Dr. J. Knox Jones, Jr. Museum of Texas Tech University, il + 315 pp. JENKINS, P. D. (1984): Description of a new species of Sy/visorex (Insectivora: Soricidae) from Tanzania. Bulletin of the British Museum of Natural History (Zoology) 47: 65-76. KERBIS PETERHANS, J. C. & HUTTERER, R. (2009): The descrip- tion of a new species of Suncus (Soricidae, Mammalia) from central Africa. Pp. 141-150 in: THORN, E. & KERBIS PETER- HANS, J. (eds), Small mammals of Uganda. Bonner Zoologis- che Monographien 55: 1-164. KERBIS PETERHANS, J. C., HUTTERER, R., KALIBA, P. & MAZ- IBUKO, L. (2008): First record of Myosorex (Mammalia: So- ricidae) from Malawi with description as a new species, Myosorex gnoskei. Journal of East African Natural History 97: 19-32. MEESTER, J. (1963): A systematic revision of the shrew genus Crocidura in southern Africa. Transvaal Museum Memorr 13: 1-127. MUKINZI, I., HUTTERER, R. & BARRIERE, P. (2009): A new species of Sylvisorex (Mammalia: Soricidae) from lowland forests north of Kisangani, Democratic Republic of Congo. Mam- malia 73: 130-134. Ray, J. C. & HUTTERER, R. (1996): Structure of a shrew com- munity in the Central African Republic based on the analysis of carnivore scats, with the description of a new Sy/visorex (Mammalia: Soricidae). Ecotropica 1: 85-97. RIDGWAY, R. (1912): Color standards and color nomenclature. Washington, D. C., published by the author. RIEGERT, J. SEDLÁCEK, O. & HUTTERER, R. (2008): Diet of sym- patric African grass owl (Tyto capensis) and Spotted eagle owl (Bubo africanus) in Bamenda Highlands, NW Cameroon. African Journal of Ecology 46: 426-431. STAGER, J. C. & ANFANG-SUTTER, R. (1999): Preliminary evi- dence of environmental changes at Lake Bambili (Cameroon, West Africa) since 24,000 BP. Journal of Paleolimnology 22: 319-330. STANLEY, W. T., ROGERS, M. A. & HUTTERER, R. (2005): A new species of Congosorex from the Eastern Arc Mountains (Tan- zania), with significant biogeographical implications. Journal of Zoology 265: 269-280. VAN DER STRAETEN, E. & HUTTERER, R. (1986): Hybomys eisen- trauti, une nouvelle espece de Muridae du Cameroun (Mam- malia: Rodentia). Mammalia 50: 35—42. Authors’ addresses: Dr. Rainer HUTTERER, Zoologisches Forschungsmuseum Alexander Koenig, Section of Mam- mals, Adenauerallee 160, D-53113 Bonn, Germany, e- mail: r.hutterer.zfmk@uni-bonn.de; Jan RIEGERT, Univer- sity of South Bohemia, Faculty of Biological Sciences, Department of Zoology, BraniSovskä 31, CZ-370 05 Ceské Budéjovice, Czech Republic, e-mail: honza@riegert.cz; Ondfej SEDLACEK, Charles University in Prague, Faculty of Science, Department of Ecology, Viniéna 7, CZ-128 44 Praha, Czech Republic, email: zbrd@email.cz. Received: 15.12.2008 Accepted: 30.03.2009 Bonner zoologische Beiträge | Band 56 Heft 3 Seiten 159-173 Bonn, September 2009 Altitudinal distribution and anthropogenic influence on small mammal assemblages on Mount Kupe, SW Cameroon Christiane Denys!, Alain Didier Missoup!, Barthelemy TCHIENGUE3, Gaston ACHOUNDONG}, Atanga Exo0BO*, Charles Felix BILONG BILONG2, Dieudonné Massoma LEMBES & Violaine NICOLAS! ! Museum national d’Histoire naturelle, Dept. Systématique et Evolution, Paris, France 2 Department of Biology of Animal Organisms, University of Yaounde I, Yaoundé, Cameroon 3 Herbier National du Cameroun, Yaoundé, Cameroun 4 WWE, Coastal Forests Programme, Limbé, Cameroon 5 Département de Biologie des Organismes Animaux, Université de Douala, Douala, Cameroun Abstract. We conducted a taxonomical inventory of small mammal biodiversity on Mount Kupe, SW Cameroon, the se- cond inventory on that mountain after M. Eisentraut's more than 50 years ago. Our survey yielded a total catch of 19 species of mammals: 16 rodents, two bats, and one shrew. For each species we summarize data on natural history and discuss taxonomical and distributional aspects. We observed a difference between the rodent assemblages of the lowland (below 1000 m) (including Praomys misonnei) and the upper sub-montane and montane forest (1500-2000 m) (inclu- ding Hylomyscus cf. walterverheyeni, identified for the first time from the Cameroon Mountains). This seems to corre- spond to previously described vegetation changes between 1000 and 1500 m. The inhabited and cultivated zones at the base of Mt. Kupe yielded eight widely-distributed rodent species. A number of taxonomical problems concerning mon- tane forest species remain to be solved. Keywords. Africa, Cameroon, Chiroptera, Rodentia, Soricomorpha, taxonomy, diversity. 1. INTRODUCTION Tropical regions are well known to have a higher biodi- versity than temperate regions. The forests of west-cen- tral Africa have been described as one of the tropical “hot spots” (MYERS et al. 2000; Kuper et al. 2005). African mountains seem to harbour a very important endemic di- versity and are of exceptional conservation importance (HEIM DE BALSAC 1968; EISENTRAUT 1973; GENTRY 1992; STANLEY et al. 1998, 2005; KASANGAKI et al. 2003; HER- RMANN et al. 2005; BURGESS et al. 2007). Both lowland and mountain forests are presently subject to intense de- forestation. Several authors have emphasized the role of African forest refugia in promoting speciation during Qua- ternary climatic changes (MALEY 1987, 1996; QUEROUIL et al. 2003, PLANA et al. 2004; BEHNIN 2006, HUHNDORF et al. 2007). However, the knowledge of the mechanisms by which so many species arise and differentiate in such regions depends directly on a precise characterization of their biodiversity in relation to altitudinal and vegetation shifts along elevational gradients. The altitudinal effect on small mammal diversity is not well known and available studies differ in their results. According to some authors species-richness is not uniformly distributed on mountains, but the base and the top hold a higher diversity than the intermediate zones (BROWN 2001; LOMOLINO 2001). Oth- er authors found a continuous decrease of diversity with elevation (Diaz et al. 2002; KASANGAKI et al. 2003) or con- trasting patterns in different taxonomic groups (HOFER et al. 2000). However, a positive correlation between species turnover and elevation was recently evidenced for rodents of different tropical mountains (MENA & VASQUEZ- DOMINGUEZ 2005). The authors considered climate, pro- ductivity and landscape heterogeneity as well as histori- cal factors as possible explanations for elevational gradi- ents. There are many other vertebrate groups for which zona- tion has been described as a function of altitude in African mountains (e.g. EISENTRAUT 1968; AMIET 1987; STUART et al. 1987; Kıryo & CLAUSNITZER 2001; KASANGAKI et al. 2003) but the observed changes seem to vary locally and their causes are not well understood. It is well known that the composition of plant communities changes along altitudinal gradients but the respective roles of vegetation or available streams versus altitudinal change remain un- 160 Christiane DENYS et al.: Small mammales of Mt. Kupe, Cameroon clear (HOFER et al. 2000). Because most terrestrial small mammal species are primary consumers or have limited dispersal capacities (restricted ranges and/or habitats) they are good models for studying altitude-diversity gradients. Situated in the Central African forests blocks of the Guineo-Congolese hot spot of biodiversity and extending for about 1600 km, the Cameroon Volcanic Line (CVL) is a succession of high volcanoes and plateaus ranging in age between 30 and one Myr (UBANGOH et al. 1998). At a distance of about 3000 km from the Western Rift Val- ley of East Africa, it represents the second highest range of mountains in Africa, with Mount Cameroon at 4100 m and Mount Oku at 3011 m. Because of difficult access the Cameroon Highlands ecoregion has been poorly sampled for its small mammal fauna. Small mammal expeditions were launched by EISENTRAUT (1957, 1961, 1963, 1968), LAMOTTE (1950-1970, unpublished), and SCHLITTER et al. (1999), among others. These studies showed a high en- demism of the region, particularly in Afro-alpine grass- lands (above 2000-2500 m) and in mountain forests (EISENTRAUT 1963; HEIM DE BALSAC 1968; LAMOTTE & PETTER 1981; PETTER 1986; DIETERLEN & VAN DER STRAETEN 1992; HUTTERER et al. 1992; VERHEYEN et al. 1997). Studies on other vertebrates and plants have also described many endemics, emphasizing the importance of this region (STUART 1986; AMIET 1987; SOSEF 1994; LAR- ISON et al. 2000; CHEEK et al. 2004; HERRMANN et al. 2005). Most previous biodiversity inventories focused on the two highest mountains (1.e. Mount Cameroon and Mount Oku). The biodiversity of the other mountains in the region is poorly known (summarized by HUTTERER et al. 1992). Among the CVL highlands, Kupe Mountain (2064 m) harbours a well preserved mountain forest in the Bakossı Highlands. This forest is conserved on the west- ern side of the mountain, thanks to the Bakossi people and a nature reserve project monitored by the World Wildlife Fund (WWF). While its vegetation has been extensively studied (CHEEK et al. 2004; TCHIENGUE 2004), its small mammal diversity is less well known; EISENTRAUT (1958, 1963) published a faunal list for this mountain. Mt. Kupe is famous for the presence of ten primate species, among them drill (Mandrillus leucophaeus) and chimpanzee (Pan troglodytes) populations, which are threatened with extinc- tion (WILD et al. 2004, 2005). The forest elephant (Lox- odonta africana cyclotis) is also present on Mt. Kupe (WILD 2004). Numerous woody plant species of the low- land forest exhibit distinct sub-montane forest ‘forms’ (LETOUZEY 1985). Most of them still have no taxonomic status and were not included in assessments of diversity, endemism and the impacts of ecological change. Several studies (e.g. BRETELER 1999; Mc Key 2000) have demon- strated the distinctness of sub-montane forest species, as well as their key role in understanding the history of the lineages concerned. With this information as a background we concentrated our small mammal survey on the montane and sub-mon- tane species. Our study should be of conservation value and provide further biogeographical hypotheses. In this pa- per we describe the composition of the small mammal community of Mt. Kupe along an altitudinal gradient. In order to better describe the Mt. Kupe small mammal com- munity we made a morphological and molecular inven- tory of small mammal species to accurately describe the degree of endemism of the Kupe fauna, and eventually al- so look at a possible altitudinal difference between sub- montane and montane communities. This should further allow us to evaluate the importance of the CVL for small mammal biodiversity conservation and diversification. 1.1. Background Previous information on the Mt. Kupe small mammal fau- na: Small mammals from the Cameroon Volcanic Line (CVL) were described by SANDERSON (1940) and EISEN- TRAUT (1957, 1961, 1963, 1968, 1969). They mainly stud- ied the small mammals from Mt. Cameroon and Bioko Is- land, but also collected specimens from the massif of Mt. Kupe. M. Lamotte’s surveys (1970-1975) followed and his collections allowed the description and rehabilitation of endemic species in the Bamenda Highlands (LAMOTTE & PETTER 1981; PETTER 1986; MUSSER & CARLETON 2005), but not the Bakossi Highlands. In an unpublished report DOWSETT-LEMAIRE & DOWSETT (1998) provided a faunal list of the Bakossi Mountains. This list contained a great number of unidentified species. The study area: Mt. Kupe is situated on the western slopes of the CVL. It is a steep-sided massive horst block formed during Cameroon’s third eruptive phase in the Quaternary (WILD 2004). Mt. Kupe is the youngest mountain of the Bakossi Highlands. The mountains of this highland are among the wettest areas in Africa (around six to seven me- tres annual rainfall, CHEEK et al. 2004). Approximately 80 % of it occurs in the April-October wet season but in gen- eral every month of the year experiences rainfall. Never- theless, there is a dry season between November and March, and a single week or two of bright sunshine with- out rain usually occurs in July (the “short dry season”). Humidity is exceptionally high. The minimum relative hu- midity (81 %) is recorded during the dry season. Descriptions of the Kupe forest vary from one author to another. All authors agree that the lowland evergreen for- est occurs between 150 to 800 m and is restricted to the western and southern part of Mt. Kupe. The sub-montane forest is found from 800 m asl to the summit; CHEEK et al. (2004) consider it to be constituted by mid-altitude for- est and Afro-montane rain forest. Previously, THOMAS Bonner zoologische Beiträge 56 161 (1986) recognized only two forest types: the lowland for- est below 1600 m and the sub-montane forest above. This classification is not followed by LANE (1994) or by CHEEK et al. (2004). The sub-montane forest is characterized by the presence of Podocarpus milanjianus (Podocarpaceae) around 1600-2000 m, Santiria trimera (Burseraceae) and numerous large Guttiferae (Clusiaceae) of the genera Al- lanblackia, Pentadesma, and Symphonia between 1000 m and 2000 m, and Cola (Sterculiaceae) also between 1000 m and 2000 m (LETOUZEY 1968). LETOUZEY (1968) divides the sub-montane forest into three blocks. According to CHEEK et al. (2004) this point of view was based mainly on aerial observation and it is possible that the band of vegetation influenced by human activity in this area was misinterpreted as sub-montane thicket. CHEEK et al. (2004) divided the sub-montane forest into two altitudinal types: the lower sub-montane forest (800-1400 m), harbouring a high epiphyte and saprophyte diversity with some of the Bakossi Highlands endemics, and the upper sub-montane forest (1400-1900 m) which is less diverse. According to CHEEK et al. (2004) there is also a montane forest (from 1900 m to the summit), characterized by the absence of Clausena anisata, Laesa lanceolata, and Pavetta hooke- riana, all otherwise frequent in montane forests of the Cameroon Highlands. On the other hand, it contains many endemic species such as Leptoderris fasciaculata, Bate- santhus purpureus (also sub-montane), Ficus oreodryad- um, Zanthoxylum leprieurii, and Aframomum zambezi- acum. A patch of mountain grassland is found on each of the three peaks of the summit which are up to 100 m wide only (CHEEK et al. 2004). These mountain grasslands are constituted by two species only: Panicum hochstetteri and Gladiolus aequinoctialis. 2. MATERIALS AND METHODS Small mammals were trapped during three periods of fieldwork. December 2005: around Nyasoso (N 04° 50” / E 09° 40”) and on the mountain below at 950 m; Febru- ary 2006: between 1500 m and 2000 m; August 2006: around 1000 m (Table 1). Lines of 250 Sherman traps bait- ed with peanuts, flour, oil and dry fish were used each time and were set in six habitats: cultivated lowland and sec- ondary forest (100-800 m), lower sub-montane forest (900-1000 m), upper sub-montane forest (1500 m), mon- tane forest and montane grassland (2000 m). The traps were checked twice per day. Additionally, a number of tra- ditional traps set by hunters were put randomly in the sur- roundings of Nyasoso village and at the summit. We al- so put two mist nets at the entrance of a cave (900 m asl) during two nights in December 2005 in order to catch Chi- roptera. Due to logistic constraints the number of trap nights varied between habitats. Standard external meas- urements (weight [W], head and body length [HB], tail length [TL], ear [E] and hindfoot lengths [HF]) and the sex of each animal were taken. All animals were autop- sied. The liver was kept in 70 % ethanol for molecular identifications and the carcasses preserved in 4 % forma- lin, and the skulls cleaned. All this material was includ- ed in the MNHN collections, Paris. Comparisons were per- formed with specimens previously collected from Cameroon (Nditam and Dja regions), Gabon (Makokou, Belinga) and Central African Republic (La Maboké). All species were identified based on information published in ROSEVEAR (1969), VIVIEN (1991) and WILSON € REEDER (2005). For species identification we performed classical morpho- metric analysis on both external and cranial distances, us- ing a digital caliper (MITUYOTO, precision 0,01 mm) and the protocol described by DENYS & TRANIER (1992). Due to the abundance of sibling species in tropical Africa, 81 specimens were sequenced by one of us (ADM) for con- firmation of identification (Cytochrome b and/or 16S genes). These specimens were compared using the Neigh- bour Joining Tree Analysis and the sequences available in Genbank. To study the small mammal communities, we used the Shannon-Wiener diversity index H = - y p;Inp; as a direct measure of species diversity in each habitat, as well as a correspondence analysis on presence-absence data. The correspondence analysis CFA (BENZECRI & BENZECRI 1984) was performed on the presence-absence of species in different habitats. By reducing the number of variables and using the chi-square distance, this method allows the representation of the relationship between variables (here habitats) and individuals (here species) on the same graph (scatter plot type) and gives an overview of the distribu- tion of species diversity. All statistical analyses were per- formed by using XLSTAT 7.1 software (Addinsoft). 3. RESULTS 3.1. Trapping success, faunal lists In total, 98 small mammals belonging to 19 species were captured on Mt. Kupe. Overall trap success with Sherman traps was 3,03 %. This varied among habitats: trap suc- cess was always low in the forest, varying from 1,86 % to 2,88 %, depending on the altitude; it was higher in habi- tats under anthropogenic influence (5,51 %; Table 2). The Shannon diversity index indicates quite an equal distribu- tion of species abundances in the habitats under anthro- pogenic influence and the low diversity of the sub-mon- tane and montane forests. 162 Christiane DENyYs et al.: Small mammales of Mt. Kupe, Cameroon Table 1. Characteristics of the different habitats investigated and periods when they were sampled. Sampling period Elevation range (m) Main vegetation Main habitats GPS coordinates Collecting method investigated Dec 2004 beginning of 850 anthropogenic farmland - Traditional traps dry season lowland forest closed canopy — Traditional traps anthropogenic fallow, farmland N 04° 50° 10,1”/ Sherman traps (60) lines A,B,C,F E 09° 40” 48,5” Dec 2005 beginning of 850-950 anthropogenic home garden N 04° 50” 09,6”/ Sherman traps (40) dry season E 09° 40° 51,2” lowland forest closed canopy N 04° 49” 26,0”/ Sherman traps (160) lines D,E,T,U,V E 09° 41” 11,9” 850 anthropogenic fallow, farmland, N 04° 50” / Sherman traps (100) line C home garden E 09° 40° Febr 2006 end of dry season 1500-1600 sub-montane forest closed canopy, N 04° 49° 08,7”/ Sherman traps (250) lines O,P,Q,R,S dense under story E 09° 42° 27,1” 1900-2000 montane forest closed canopy N 04° 47’ 51,1”/ Sherman (250) and lines K,L,M E 09° 42° 11,2” traditional traps montane grassland woodland, N 04° 47° 52,7”/ Sherman traps (250) lines 1,J savannah E 09° 42’ 23,9” August 2006 little dry season 850 lowland forest closed canopy N 04° 49° 26,0°7/ Sherman traps (250) E 09° 41° 11,9” Table 2. Details of trapping efforts and success in each habitat sampled. The trapping success corresponds only to animals cap- tured in Sherman traps and the Shannon Index was calculated on pooled Sherman + Traditional trap captures in all habitats. Species/Trap Anthropogenic Lowland forest Sub-montane forest Montane success/Diversity/H’ forest 2000m Sher Trad Sher Trad Sher Trad Sher Trad Crocidura poensis complex 4 0 | 0 0 0 0 0 Cricetomys cf. emini 0 | 0 0 0 0 0 3 Dendromus sp. 0 | 0 0 0 0 0 0 Deomys ferrugineus 0 0 0 0 2 0 0 0 Grammomys poensis 2 | 0 0 0 0 0 0 Hybomys cf. univittatus 0 0 l 0 0 0 0 0 Hylomyscus walterverheyeni 0 0 10 0 12 0 12 0 Hylomyscus sp. 0 0 l 0 0 0 0 0 Lemniscomys striatus 2 0 0 0 0 0 0 0 Lophuromys cf. sikapusi 0 3 0 0 0 0 0 0 Malacomys longipes 0 0 3 0 0 0 0 0 Mastomys natalensis 1 0 0 0 0 0 0 0 Mus musculus 0 2 0 0 0 0 0 0 Mus setulosus 4 | l 0 0 0 0 0 Oenomys hypoxanthus 8 3 0 0 0 0 0 0 Praomys jacksoni 2 l 0 0 0 0 0 0 Praomys misonnei 0 0 13 0 0 0 0 0 Sherman trap nights 471 - 1041 - 751 - 637 = Sherman captures 23 13 30 0 14 0 12 3 Trap success per night (%) 5.51 - 2.88 - 1.86 - 1.88 Diversity 7 8 7 0 2 0 1 1 H’ (pooled Sher+Trad/habitat) 0,921 0,456 0,178 0,217 Bonner zoologische Beitráge 56 Table 3. Means and ranges for the standard external measurements [mm] of the Mt. Kupe rodents. Measurements HB Oenomys hypoxanthus (n=11) 149 (110-168,5) Malacomys longipes (n=3) 137,7 (130-142) Lemniscomys striatus (n=2) 109-122 Mus setulosus (n=5) 74,9 (63,5-84) Hybomys cf. univitatus (n=1) 160 Grammomys poensis (n=3) 122 (116-133) Hylomyscus walterverheyenei 89,2 (n=28) (74-103) Hylomyscus sp. (n=1) 73 Praomys jacksoni (n=3) 120,3 (100-132) Praomys misonnei (n=10) 117,9 (100-197) Dendromus sp.(n=1) 33 Cricetomys emini (n=3) 347,7 (340-358) Lophuromys cf. sikapusi (n=2) 130-142 Deomys ferrugineus (n=2) 119-137 Mastomys natalensis (n=1) IES 163 TL HF E 177,3 31 18 (151-194) (23-36) (13-20) 161 35,3 20,3 (151-167) (32-35) (20-21) 111-115 25-26 15-16 33,5 14,8 117 (50-57,5) (14-16) (10,5-14) 93 28 IS 178,7 23 15,3 (171-189) (21-28) (15-16) 129,34 18,22 14,56 (112-150) (12-21) (13-16) 112 19 16 125 24,3 18 (113-139) (24-25) (16-19) 133 23,65 17,25 (115-160) (21,5-28) (15,5-19) 84 19 10 371 66 39,7 (350-391) (65-67) (39-40) 59-70 20 17 172-191,5 33-36 25 112 20 17 In total, 16 rodent species were captured. The most abun- dant species was Hylomyscus walterverheyeni (35, 80 % of captures), followed by Praomys misonnei (13,68 %) and Oenomys hypoxanthus (11,58 %). Only one shrew species (Crocidura cf. poensis) and two bat species (Epomops franqueti and Myonycteris torquata) were captured. The maximum diversity was found in the habitats under an- thropogenic influence and in lowland forest. Eight species were trapped in the former habitat type, with Oenomys hy- poxanthus and Mus setulosus being the most abundant. The absence of Hylomyscus cf. walterverheyeni from this habitat type is noteworthy, compared to its general abun- dance in the natural types. In the lowland forest Ay- lomyscus walterverheyeni and Praomys misonnei domi- nate the assemblages. The montane grassland yielded no captures. The sub-montane and the montane forest habi- tats’ diversity are low with two species each and are char- acterized both by the presence of Hylomycus walterve- heyeni. 3.2. Species records, characteristics and systematic notes In this species account we provide morphological and/or habitat characteristics for each species. Where confusion with another species is possible we also provide results from skull analysis and cyt b or 16S DNA sequences analysis. Order Rodentia Family Muridae Illiger, 1811 Oenomys hypoxanthus Pucheran, 1855 Six males and five females (one juvenile) of this species were recorded. Standard measurements are provided in Table 3. Our specimens displayed the characteristic red nose of the genus and an underside of white-orange col- ors ventrally, with a more intense orange-red part in the genital region. The back fur displays a mixture of long black guard and short red-yellow hairs. The tail is cov- ered with small black scales and is naked except for a few scattered, very short hairs. The Kupe specimens fall with- in the size range of specimens from Mount Cameroon (EISENTRAUT 1963) and S Nigeria (HAPPOLD 1987). Oeno- mys hypoxanthus shows appreciable geographic variation in fur color and body size; in Cameroon it was also cap- tured in Nyasoso, Buea, Musake, Malende and Mueli (11 specimens) (EISENTRAUT 1963). During this inventory of the Mt. Kupe small mammal fauna, this species was on- ly trapped at low altitude in Nyasoso village and in a fal- low area with Ageratum spp. These results are congruent with the observations of KINGDON (1997) and HAPPOLD (1987), according to which O. hypoxanthus commonly nests in fallow ground and village gardens. It is general- ly restricted to forest below 2000 m. 164 Christiane DENYS et al.: Small mammales of Mt. Kupe, Cameroon Malacomys longipes Milne-Edwards, 1877 Three specimens (two males and one sub-adult female) were trapped in the forest along a river at an altitude of 850 m. They have a hindfoot longer than 34 mm (Table 3). The specimens from Nyasoso have a shorter ear (20-2lmm) than those from Efulan (Cameroon) or Makokou/Belinga (Gabon) (25 mm), and their skull and body sizes are also smaller. These observed differences may have been a result of the low sample size of speci- mens captured from Nyasoso. MUSSER & CARLETON (2005) recognized the existence of three species within the genus Malacomys, among which only M. longipes is men- tioned in Central and South Cameroon. However, M. longipes and M. edwardsi occur both in South Nigeria (HAPPOLD 1987). EISENTRAUT (1963) found only seven specimens of M. longipes in Batoki, Malende and Isobi (Cameroon) and only one on Mt. Kupe at 900 m, but men- tioned a great variety of pelage colours in his sample. Sim- ilarly HAPPOLD (1987) also mentioned that Cameroon specimens of longipes show considerable variation, and some individuals are almost the size of the smaller ed- wardsi. However, the comparison of our cyt b sequences to those available in Genbank allows the definite identi- fication of the three specimens as M. longipes. Lemniscomys striatus Linnaeus, 1758 Two male striped rats were trapped in a maize field at Nya- soso village, 800 m asl. They fall within the size variabil- ity and color of L. striatus from Central African Repub- lic (CAR), Congo and Cameroon and have molecular affinities with representatives from the region (NICOLAS et al. 2008a). A few earlier specimens had been collected in Nyasoso, Mt. Manengouba (EISENTRAUT 1973) and the Mamfe region (SANDERSON 1940). Mus (Nannomys) setulosus Peters, 1876 EISENTRAUT (1957) attributed the Nyasoso Mus specimens to M. musculoides, but mentioned that they have a brow- ner pelage than specimens from Buea and Mt. Cameroon. Later, EISENTRAUT (1968) corrected the identification and attributed the M. musculoides specimens to M. setulosus, which we confirm here on morphological and molecular grounds. The type specimen of M. setulosus comes from Limbe (formerly Victoria), less than 100 km S of Mt. Kupe. We obtained six pygmy mice (two males and four females) attributed to Mus (Nannomys) setulosus, based on their dark pelage, rather large size (Table 3), and the presence of a second lobe on the lower M3 (KOUASSI et al. 2008). They have been molecularly typed to confirm the identification (KOUASSI et al. 2008); TL is somewhat shorter than HB. All six specimens were trapped in zones influenced by human activities and in lowland forest. One subadult female had 2+2 mammae, the other female was a juvenile. Mus musculus Linnaeus, 1758 Two M. musculus males were trapped by inhabitants in Nyasoso village. Cyt b analysis confirmed their identifi- cation. Hybomys cf. univittatus (Peters, 1876) One sub-adult male was recorded in lowland forest. Both skull morphology and cyt b analyses identified it as Hy- bomys (Table 4). EISENTRAUT (1968, 1973) described on- ly A. univittatus badius from Nyasoso and Mt. Cameroon. But according to the same author Hybomys u. badius and H. u. univitattus coexist in Mamfe region. MUSSER & CARLETON (2005) mentioned taxonomic problems con- cerning the Central African Hybomys species, but con- firmed three other forms in the Cameroon Highlands: A. badius on the Bakossi Highlands (Mt. Cameroon, Rumpi Hills), A. eisentrauti (described from Mt. Lefo, syn- onymized by MUSSER & CARLETON with H. badius) on the Bamenda Highlands, and H. basilii on the Island of Bioko. Further revision and discoveries of new specimens from both lowland and highland forests of the CVL pending, we attribute the Mt. Kupe specimen from lowland forest provisionally to the univitattus complex. Grammomys poensis (Eisentraut, 1965) Three specimens were caught in lowland forest and cul- tivated areas at Nyasoso village (Table 2). One of the two males was sexually active and the female was pregnant, carrying two embryos and displaying 0+2 mammae. Spec- imens were identified based on their very long tail (Table 3), ending with a small tuft, and by typical skull charac- teristics, notably the globular aspect of the braincase, mi- crodonty, as well as the existence of a stephanodont crest on the molar cusps and presence of a t7 on the upper M12 (see review in PETTER & TRANIER 1975). Comparison with G. gazellae (put in synonymy with G. macmillani by MUSSER & CARLETON 2005) specimens from CAR and with G. “rutilans” (a synonym of G. poensis) from Gabon and with the holotype from Fernando Poo allows assig- nation of our new Kupe specimens to G. poensis. Yet, they are slightly smaller than the holotype which is a young adult. Two specimens were sequenced (cyt b) but a revi- sion of the whole genus in West Africa is required before definite conclusions can be drawn. According to EISEN- TRAUT (1968) this species is absent from Mount Cameroon, but present in Mamfe (SANDERSON 1940) and on Bioko Island (EISENTRAUT 1965). Hylomyscus walterverheyeni Nicolas et al. 2008 Thirty-four specimens of this rodent were trapped on Mt. Kupe at different altitudes (Table 4). Among the 12 fe- males we found two juveniles, the other being adult with IT, DAA Bonner zoologische Beitráge 56 165 2+2 mammae. Three females were each carrying four to five embryos. Ten of the 20 males were sexually active. Our identifications were confirmed through molecular analyses (cyt b and 16S gene sequencing). According to our preliminary molecular analyses, specimens from Mt. Kupe are the sister clade of the newly described species H. walterverheyeni (NICOLAS et al. 2008a, Missoup 2009). This species was described from the region between the Sanaga River and the Oubangui and Congo rivers in Cameroon, Gabon, Central African Republic and Repub- lic of Congo. However, the authors stressed the importance of conducting additional sampling in Western Cameroon, Nigeria and the Dahomey Gap region in order to precise- ly define the western limit of its distributional range. Re- fined molecular analyses and multivariate morphometri- cal analysis of cranio-dental measurements are necessary to confirm whether the Kupe individuals represent a dis- tinct montane forest clade. Hylomyscus sp. One adult female of Aylomyscus was captured in the zone under anthropogenic influence and was molecularly sim- ilar to four specimens called “Hylomyscus taxon 2” by NICOLAS et al. (2006). These four specimens all came from the Korup National Park in western Cameroon. They un- doubtedly belong to a new species. Waiting for a gener- al revision of the genus and especially of the taxa found along the CVL, we provisionally call it “Hylomyscus sp.” This female had a mammae formula of 2+2. Mastomys natalensis (Smith, 1834) One adult male specimen was trapped in Nyasoso village. An analysis of the cyt b gene confirmed its identification as M. natalensis. Because of the high morphological vari- ability observed in this group and the low degree of spe- cific distinctiveness (DENYS et al. 2007), we did not com- pare this specimen with other specimens morphometrical- ly. Previously only one specimen of this species was trapped (EISENTRAUT 1968), and therefore it does not seem to be common in the CVL while it is abundant in West Africa (DENYS et al. 2005) and in Nigeria (HAPPOLD 1987). Praomys Thomas, 1915 Based on molecular analyses and the identification key of LECOMPTE et al. (2001) the Mt. Kupe specimens belong to two different species. EISENTRAUT (1968) only record- ed one Praomys species, P. morio, from Mt. Cameroon. Praomys jacksoni (de Winton, 1897) In our study this species is represented by one female, one male and one juvenile which were all trapped and brought to us by Nyasoso villagers (Table 4). The mammae for- mula was 1+2. This species is recognized by its well-de- veloped supra-orbital crests and the presence of a clear- ly separated t3 on MI (LECOMPTE et al. 2001). A compar- ison of our 16S and cyt b sequences with those published by NICOLAS et al. (2005) and by LECOMPTE et al. (2002) confirmed our identification. Praomys misonnei Van der Straeten & Dieterlen, 1987 Five females and eight males were trapped in the lowland forest (Table 2). Based on the absence of supra-orbital crests and their molar morphology they clearly belong to the so-called tullbergi complex. The females have a mam- mae formula of 1+2. A comparison of our 16S and cyt b sequences with those published by NICOLAS et al. (2005) and LECOMPTE et al. (2002) confirmed our identification. Subfamily Deomyinae Thomas, 1888 Deomys ferrugineus Thomas, 1888 Two specimens were captured at c. 1500 m asl on Mt. Kupe (Table 2). This is surprising since EISENTRAUT (1963) only obtained this species in the lowland forest, six specimens in Nyasoso and at “Lager V” (western part of Mt. Cameroon) at 600 m. Lophuromys cf. sikapusi (Temminck, 1853) We trapped only one male and one female; one badly dam- aged specimen (sex unknown) was brought to us by the villagers. All specimens were trapped in the village. They are characterized by a red belly and a short tail (Table 4). The female had 1+1 mammae. Recently, Lophuromys tax- onomy and systematics in Central, West and East Africa has been under debate. According to MUSSER & CARLETON (2005), the Lophuromys sikapusi complex may be repre- sented in S Cameroon by different species, following VER- HEYEN et al. (1997, 2000). Numerous sibling species are found within this complex. A molecular revision is nec- essary in order to precisely identify the specimens from Mt. Kupe. Family Nesomyidae Major, 1897 Subfamily Dendromurinae G. M. Allen, 1939 Dendromus sp. Only one adult male was captured in a crop field. Accord- ing to MUSSER & CARLETON (2005) and EISENTRAUT (1968, 1973), D. messorius in the Cameroon highlands is 166 Christiane DENYS et al.: Small mammales of Mt. Kupe, Cameroon known from Manengouba and Nyasoso, and from the holotype from Efulen. The species is a typical inhabitant of tropical lowlands and lower mountain slopes. In the same region, Eisentraut found Dendromus oreas at high- er altitudes (1850 m and 3000 m on Mt. Cameroon). MUSSER & CARLETON (2005) mentioned the high variabil- ity in D. oreas from West Central Africa. Our “new” spec- imen is smaller than D. messorius and D. oreas of the CVL. In size and skull morphology it is close to speci- mens in the Paris Museum from Belinga, Gabon, identi- fied as D. mystacalis. The specimen from Mt. Kupe is al- so smaller than D. messorius from CAR. Pending further comprehensive revision of the genus in West-central Africa, we leave this specimen as unidentified. Subfamily Cricetomyinae Roberts, 1951 Cricetomys ct. emini Wroughton, 1910 Four Cricetomys individuals (three males and one unde- termined) were brought to us by hunters. They were ob- tained by traditional traps placed in Nyasoso village and in the forest near the top of the mountain, at approximate- ly 1950 masl. EISENTRAUT (1968) described 11 specimens from Buea, Nyasoso, Kumba and Mt. Cameroon as C. gambianus poensis. MUSSER & CARLETON (2005) put poensis in synonymy with emini and recognize C. gam- bianus only in the sub-Saharan savannah belt. Our spec- imens fit within the external measurements range of vari- ability provided by EISENTRAUT (1968), except for the shorter tail length. According to OLAYEMI (2008) both C. gambianus and C. emini are present in South Nigeria. Fur- ther molecular work will help to refine the taxonomy with- in this genus. Table 4. External measurements [mm] of Mt. Kupe Crocidu- ra. HB TL HF E WwW mean 94,2 53,4 14,6 8,6 16,7 range 82-105 48-62 10-17 7-11 13-19 Table 5. External measurements [mm] of Mt. Kupe Chiropte- ra. Species W HB TL HF + claw FA Epomops 1 66 122 10 20352 80 Epomops 2 50,5 113 16 20 +2 TT Myonycteris 14 65 10 + 0,5 de Order Soricomorpha Gregory, 1910 Family Soricidae G. Fischer, 1814 Crocidura cf. poensis (Fraser, 1862) Five medium-sized shrews were captured in Sherman traps (Table 2). We recovered four males and one female with 0+2 mammae. At least eleven species of Crocidura are known from SW Cameroon (HUTTERER 2005). According to size and morphology our new specimens fit within the C. poensis complex (P. Barriere, pers. comm.). It should be noted that EISENTRAUT (1973) recorded 5 species of shrews, Sv/visorex megalura, Sylvisorex ollu- la, Crocidura nigeriae/poensis, C. attila, and C. hilde- gardeae from Nyasoso at 850 m. Order Chiroptera Blumenbach, 1779 We used mistnets to sample the bat population of a cave 950 m asl in the lowland forest of Mt. Kupe. Two fruit bat species were collected. Family Pteropodidae Gray, 1821 Epomops franqueti (Tomes, 1860) This species displays white basal ear tufts, and males al- so have white epaulets. We collected two adult females of this species (Table 5). This species was quite common- ly trapped by EISENTRAUT (1968) on Mt. Cameroon and at Lake Barombi Mbo, and he attributed these specimens to the subspecies franqueti, mentioning the considerable variability of the species over all of West Africa. Our spec- imens from Nyasoso are much larger than the average for his specimens (FA=57,4, TL=12, HB=91, HF=15; EISEN- TRAUT 1968) but are smaller than Nigerian specimens (FA=82-91; HAPPOLD 1987). Myonycteris torquata (Dobson, 1878) One male of the little collared fruit bat was trapped. This species has also been sampled by EISENTRAUT (1968) at “Lager V” and “Lager VI” of Mt. Cameroon and was list- ed in an unpublished report by DOWSETT-LEMAIRE & DowseTT (1998). Our specimen has a smaller forearm length (44 mm) than specimens from Mt. Cameroon (av- erage 57,4 mm; EISENTRAUT 1968) and Nigeria (HAPPOLD 1987) (Table 5). Bonner zoologische Beiträge 56 167 4. DISCUSSION 4.1. Rodent species diversity on Mt. Kupe compared to results of previous studies Our study confirms the presence of £. cf. sikapusi, L. stria- tus, M. setulosus, O. hypoxanthus, H. cf. univittatus and D. ferrugineus on Mt. Kupe (Table 6). The differences be- tween our species list and those of previous studies main- ly concern genera for which the taxonomy is under revi- sion: Praomys, Hylomyscus, Mus, Nannomys and Crice- tomys. Concerning the genus Praomys, we could not con- firm the presence of P. morio on Mt. Kupe, as suggested by EISENTRAUT (1968). This species was described from Mt. Cameroon where it ranges at altitudes up to 3000 m. The validity of P. morio was never assessed by molecu- lar data. Our results show that the specimens from Mt. Kupe correspond to P misonnei, not to P. morio. If future molecular studies confirm the validity of P morio it should be restricted to the high altitudes of Mt. Cameroon. We also confirmed the presence of P. jacksoni, as mentioned by EISENTRAUT (1973) and DOWSETT-LEMAIRE & DOWSETT (1998). These authors also cited Hylomyscus stella from Mt. Kupe. However, NICOLAS et al. (2008b) showed that H. stella is restricted to East Africa, and that the speci- mens from West Central Africa (from the Sanaga River to the Oubangui-Congo River) represent a new species, H. walterverheyeni. Our results suggest that this species could also be present on Mt. Kupe. More detailed molec- ular and morphometrical analyses, though, are necessary to confirm this conclusion. EISENTRAUT (1968) attributed Mus specimens from Nyasoso to M. setulosus, which we confirm here on morphological and molecular grounds. Concerning Cricetomys, DOWSETT-LEMAIRE & DOWSETT (1978) mentioned the presence of C. emini at 1500 m. However, molecular analyses in progress seem to indicate quite a large genetic variability. Our inventory adds three species to the fauna of the Mt. Kupe: Grammomys poensis, Mastomys natalensis, and Hylomyscus walterverheyeni (previously recorded under the name H. stella) (Table 6). Despite its presence on Mt. Cameroon, the genus Mastomys was not previously recorded on Mt. Kupe. Similarly, EISENTRAUT (1957, 1968) did not list Grammomys poensis but indicated that it was trapped by SANDERSON (1940) in the Mamfe region. We also confirm the presence of Malacomys longipes, Oenomys hypoxanthus, Deomys ferrugineus, and Mus se- tulosus in Nyasoso. We failed to collect Stochomys long- icaudatus (4 specimens collected in Nyasoso by Eisen- traut) and Hylomyscus aeta, both easily identifiable based on morphological characters. Stochomys longicaudatus is known to be present in fragmented populations through- out its distributional range, and may be locally abundant (KINGDON 1997). In the Kupe area this species may be rare or may even have disappeared since 1963. Like DOwsETT-LEMAIRE & DOWSETT (1998), we found the bat species Epomops franqueti and Myonycteris torqua- ta in a cave in the lowland forest of Mt. Kupe. EISENTRAUT (1973) and FEDDEN & McLEoD (1986) collected both species at the same altitude. They seem to be quite com- mon on Mt. Bakossi and Mt. Cameroon. The yield of shrews was poor as compared to previous studies (EISENTRAUT 1973) and does not allow any con- clusion. Our study indicates that the rodent fauna on Mt. Kupe has not changed much since the pioneer study of EISENTRAUT (1957). This may be a sign of the quality of conservation of this forest. It has also shown that the rodent diversity is quite high in comparison with other mountains of the CVL (Mt. Bakossi, Mt. Manengouba, Mt. Lefo, Mt. Go- tel, Mt. Cameroon — see Missoup et al. 2006). With 16 species, however, it is lower than on Mt. Oku (Missoup et al. 2006). This may be due to the fact that few inven- tories have been performed on these mountains. All moun- tains may harbour more species than are currently known. However, our results may also be biased by the common species because of the low number of individu- als captured despite our trapping efforts. All the possible habitats of the lowland and montane forest were investi- gated but our trapping success remained very low in all lines and in all seasons (Table 2). Trapping success was lower than that obtained for Mount Elgon montane for- est (7,7-10%) (Kityo & CLAUSNITZER 2001), Kilimand- jaro (4,5-36 %) (MULUNGU et al. 2008) or montane forests of the Albertine Rift mountains, SW Uganda (3,41-11,4 %) (KASANGAKI et al. 2003). In all these studies, the high- est trapping success was obtained in disturbed or degrad- ed habitats. 4.2. Community analysis It is generally assumed that the diversity of small mam- mals decreases with altitude, or at least changes with changes in vegetation type. Vegetation types may play a crucial role in structuring small mammal assemblages. In order to test if these hypotheses apply to Mt. Kupe we per- formed a correspondence analysis on presence-absence da- ta for rodents and shrews as a function of altitudinal veg- etation levels. The graph of axis Fl x F2 of the correspondence analy- sis displays a quite good separation between all habitats (Fig. 1). It is clear that zones under direct human impact are well characterized by a cohort of commensals (like Mastomys, Mus and Rattus), but also species which are frequent in secondary forest and crops (e.g. Lemniscomys, Lophuromys, Dendromus, P. jacksoni). The true forest habitats display a different diversity with marked differ- 168 Christiane DENYS et al.: Small mammales of Mt. Kupe, Cameroon Table 6. Comparisons of the faunal lists for Mt. Kupe based on different studies (x= presence). Species This study Eisentraut 1968, 1973 Dowsett-Lemaire Eisentraut & Dowsett 1998 1957 Crocidura poensis x X x x Crocidura attila x 3 x Crocidura sp. x Cricetomys gambianus x x C. emini x x Dendromus sp. x x Grammomys poensis x Lophuromys sikapusi x x x x Deomys ferruguineus x x x x Lemniscomys striatus x x x x Mastomys natalensis x Mus musculus X X Mus setulosus x x x Mus musculoides x x Oenomys hypoxanthus x x x x Praomys jacksoni X x Praomys morio x x Praomys misonnei X Hybomys cf. univitattus 2 x x x Hylomyscus walterverheyeni x ? Hylomyscus sp. x Hvlomyscus aeta x x Hvlomyscus stella? x Malacomys longipes x Stochomys longicaudatus x X Rattus rattus X x Micropterus pusillus x x Epomops franqueti x x Myonycteris torquata x x Nannonycteris veldkampi? x Lissonycteris angolensis X Roussettus aegyptiacus x Megaloglossus woermanni x x x Kerivoula smithii X x Hipposideros caffer x Hipposideros ruber x Hipposideros cyclops x Rhinolophys alcyone x Pipistrellus eisentrauti? x Tadarida thersites? pS ences between the lowland forests on the one hand, and the sub-montane and montane forests (1500-2000 m) on the other along the F2 axis. Hybomys, Hylomyscus sp., Malacomys, and P. misonnei are characteristic of the low- land forest and are true forest taxa also found in the low- land forests of Gabon (DUPLANTIER 1982; NICOLAS & CoLYN 2003). H. walterverheyeni and Deomys ferrugineus are the only species found at 1500 and 2000 m (upper sub- montane and montane forests) on Mt. Kupe. EISENTRAUT (1957) showed for Mt. Cameroon that Malacomys and De- omys are lowland forest taxa (< 900 m) and that only Praomys morio and Hylomyscus aeta reach the higher lev- els (up to 3100 m and 2000 m, respectively). Other species of Hvlomyscus are commonly trapped at high altitudes in East Africa: On Mt. Elgon H. denniae is found in the for- est and bamboo zones up to 2875 m and 3100 m (Kıryo no - °° °° °— © Bonner zoologische Beitrage 56 169 CFA (axis F1 x F2 : 80,90 %) 2,5 0,5 Anthropogenic M. natalensis Ps eee Dendromus sp. -- axis F2 (36,21 %) --> oO -1 O. hypoxanthus G. poensis L. striatus -1,5 L.cf. sikapusi -2 -2,5 -2,5 -2 -1,5 -1 -0,5 C. poensis cplx e M. setulosus M. longipes Hylomyscus Sp. H. cf. univittatus e P. misonnei @ Lowland Forest H. cf. walterverheyeni 4 Montane Forest 2000m ambianus 4 D. ferrugineus Sub-montane Forest 1500m 0 0,5 1 1,5 2 2; -- axis F1 (44,70 %) --> Fig. 1. Correspondence analysis scatter plot of axes | and 2 representing 80.9 % of the total variability based on presence-absen- ce data for 16 species in the four general habitat types surveyed on Mt. Kupe. & CLAUSNITZER 2001), while on Mount Rungwe A. arci- montanus is only found between 600 m and 2000 m (Car- LETON & STANLEY 2005). In West and East Africa Lo- phuromys roseveari, L. eisentrauti, L. flavopunctatus, Praomys jacksoni, P. morio, P. delectorum, and Colomys goslingi are found above 2000 m (DIETERLEN 1979; VER- HEYEN et al. 1997; Kityo & CLAUSNITZER 2001; MULUN- GU et al. 2008). In our study Lophuromys and P. jacksoni were only caught in zones under anthropogenic influence close to the vil- lage at around 800 m elevation. EISENTRAUT (1957) found Stochomys, Cricetomys, and Dasymys to be restricted to an altitude below 900 m, Oenomys, Mus, and Lophuromys reached 2100 m, and Dendromus oreas and Otomys bur- toni were found above 2000 m. On Mt. Kupe we trapped Deomys at 1500 m, while Cricetomys is known from the summit (2000 m and above). On Mt. Elgon Cricetomys is also present at high altitude (3500 m) (KiTyo & CLAUS- NITZER 2001). In Bwindi, SW Uganda, species such as Grammomys dryas, Hybomys sp., L. medicaudatus, rah- mi, woosnami, H. aeta, denniae, vulcanorum, Mus bufo, and 7. venustus live in forests above 1800 m (KASANGA- KI et al. 2003). In lower elevation forests H. stella, Praomys jacksoni, Praomys sp., S. longicaudatus, and T. kempi occur. Other taxa like Deomys, H. univittatus, O. hypoxanthus found on Mt. Kupe, live in a wide range of habitats and may be found on the edge of forests. Rodent species diversity in the area studied decreases dras- tically from lowland to sub-montane forest (Figure 1, Table 2) between 900 and 1500 m. This may correspond to the differences observed in vegetation composition. Both LETOUZEY (1985) and TCHIENGUE (2004) found floristic differences between the lowland forest and the montane forest, with some endemics occurring only above 1000m. CHEEK et al. (2004) noted a floristic change from lowland to sub-montane forest on Mt. Kupe at 800 m. The very low diversity observed at 1500 m and 2000 m cor- responds with the difference observed between a lower sub-montane forest (= very rich and diversified) and an upper sub-montane forest (= relatively poor) between 1400 m and 2000 m (CHEEK et al. 2004). In comparing species turnover and elevation in small mammals for different 170 Christiane Denys et al.: Small mammales of Mt. Kupe, Cameroon tropical mountain regions MENA & VASQUEZ-DOMINGUEZ (2005) found the highest turnover at intermediate altitude between 1000 m and 2500 m, but they did not attempt to define causes of such changes in species diversity. EISEN- TRAUT (1957, 1968) also described a change in vertebrate community composition on Mt. Cameroon: for amphib- ¡ans between 800 and 1000 m, for reptiles around 900 m, and for small mammals at 2100 m. Studies are on the way to verify the altitudinal effect hypothesis for the other mountains of the Bakossi and Bamenda Highlands of the CVL (Missovp et al., unpubl). 4.3. Conclusion The results of our small mammal survey suggest an ab- sence of rodent species endemic to the high altitude for- est of Mt. Kupe. Because recent work has revealed an un- suspectedly high cryptic diversity, molecular analyses are now necessary to validate this hypothesis. Our study has updated the Mt. Kupe faunal list with the addition of four species not recorded in previous studies. The Mt. Kupe diversity is relatively high when compared to other Cen- tral African forest areas. We present first evidence that the recently described HA. walterverheyeni occurs at 2000 m and is absent from habitats under impact by humans, and that Deomys ferrugineus can reach altitudes of 1500 m. On the other hand, P misonnei, Malacomys, and Hybomys are restricted to lowland forest below 900 m. We recov- ered almost all the true forest species previously collect- ed from Mt. Kupe, suggesting a good preservation of the lowland forest. This is of importance for drill conserva- tion because these primates have been encountered on Mt. Kupe from 700 m up to 2000m and they use all types of habitats on Mt. Kupe. Of the species previously mentioned for this area by Eisentraut we could not collect Stochomys longicaudatus and Hylomyscus aeta durmg this study. These two genera are inhabitants of the lowland forests of Central Africa. Little is known about their habitats and demography; their conservation status is “Lower Concern” (BOITANI 2007; SCHLITTER & VAN DER STRAETEN 2007). The absence of S. longicaudatus and H. aeta may suggest a low population size. Moreover, the low trapping success we obtained during various seasons may indicate a gen- eral decrease of rodent abundance. This ought to be checked in the future because 1t may indicate an over-ex- ploitation of forest rodents for bushmeat. We observed a change in the small mammal community composition between lowland and sub-montane forest, suggesting a turnover occurring between 1000 m and 1500 m. We also observed a change in the community struc- ture between areas under human impact and lowland forests, with different genera or species present and a rather good diversity in the undisturbed lowland forest. This stresses the importance of regular small mammal bio- diversity surveys in close relationship with local vegeta- tion associations on Mt. Kupe and more generally in hotspot zones to follow the forest condition. Acknowledgements. Special thanks are due to our drivers Gabriel Kogé and René Tsiguia. In Nyasoso village we grate- fully acknowledgement the help of : Mara and the women's cen- ter, Chiefs Ekindé and Bick Jack, the WWF collaborators Theophilus Ngwané, Messape Derrick, our ecoguides Gabriel Assuntong, Ewané Njune Charles, Nsang Félix, as well as the whole community. We are grateful to Dr M. Tchamba (WWF Yaoundé Cameroon) and Mr T. Okah (WWF Limbe), F. Riviere and X. Garde (IRD Yaoundé Cameroun) for all support received. This research was conducted under permit N° 41/MINRESI/B00/C00/C40 and an EGIDE grant attributed to Alain Didier Missoup by the SCAC Yaoundé (French Embassy) and MAE. We dedicate this work to the late Dr Daniel Lachaise who joined us in the field on Mt. Kupe in 2004 and 2005 and transmitted to us his knowledge of the CVL and his passion for evolution. The work was funded by a Grant from the “BQR Ray- onnant” of the MNHN Paris (2004-2006); Michel Veuille (MNHN, Department Systematics and Evolution) supported ac- tively this project. We thank Doyle Mc Key and N. Avenant for their attentive reading and suggestions on this manuscript. Zusammenfassung. Mehr als 50 Jahre nach M. Eisentrauts er- ster Bestandsaufnahme der Kleinsäuger-Fauna des Mount Kupe, SW Kamerun, unternahmen wir die zweite. Dabei kon- nten wir 19 Arten nachweisen: 16 Nager, zwei Fledermäuse, eine Spitzmaus. Für jede Art werden Daten zur Biologie, Verbreitung und Taxonomie zusammengefasst. Zwischen der Nager-Artenge- meinschaft des Tieflandes (< 1000 m) (inkl. Praomys misonnei) und der der höher gelegenen Wälder (1500-2000 m) (inkl. Hy- lomyscus cf. walterverheyeni als Erst-Nachweis für das Kameruner Bergland) bestehen Unterschiede, die wahrschein- lich mit den bekannten Veränderungen der Vegetation zwischen 1000 und 1500 m Meereshöhe zusammenhängen. In bewohn- ten und landwirtschaftlich genutzten Gebieten am Fuße des Mount Kupe konnten wir acht weitverbreitete Nager-Arten nach- weisen. 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Conservation of drill populations in Bakossiland, Cameroon: historical trends and current status. International Journal of Primatology 26: 759-773. WILSON, D. E. & REEDER, D. M. (eds.) 2005. Mammal Species of the World. A Taxonomic and Geographic Reference. 3rd edition. The Johns Hopkins University Press, Baltimore. Authors’ addresses: Prof. Christiane DENYS (correspon- ding author, e-mail: denys@mnhn.fr) and Violaine Nico- LAS, MNHN, Dept. Systématique et Evolution, UMR7205, 55, rue Buffon, 75005 Paris, France; Alain Didier Missoup and Prof. Charles Felix BILONG, Department of Animal Bi- ology and Physiology, Faculty of Sciences, University of Yaoundé I, P.O. Box 812, Yaoundé, Cameroon; Dr. Barthelemy TCHIENGUE and Dr. Gaston ACHOUNDONG, Herbier National du Cameroun, Institut de la Recherche Agricole pour le Développement, B.P. 1601, Yaounde, Cameroun; Dr. Atanga EKOBO, World Wild Fund for Na- ture, Coastal Forests Programme, Limbé, Cameroon; Dr. Dieudonné Massoma LEMBE, Département de Biologie des Organismes Animaux, Faculté des Sciences, Université de Douala, B.P. 24157, Douala, Cameroun. Received: 05.11.2008 Accepted: 19.11.2008 Bonner zoologische Beitráge Band 56 | Heft 3 Seiten 175-183 Bonn, September 2009 A new species of Surdisorex Thomas, 1906 (Mammalia, Soricidae) from western Kenya Julian KERBIS PETERHANS!2, William T. STANLEY2, Rainer HUTTERER3, Terrence C. DEMos* & Bernard AGWANDAS ! Roosevelt University, Chicago, U.S.A. 2 Division of Mammals, Field Museum of Natural History, Chicago, U.S.A. 3 Section of Mammals, Zoologisches Forschungsmuseum Alexander Koenig, Bonn, Germany 4 Department of Biology, The Graduate Center, City University of New York, New York, U.S.A. > Section of Mammals, National Museums of Kenya, Nairobi, Kenya Abstract. The genus Surdisorex represents Kenya’s only endemic genus of mammal. It has heretofore included two species from the highlands of central Kenya. Here we add a third species, also from Kenya, based on a specimen from the eri- caceous zone of the eastern slopes of Mt. Elgon. Although this species further aligns Mt. Elgon with the Kenya High- lands, the mammal fauna of Mt. Elgon illustrates a mixture of faunal origins and associations. Keywords. Surdisorex, mole shrews, endemism, fossorial mammals, Mt. Elgon. 1. INTRODUCTION In 1906, Thomas erected the genus Surdisorex for a high- ly specialized burrowing shrew, without external pinnae, from the Aberdare Mountains of central Kenya. Ten years later, a second species of the genus was named by HoL- LISTER (1916) from nearby Mt. Kenya. For over 100 years, the genus Surdisorex has been Kenya’s only endemic genus of mammal. Even with the acquisition of large se- ries of each taxon, the two species remain readily diag- nosable despite their geographic proximity. HOLLISTER (1918) stated that both species were confined to high al- titudes (above 9,000’). Their occurrence on mountain tops appears to account for their isolation and distinction, de- spite a geographic disjunction of less than 50 miles. In 1984, a specimen of the genus Surdisorex was picked up on a footpath in the ericaceous zone of the Kenyan slope of Mt. Elgon. We describe it as a third member of the genus Surdisorex. It is surprising that this distinctive animal had gone uncollected from this often-studied mountain until now. 2. MATERIALS AND METHODS The sole representative of the new species was found dead on the footpath by Dr. Henning Grossman at 3,150 m in the ericaceous zone of Mt. Elgon in December, 1984 and donated to Professor Kim M. Howell, Department of Zoology, University of Dar es Salaam who assigned it the field number KMH 3213. Professor Howell donated the specimen to the Field Museum of Natural History in 2005. The specimen was partially eviscerated with the loss of internal viscera and external reproductive organs, but we do not know how or when this occurred. It was kept in the Tanzanian collection, and stored in ethanol for some 20 years under the name *Myosorex sp.’ Exposure to light may have discolored the pelage. We compared the specimen with specimens in the Amer- ican Museum of Natural History (AMNH), The Natural History Museum, London (BMNH), Field Museum of Natural History, Chicago (FMNH), the National Museum of Natural History, Smithsonian Institution, Washington, DC (USNM), and the Zoologisches Forschungsmuseum Alexander Koenig, Bonn (ZFMK). The following exter- nal measurements were recorded for each specimen: to- tal length (TOL; tip of nose to end of caudal vertebrae), head and body length (HB; tip of nose to base of tail), tail length (TL; from base of tail to end of caudal vertebrae), hind foot length (HF; from heel to tip of claw), ear height (E; the longest dimension of the opening of the auditory meatus) and weight (WT). All measurements are in mm except weight (grams), and generally follow DEBLASE & MARTIN (1974). The cranial and dental variables measured are condylo-incisive length (CI), post-palatal length (PPL), length of entire upper toothrow (UTR), length of 176 Julian KERBIS PETERHANS et al.: A new species of Surdisorex Thomas, 1906 from western Kenya complex teeth in upper toothrow, 1.e. the distance from the anterior edge of the fourth upper premolar to the posteri- or edge of the third upper molar (P*-M?3), distance from anterior edge of first upper incisor to posterior edge of up- per canine (UNICUSP), width of third upper incisor (BW), width of canine (CW), length of third upper molar (M3L ), width of third upper molar (M3W), least interorbital width (10), bimaxillary width (MX), greatest width of the brain- case (GB), post-glenoid width (PGL), height of the cra- nial capsule (HCC), length of mandible including the in- cisor (md), height of coronoid process on dentary (cor), and length of lower toothrow (Itr). These variables are a subset of those utilized by HUTTERER & Kock (2002), Dip- PENAAR (1977), and CARRAWAY (1990) and are figured in STANLEY et al. (2005). The infraorbital bridge was meas- ured at its narrowest point; the structure was figured by MEESTER (1963). Digital callipers were used to measure skulls to the nearest 0.01 mm. Standard descriptive statistics (mean, range, and standard deviation) were calculated for each species. One-way analysis of variance (ANOVA) was used to determine whether there were significant differences in cranial char- acters between sexes in each species. Principal compo- nents were extracted from a variance-covariance matrix using the cranial variables converted to natural logarithms. All univariate and multivariate statistical analyses were conducted using Systat (version 11.00, 2004). 3. RESULTS External measurements indicate that S. polulus has a short- er head and body, but longer tail and hindfoot than S. no- rae (Table 1). Unfortunately, no external measurements were recorded from the specimen found on Mt. Elgon at the time of collection. The tail and hindfoot measurements taken from the preserved specimen indicate this specimen had a longer tail than either of the other two species, but the hindfoot length was intermediate between S. norae and S. polulus. In general, the specimen from Mt. Elgon falls between the means of 5. norae and $. polulus in various length meas- urements of the skull (Table 2). One-way Analysis of Vari- ance indicated there was no sexual dimorphism in cranial characters between sexes in either $. norae or S. polulus. The one exception was in 5. norae, where the males had a longer toothrow on the dentary than females (F = 5.14; P < 0.05). Based on the fact that this was the only char- acter exhibiting significant differences between sexes, and that this difference was found in only one species, the sex- es were combined in all other analyses. Table 1. External measurements (in mm) for samples of Sur- disorex, given as mean, standard deviation, range and sample si- ze. Measurements of S. norae and $. polulus taken from speci- mens upon capture, and those of S. schlitteri from the preser- ved specimen. Head & body Tail length Hindfoot External measurement Surdisorex norae 102.2 + 4.1 28.2 + 3.5 156; 1.2 95-110 21-33 14-17.5 n=19 n= 19 n= 19 Surdisorex polulus 94.8 + 3.1 28.5 #23 16:3=:0:5 89-100 2431 16-17 n=11 n=11 n=11 Surdisorex schlitteri = 28.6 16.1 Principal components analysis resulted in skull length measurements generally having positive and moderately high loadings with the first component. For example, the variable with the highest loading on PC 1 was the condy- lo-incisive length (CI). PC 2 was most heavily influenced by the length of the row of upper unicuspids (length from first upper incisor to the canine), which had positive load- ings. Equally influential on the second component (but with negative loadings), was the width of the third upper incisor (BW). The third component was most influenced by the width of the third upper molar, (MW) which ex- hibited negative values. The first three components ex- plained 40.1, 22.3 and 16.1% of the variation, respective- ly. A plot of the specimen scores on the first two compo- nents extracted is shown in Fig. 1. 3 2 < a 1 N SO N oO 0 a E SPECIES -1 e norae x polulus 2 + schlitteri -2 -1 0 1 2 PC 1 (40.1%) Fig. 1. Plot of the specimen scores on the first two principal components, based on cranıal measurements. Table 2. Comparison of selected cranial measurements (mm) for 3 species of Surdisorex, given as mean, + standard deviation, sample size and range. Character Condylo-incisive length (CI) Post-palatal length (PPL) Upper toothrow length (UTR) Length of P4-M3 (P4-M3) Length of upper unicuspids (UNICUSP) Greatest width of upper P (PW) Greatest width of upper canine (CW) Greatest length of upper M3 (M3L) Greatest width of upper M3 (M3W) Interorbital width (1O) Maxillary breadth (MX) Greatest breadth of cranium (GB) Post-glenoid width (PGL) Height of cranial capsule (HCC) Length of mandible (md) Coronoid height (cor) Lower toothrow length (Itr) Bonner zoologische Beiträge 56 S. schlitteri (n= 1) 25.15 11.28 4.45 0.72 0.77 0.87 12.90 16.41 S. norae 25.79 + 0.37 (n= 13) 25.24-26.75 11.38 + 0.24 (n= 10) 11.05-11.86 11.03 + 0.22 (n=21) 10.55-11.48 6.28+0.15 (n= 17) 6.02-6.59 4.75 + 0.13 (n= 17) 4.53-5.02 0.71 + 0.05 (n=17) 0.62-0.76 0.82 + 0.04 (n= 17) 0.72-0.87 1.58 + 0.07 (n= 17) 1.42-1.68 0.87 + 0.05 (n= 17) 0.80-0.97 5.30 + 0.14 (n = 16) 5.08-5.50 7.26 + 0.18 (n = 20) 6.93-7.56 13.17 + 0.29 (n= 13) 12.50-13.72 8.29 + 0.25 (n = 17) 7.86-8.76 6.62 +0.18 (n= 16) 6.36-7.03 16.58 + 0.36 (n = 20) 15.83-17.34 6.64 + 0.22 (n=21) 6.36-7.13 9.99 + 0.19 (n = 21) 9.65-10.42 S. polulus 24.68 + 0.30 (n= 8) 24.35-25.06 10.99 + 0.21 (n= 7) 10.59-11.25 10.43 + 0.20 (n= 18) 10.01-10.75 6.02 + 0.14 (n= 15) 5.81-6.29 4.40 + 0.14 (n=15) 4.13-4.61 0.70 + 0.05 (n= 15) 0.61-0.79 0.78 + 0.03 (n= 15) 0.72-0.84 1.49 + 0.08 (n = 14) 1.40-1.68 0.84 + 0.05 (n= 15) 0.76-0.94 5.22 +0.19 (n= 13) 4.82-5.54 6.84 + 0.16 12.04-12.68 7.98 + 0.18 (n= 11) 7.59-8.18 6.38 + 0.19 (n=8) 6.03-6.60 15.81 + 0.17 (n= 17) 15.58-16.28 6.41 + 0.17 (n= 18) 6.17-6.73 9.46 +0.19 (n= 18) 9.13-9.71 178 Julian KERBIS PETERHANS et al.: A new species of Surdisorex Thomas, 1906 from western Kenya 3.1. The Mt. Elgon Mole Shrew Surdisorex schlitteri n. sp. Holotype. FMNH 195069 (KMH 3213). Old adult, teeth heavily worn, sex indeterminate although suspected to be male since no teats are visible. Secondary number KMH 3213. Collected by H. Grossman in December 1985. Spec- imen prepared as fluid-preserved body with skull removed. Type locality. Kenya, eastern flank of Mt. Elgon, erica- ceous zone at 3,150 m. Etymology. The specific epithet honors Dr. Duane Schlitter, recognizing his important contributions to the knowledge of African small mammals and his personal and scientific engagement in Kenya. Diagnosis. Intermediate in cranio-dental dimensions compared with the other two species in the genus, S. polu- lus and S. norae. Braincase smaller and absolutely short- er. The more anterior position of the pair of anterior palatal foramina is unique within the genus, as is the position of the last palatal foramen, situated between the second pair of unicuspids. Infraorbital bridge narrow. The lingual ex- tension of the upper P* is longer than in the other Fig. 2. Fore- and hindfoot of the holotype specimen of Surdisorex schlitteri n. sp. (FMNH 195069); length of hindfoot (including claws) ıs 16.1 mm. Bonner zoologische Beitráge 56 179 Fig. 3. Skull of Surdisorex schlitteri n. sp. (FMNH 195069, holotype); cranium in lateral, dorsal and ventral view, mandible in lateral view. members of the genus, and almost in contact with the an- terior edge of M!. Lower P, narrow. Mandibular condyle heavy and straight-sided, forming a near-triangle without sinuosity. Description. The new species exhibits characters typical for the genus. The tail is short (28.6/84=34% of Head and Body). Fur is thick and woolly, length of single hairs on dorsum 9 mm. Color faded, presumably because of long- term preservation in alcohol, base of hairs dark brown, ter- minal 2 mm lighter. The foreclaws greatly enlarged, hind- claws moderately so (Fig. 2). Ear conch absent. Eyes re- duced and covered in membrane. Skull hexagonal, typi- cal of burrowing Soricidae (e.g. Myosorex, Anourosorex, Blarina) with particularly well-defined anterior facets of the braincase (Fig. 3). Interorbital region elongate (be- tween posterior processes of maxilla and anterior aspect of the braincase). Bridge across infraorbital foramen nar- row (1.87 mm). Nuchal crests well-developed, sagittal crest slight. Braincase small and short, as reflected in the measurement from the anteromost corner of the braincase to the corresponding occipital condyle and in the distance of the antero-lateral component of the hexagon-shaped braincase. Foramen magnum oval-shaped. Paired frontal foramina within interorbital basin. This basin provides a sinuous curve to the lateral profile of the skull. Three upper and two lower unicuspids, both unicuspid rows void of the vestigial unicuspids typical of Myosorex. Unicuspids broad, each with a medial accessory cusplet. Protocone of P* very prominent, forming a prominent shelf with parastyle. Upper M! and M? without metaloph. Low- er incisor denticulate, with chisel-like tip. Mental foramen beneath second cusp of lower Pz. Lower P, fairly narrow, exhibiting the inverted v-shape pattern typical of the Myosoricinae (HUTTERER et al. 2002). Lower M, with well-developed and deep entoconid basin (MEESTER 1963). MEESTER (1958) found that palatal foramina are useful in distinguishing closely related species of primitive African Soricidae (1.e. Myosorex). For the new taxon, the three principle anterior incisive foramina are tightly clustered in a triangle. The anterior pair is positioned slightly ante- rior to the medially positioned third member of this tri- ad, this pair being located in a plane even with the antero- most portion of the first unicuspid (Fig. 3). The third fora- men of this triad is positioned just below the anterior pair and is still situated within the anterior half of the first uni- cuspid. These foramina are followed by two reduced foramina, located approximately midway down the first unicuspid. A final foramen, the sixth, is located midway down from the second unicuspid. 4. COMPARISONS In most cranio-dental measurements, Surdisorex schlitteri is intermediate in size between S. norae, the largest mem- ber of the genus and $. polulus, the smallest (Table 2). This 180 Julian KERBIS PETERHANS et al.: A new species of Surdisorex Thomas, 1906 from western Kenya Table 3. Comparison of selected cranial measurements (mm) for 3 species of Surdisorex, given as mean, + standard deviation, range and sample size. Character S. norae S. polulus S. schlitteri Condylo-incisive length 23:8 0.3 24.6 + 0.3 25.1 25.2-26.7 23.8-25.1 (n=21) (n=15) Post-palatal length 11.4 + 0.2 10.9+0.3 113 11.0-11.9 10.5-11.2 (n= 18) (n=13) Upper toothrow length 11.0 + 0.2 10.4+0.2 10.5 10.6-11.5 10.0-10.7 (n=21) (n = 15) Length of P4-M3 6.3 +0.2 6.0+0.1 6.1 6.0-6.6 5.8-6.3 (n= 18) (n = 14) Length of upper unicuspid teeth 4.9+0.2 46+04 4.4 4.6-5.5 4.2-5.4 (n = 18) (n = 14) Greatest width of upper IP 0.7 +0.0 0.7+0.1 0.7 0.6-0.8 0.6-0.8 (n= 18) (n= 14) Greatest width of upper canine 0.8 + 0.0 0.8 + 0.0 0.8 0.7-0.9 0.7-0.8 (n= 18) (n = 14) Greatest length of upper M3 1.6+0.1 SEO 1.6 1.4-1.7 1.4-1.7 (n = 18) (n = 14) Greatest width of upper M? 0.9+0.1 0.8 + 0.0 0.9 0.8-1.0 0.8-0.9 (n= 18) (n= 14) Interorbital width 5.3+0.2 5.3: 0:2 5.4 5.1-5.6 5.0-5.6 (n = 21) (n= 15) Maxillary width 7.3 +0.2 6.9 + 0.2 7.0 7.0-7.6 6.5-7.2 (n=21) (n=15) Greatest width of cranium 13.2 + 0.3 12.6 + 0.3 129. 12.5-13.7 12.0-13.2 (n= 18) (n= 15) Post-glenoid width 8.3+0.2 8.0 + 0.2 8.1 7.9-8.9 7.6-8.5 (n= 21) (n=15) Height of cranial capsule 6.7 + 0.2 6.4 + 0.2 6.2 6.4-7.0 6.0-6.7 (n=21) (n = 14) Length of mandible 16.6 + 0.3 15.8 +0.1 16.4 16.1-17.3 15.5-16.0 (n=21) (n= 15) Coronoid height 6.6 + 0.2 6.4 + 0.2 6.7 6.3-7.0 6.0-6.8 (n= 21) (n= 15) Lower toothrow length 10.0 + 0.2 9.4+0.2 93 9.6-10.4 9.1-9.7 (n= 21) (n= 15) ur Bonner zoologische Beiträge 56 181 intermediate position is reflected in its position between the other two species along the primary axis of the first Principle Component (Fig. 1), an axis that illustrates size differences. Its infraorbital bridge is narrower (1.87) than either norae (1.97 and 2.26) or polulus (2.13 and 2.16). Its mandibular condyle is heavy and straight-sided and more or less triangular, without the sinuosity seen in the other two species. The new species can be distinguished from S. polulus by its generally larger size in most cranial and dental dimen- sions (CI, MX, GB, md, Itr: Table 2). In the new species, the central foramen of the anterior triad is located poste- rior to the triad pair whereas in S. polulus, it is situated slightly ahead, between the incisor and the first unicus- pid. The second pair of palatal foramina is in a compara- ble position between these species, but in the new species, the sixth and postero-most palatal foramen is located be- tween the second unicuspids instead of between the first and second unicuspids, as in $. polulus. From Surdisorex norae, the new species can be distin- guished by its generally smaller size in most dimensions: (CI, MX, GB, I1-P3 length, ltr; Table 2). In the new form, the antero-lateral dimension of the hexagonal edge of the braincase is 4.63, compared with 4.79 and 5.15 in S. no- rae. The distance from the antero-lateral corner of the braincase to the corresponding occipital condyle is 10.11 vs. 10.18 and 10.93 in S. norae. The lower P, is narrow- er than in S. norae. The shorter length of the upper unicuspids in the new species gives them a relatively broader apppearance than those of S. norae. This is especially clear in the first and third unicuspids. The angle in the posterior fovea of the upper P* is about 90° in the new form vs. 120° in Surdis- orex norae; therefore there is no medial gap between the P* and M! in the new form compared with norae. The anterior pair of palatal foramina in Surdisorex norae is positioned more posteriorly than in the new species, such that they form a straight line across the anterior half of the first unicuspid. The next pair is positioned compa- rable to the condition seen in the new species, whereas the final foramen is located midway down the third unicus- pid. A distinct sinus canal foramen (MEESTER 1963) 1s visible on the left side of the pterygoid region of the new species. It is less well-developed but evident on the right side. In other members of the genus, a canal replaces the foramen. A larger series is necessary to determine the significance of this feature. There is no east to west size cline among the species of Surdisorex, further demonstrating their mutual distinction. The largest species 1s Surdisorex norae from the Aberdare Mountains, which is located between the populations of S. schlitteri and S. polulus. Ecology of Surdisorex spp. The ecology of Surdisorex spp. 1s poorly known; to date there has only been a sin- gle published paper on the subject (DUNCAN & WRANG- HAM 1971). We have little data to report for S. schlitteri. Represented by a single specimen, $. schlitteri is only known from the ericaceous zone of Mt. Elgon. The Kenyan slope of Mt. Elgon was specifically surveyed for subterranean insectivores by DUNCAN & WRANGHAM (1971) without finding this species. However, these au- thors note that on Mt. Elgon, burrows of the golden mole (Chrysochloris stuhlmanni fosteri; species rank suggest- ed by THORN & KERBIS PETERHANS 2009) are “common in fairly open ground in the ericaceous zone up to 10,000 feet <3048m >". The golden mole is known from the eri- caceous zone of Mt. Elgon and the nearby Cherangani Hills. The study of DUNCAN & WRANGHAM (1971) showed that Chrysochloris fosteri feeds mainly on earthworms, supplemented by smaller oligochaetes. Earthworms were predominant in the diet of S. norae (Aberdares range) and S. polulus (Mt. Kenya). They concluded that Surdisorex and Chrysochloris share an earthworm diet and mole-like habits. In turn, this may have contributed to ecological dis- placement, mutually excluding these vermivores from overlapping ranges during times of climatic amelioration. Though this is a reasonable conclusion, the new record of S. schlitteri contradicts this view. The occurrence of S. schlitteri on Mt. Elgon also calls into question the recog- nition of Chrysochloris vs. Surdisorex burrows by visual recognition alone. Reporting on the Oxford University Expedition of 1969, DUNCAN & WRANGHAM (1971) documented $. norae and S. polulus in bamboo, mixed Podocarpus/bamboo and swamp. In the Aberdares, S. norae was collected at Kian- dongoro Gate (specimens in BMNH) along a swamp in moorland at 9,400’ (=2,865 m). Our 2006 efforts (JCK, TCD, BA) documented a single S. norae on the Aberdares at 2,700 m (=8860°) in secondary forest, the lowest vouchered record for the genus, but none at 2,200 m (=7220”). During our surveys, S. norae was extremely common at 3,300 m (10,830”) in Hagenia forest, Carex tussock grass and alpine shrubs and meadows. Heller col- lected $. norae on the summit of the Aberdares at 11,000” (specimens at USNM; Appendix). Our collated data (Ap- pendix) indicates that S. norae ranges between 2,700 m and 3,350 m. 182 Julian KERBIS PETERHANS et al.: A new species of Surdisorex Thomas, 1906 from western Kenya With a single exception, all records of 5. polulus originate from the west slope of Mt. Kenya. This may be an arte- fact of site access and collecting bias. The lowest record from Mt. Kenya (AMNH 82479) is at 9,000” (2,743 m) and the highest records are from 12,100” (= 3,690 m, US- NM records). Records of S. polulus from Mt. Kenya in the BMNH collected by the Oxford Expedition of 1969 originate near the Naro Moru Track at 10,500° (= 3,200 m). They derive from a range of bamboo associated habi- tats including mixed bamboo, bamboo/Podocarpus forest, bamboo mixed with 6” grass and swamp near bamboo (Oxford Expedition of 1969, specimens in BMNH). Lor- ING (1910) describes them as common in grassy openings within bamboo thickets at 10,000”. At other elevations (10,700” and 12,100”) they frequented Otomys sp. runs in tall marsh grass. 5. DISCUSSION This remarkable discovery suggests a relictual biotic as- sociation between Mt. Elgon and the nearby Kenyan High- lands to the east (The Aberdares and Mt. Kenya). The three known species of Surdisorex ssp. are confined to upper montane forest (uncommon), bamboo, and ericaceous and alpine zones above 2,700 m. The three taxa are mutually isolated due to their confinement to high montane habi- tats. Mt. Elgon has been well surveyed over the past 100 years. For the Ugandan slope, this includes: LOVERIDGE in 1933-1934 (ALLEN & LAWRENCE 1936), DAVENPORT et al. (1996); CLAUSNITZER & KiITYO (2001); CLAUSNITZER (2003); CLAUSNITZER et al. (2003). In addition to the re- port of DUNCAN & WRANGHAM (1971), historical expedi- tions to the Kenyan slope of Mt. Elgon to collect small mammals include those of KEMP in 1909 (reviewed in LOVERIDGE 1937), BAYER in 1914 (LONNBERG 1918), the Swedish Expedition of 1920 (GRANVIK 1924), and LOVERIDGE in 1934 (ALLEN € LAWRENCE 1936). The ab- sence of Surdisorex in prior reports is surprising. The genus is extremely common in bamboo/alpine zones of the Aberdares and Mt. Kenya. Perhaps its co-occurrence with Chrysochloris fosteri on Mt. Elgon limits its abun- dance. Mt. Elgon lies at the confluence of three major faunistic regions: the Albertine Rift and Congo basin to the west, the Kenya Highlands to the east, and the northern high- lands of Tanzanıa to the south. It houses representatives of two or three of these systems (summarized in CARLETON et al. 2006): Sylvisorex granti group, Colomys goslingi (Ruwenzoris, Albertine Rift); Crocidura elgonius, Cro- cidura fumosa, Surdisorex sp., Rhabdomys dilectus, and possibly Cricetomys ansorgei (Kenya Highlands); and per- haps from the third as well (northern highlands of Tanza- nia): Lophuromys aquilus (MUSSER & CARLETON 2005), Hylomyscus anselli (BisHop 1979). Mt. Elgon also hous- es a few strict endemics of its own (Otomys barbouri, Oto- mys jacksoni). Although our knowledge of the alpha tax- onomy of the small mammals in the region is incomplete, the central position of this extinct volcano reflects influ- ences from multiple montane biotic systems. Acknowledgements. We thank Dr. H. Grossman for collecting the first member of the new species and to K. Howell for pro- viding access to the specimen. We thank the Government of Kenya, including the Office of the President for the research per- mit (MOEST 13/001/30C/57) to the senior author. We thank the staff of the National Museums of Kenya (Dr. Idle Farah, Direc- tor) and Mr. Ogeto Mwebi (Head, Section of Osteology) for their continued support. For assistance in the field, we recognize, F. Awuor, D. Muruthi and B. Ochieng. For permission to work in Aberdare Mountain NP, we recognize Ms. Catherine Wambani, the Senior Warden, without whose assistance supplemental field work would not have been possible. D. Ogada, Father Boniface and the Gataragwa Catholic Parish provided key logistical sup- port adjacent to Aberdare Mountain NP. Finally, we recognize the staff of the following institutions for their assistance: AMNH (D. Lunde), BMNH (P. Jenkins, D. Hills), FMNH (M. Schulen- berg, J. Phelps), USNM (M. Carleton, L. Gordon, H. Kafka), and ZFMK (T. Wagner). We acknowledge the Barbara Brown Fund of the Field Museum for financial support. Rebecca Banasiak and Dirk Rohwedder helped with preparation of figures. Zusammenfassung. Surdisorex ist die einzige endemische Sáugetiergattung Kenias. Bisher enthielt die Gattung zwei Arten, die in den zentralen Hochlándern Kenias beheimatet sind. Hier fúgen wir eine dritte Art hinzu, ebenfalls aus Kenia, basierend auf einem Exemplar von der Ericaceen-Zone der Ostflanke des Mount Elgon. Diese neue Art, Surdisorex schlitteri n. sp., verbindet Mount Elgon zoogeografisch mit den Hochlandern Ke- nias, doch weist die Sáugetierfauna des Mount Elgon auch eine spezielle Mischung faunistischer Einflússe auf. APPENDIX Specimens examined. Surdisorex norae (THOMAS, 1906): all records from Aberdare Mountain NP, Kenya. AMNH 187262, 8 mi W of Bellevue (female): BMNH 6.7.8.1 (type), east side, near Nyeri, 9500”, old female; BMNH 10.5.3.1, 11,000” (female); BMNH 1974.653-74.657, Kiandongoro Gate, 9,400” (3 females, 2 males); FMNH 190257-190266, FMNH 190623-190626, 28.5 km W & 4.9 km N of Nyeri, 3,100 m (6 males, 8 females), FMNH 190622, 3.8 km W & 2.5 km S of Gatarakwa, 2,700 m (male); USNM 165513-166515, (1 female, 2 males), 182581-182586, summit, 11,000” (2 females, 4 males), USNM 589811-589813, USNM 589815-5898 18, Fishing Lodge, 9,000°-9,500° (1 female, 6 males). Bonner zoologische Beitráge 56 183 Surdisorex polulus (HOLLISTER, 1916): all but AMNH record, from W slope, Mt Kenya, Kenya. AMNH 82479, E slope, 9,000” (female); BMNH 1974.646-1974.652, Naro Moru Track, 10,500” (3 females, 2 males, 2 ?); FMNH 43845=USNM 163980, 12100” (female), FMNH 43846=USNM 163983, 12100” (male); USNM 163975- 163979 (4 males, 1 female), USNM 163981-163982 (2 fe- males), USNM 163984 (male), USNM 163986-163991 (3 males, 3 females), USNM 163993-163994 (male, female), USNM 163997-163998 (male, female), USNM 164000 (female), USNM 164003-164004 (2 females), USNM 165509 (male), USNM 165511 (female), 9000”-12100”, USNM 589820, Naro Moru Gate, 10000” (male); ZFMK 2003.702 (female), Mt Kenya N.P., 2950 m, Chagoria trail. REFERENCES CAMERON, A, (1997): Small mammal community composition on Mt. Elgon, Uganda with reference to habitat modification by humans. Honours Dissertation for BSc Tropical Environ- mental Science. Aberdeen University. Unpublished (not seen). CARLETON, M. D., KERBIS PETERHANS, J. C. & STANLEY, W. T. (2006): Review of the Hylomyscus denniae group (Rodentia: Muridae) in eastern Africa, with comments on the generic al- location of Epimys endorobae Heller. Proceedings of the Bi- ological Society of Washington 119: 293-325, CARRAWAY, L. N. (1990): A morphologic and morphometric analysis of the “Sorex vagrans species complex” in the Pa- cific coast region. Special Publication, Museum of Texas Tech University 32: 1-76. CLAUSNITZER, V. (2001): Rodents of the afro-alpine zone of Mt. Elgon. Pp. 427-443 in: DENYS, C., GRANJON, L. & POULET, A. (eds.): African Small Mammals. Paris, IRD editions. CLAUSNITZER, V. & KITYO, R. (2001): Altitudinal distribution of rodents (Muridae and Gliridae) on Mt. Elgon, Uganda. Trop- ical Zoology 14: 95-118. CLAUSNITZER, V., CHURCHFIELD, S. & HUTTERER, R. (2003): Habitat occurrence and feeding ecology of Crocidura montis and Lophuromys flavopunctatus on Mt. Elgon, Uganda. Afrıcan Journal of Ecology 41: 1-8. DAVENPORT, T., HOWARD, P., DICKENSON, C. (1996): Mount El- gon Biodiversity Report. Republic of Uganda Forest Depart- ment, P.O. Box 1752, Kampala, Uganda. DEBLASE, A. F. & MARTIN, R. E. (1974): A Manual of Mammal- ogy with Keys to the Families of the World. W. C. Brown Company Publishers, Dubuque, Iowa. DIPPENAAR, N. J. (1977): Variation in Crocidura mariquensis (A. Smith, 1844) in southern Africa, Part 1 (Mammalia: Sorici- dae). Annals of the Transvaal Museum 30: 163-206. DUNCAN, P. & WRANGHAM, R.W. (1971): On the ecology and distribution of subterranean insectivores in Kenya. Journal of Zoology 164: 149-163 GRANVIK, H. (1924): Mammals from the eastern slopes of Mount Elgon, Kenya Colony. Lunds Universitets Ärsskrift XXI 3: 432. le) HOLLISTER, N. (1916): Descriptions of a new genus and eight new species and subspecies of African Mammals. Smithson- ian Miscellaneous Collections 66 (1): 1-8. HOLLISTER, N. (1918): East African mammals in the United States National Museum. Part I. Insectivora, Chiroptera, and Carnivora. Bulletin of the United States National Museum 99: 1-194, 55 pls. HUTTERER, R., BARRIERE, P. & COLYN, M. (2002): A new myo- soricine shrew from the Congo Basin referable to the forgot- ten genus Congosorex (Mammalia: Soricidae). Bulletin de l'Institut Royal des Sciences Naturelles de Belgique, Biolo- gie 71, Suppl.: 7-16. HUTTERER, R. & Kock, D. (2002): Recent and ancient records of shrews from Syria, with notes on Crocidura katinka Bate, 1937 (Mammalia: Soricidae). Bonner Zoologische Beiträge 50: 249-258. LONNBERG, E. (1918): Mammiferes recuellis dans la region du Mont Elgon par le Dr. Bayer, en 1914. Revue Zoologie Afri- caine 5: 172-192. LORING, J.A. 1910. Appendix C, p. 549. In: African Game Trails, T. Roosevelt. Charles Scribners’ Sons, New York. MEESTER, J. (1958): Variation in the shrew genus Myosorex in southern Africa. Journal of Mammalogy 39: 325-329. MEESTER, J. (1963): A systematic revision of the shrew genus Crocidura in southern Africa. Transvaal Museum Memoir 13: 1-125. STANLEY, W. T., ROGERS, M. A. & HUTTERER, R. (2005): A new species of Congosorex from the Eastern Arc Mountains (Tan- zania), with significant biogeographical implications. Journal of Zoology 265: 269-280. THOMAS, O. (1906): Two new genera of small mammals discov- ered by Mrs. Holms-Tarn in British East Africa. Annals and Magazine of Natural History 7 (18): 222-226. THORN, E. & KERBIS PETERHANS, J. (2009): Small mammals of Uganda. Bonner zoologische Monographien 55: 1-164. Author’s addresses: Dr. Julian C. KERBIS PETERHANS, jkerbis@fieldmuseum.org, College of Professional Stud- ies, 430 S. Michigan Ave., Chicago, IL 60605, U.S.A.; William T. STANLEY, Division of Mammals, Field Muse- um of Natural History, Chicago, Illinois, 60605-2496, U.S.A.; Dr. Rainer HUTTERER, Zoologisches Forschungsmuseum Alexander Koenig, Adenauerallee 160, D-53113 Bonn, Germany; Terrence C. DEMOS, De- partment of Biology, The Graduate Center, City Univer- sity of New York, New York, U.S.A.; Bernard AGWANDA, Mammal Section, National Museums of Kenya, P.O.Box 40658, Nairobi, Kenya. Received: 16.05.2008 Accepted: 20.03.2009 Bonner zoologische Beiträge Band 56 | Heft 3 Seiten 185-200 Bonn, September 2009 Climbing mice of the genus Dendromus (Nesomyidae, Dendromurinae) in Sudan and Ethiopia, with the description of a new species Fritz DIETERLEN Staatliches Museum fiir Naturkunde, Stuttgart, Germany Abstract. The species of Dendromus occurring in Sudan and Ethiopia are revised. The forms of climbing mice referred to Dendromus mystacalis (Heuglin, 1863) from Sudan and other parts of Africa are compared with the typical popula- tion from Ethiopia. Ethiopian D. mystacalis differ from the other African populations in morphology and habitat. On the other hand, all populations from outside Ethiopia correspond in size and appearance and live in similar habitats. It is sug- gested that the lowland populations represent a different species. A new species, Dendromus ruppi, is described from the Imatong Mts., Sudan. It is a relatively large montane form, which was isolated during climatic changes in a Pleistocene postpluvial phase, and is probably endemic to these mountains. Keywords. Sudan, Ethiopia, rodents, Dendromus, morphology, ecology, altitudinal distribution, habitat, new species. 1. INTRODUCTION The genus Dendromus (African Climbing Mice) contains at least 12 species distributed throughout sub-Saharan Africa. Most species occur in S, C and E Africa (Muss- ER & CARLETON 2005). Typical habitats are long grassland, bracken, dense scrub, grassy wetland, and subalpine or alpine vegetation. Species are characterised by small size, soft brown or red- dish-brown pelage and a long thin tail (100-160% of head and body length, depending on the species) and feet spe- cialised for climbing. Several species have a black mid- dorsal stripe, and one has three black stripes. In some species the stripe is barely visible or even absent. Most species of Dendromus climb with the aid of the long digits of the forefeet and hind feet. The opposable digit 5 of the hindfoot which can be opposed to contact digit 1 and the prehensile tail are especially important for climb- ing and balancing on twigs and grass stems. Some species have a nail rather than a claw on digit 5 of the hindfoot, and tend to be more terrestrial than others (ZIEKUR 2006). The mammary formula is 2 — 2 = 8. The skull is small with a round braincase, a narrow ros- trum, and a narrow and vertical zygomatic plate carrying a distinct masseteric knob at its lower anterior corner. In- cisive foramina extend to the middle row of the upper M1. Upper incisors are small, orthodont, each with a single lon- gitudinal groove. Cheek teeth are rather narrow; M2 is about half the size of M1; M3 is minute; cusps of M1 are biserial (two cusps in each row), with additional small lin- gual cusps on a middle lamella of M1; on M2 additional cusps are expressed less or absent. The morphology of the skull and the dentition are very uniform in the genus but the pelage shows great variabiliy in colour and marking. Differences in skull length usually are very small, and measurements often overlap broadly. Classification of the species is therefore often based on rather few details (HEIM DE BALSAC & LAMOTTE 1958; DIETERLEN 1971). The distributional limits of many of the currently recog- nized 12 species (including 44 synonyms) of Dendromus are unresolved, and karyological information is missing for most species. The genus therefore is in need of fur- ther study and revision. As concerns systematics, it is one of the most difficult genera of African rodents. Earlier, the Dendromurinae were classified as a subfamily of the Muri- dae (e. g. THOMAS 1896; SIMPSON 1945; MUSSER & CAR- LETON 1993). CARLETON & MUSSER (1984) already ques- tioned this systematic position. Results of DNA hybridi- sation and mitochondrial and nuclear gene sequence stud- ies finally supplied the basis for grouping the Dendromuri- nae together with five further subfamilies in the family Ne- somyidae within Muroidea (MUSSER & CARLETON 2005). In this study, new specimens from the Imatong Mountains in Southern Sudan are described as a new species. The sample is compared with the related Dendromus mysta- calis from Ethiopia and with several forms from East and Central Africa currently assigned to that species. 186 Fritz DIETERLEN: Climbing mice of the genus Dendromus in Sudan and Ethopia 2. MATERIAL AND METHODS The present paper continues my revisionary studies of the rodent fauna of the Sudanese Republic (DIETERLEN & Rupp 1978; HUTTERER € DIETERLEN 1981; DIETERLEN & NIKOLAUS 1985). The research is mainly based on collec- tions obtained by Hans Rupp (71979) and Gerhard Niko- laus in Ethiopia and Sudan between 1971 and 1983. Al- most all the rodent material is housed in the Staatliches Museum für Naturkunde, Stuttgart, Germany (SMNS). Comparative material was studied in the following mu- seum collections: Museum of Comparative Zoology (MCZ), Harvard University, Cambridge (Mass. ), U.S.A.; Zoological Museum of the University of Copenhagen (ZMUC), Denmark; British Museum (Natural History), London, U.K.; Zoologisches Forschungsmuseum Alexan- der Koenig, Bonn (ZFMK), Germany. In addition to the relevant literature, the author’s fieldwork and taxonomic studies of several species of Dendromus in the Congolese Kivu Highland (Albertine Rift Valley) served for compar- ison (DIETERLEN 1969, 1971, 1976). 3. RESULTS 3.1. Species of Dendromus in the Sudan Republic Only two species were recorded from the territory of the Sudan Republic so far: D. mystacalis by SETZER (1956), and D. messorius by MUSSER & CARLETON (2005). A record of D. mesomelas subspec. (SETZER 1956) was based on an incorrect determination. Most of the 17 specimens of D. mystacalis preserved in museum collections were collected by J. S. OWEN and H. HOOGSTRAAL. A third (new) species, to be described below, was collected by J. S. OWEN, H. Rupp, and G. NIKOLAUS. Dendromus mystacalis lineatus Heller, 1911 Material from South Sudan (17). All nine localities are in the Equatoria Province, seven of them E of the Bahr el Jebel (White Nile) and S of Torit (Fig. 1): Issore (40 miles S of Torit), ZMUC 1753; Katire, ZMUC 14051, 14125; te Mo i 1 Ed \ & => E - = a IX = u on , elle on N adi A. E | ee | e x he En Lota ao | ER N + Lad TOO et E BO - «al E eae Mongalla a SER don SR $ y | oO. N en 9-7 | a \ \ “i IL ‘ 25 AL 3 £uri B- A a af x Gworo-e ESTE .oK! oeta = SORT Y ‘Juba, Nigeangalas i Da a vs 2 one ÑO E ee ES > : ae = Nefirangole a | o A » E ie a ‘Torit A Si 2 ea SE JGumbiris, , Se \ Na äshote er Ae oe Loko 4 Ye Fare | page, sok 0 o RER y Negtarpeto 20 Bra Lomein eee) KENYA — z Ss _Thoto Hy ge AS RE 10 Marada A, | ee Al Y ER y Be ‘ o Y | | paso Ne Langa Paimol a wege a O Rhino a P ag ed ER Arız°‘ Bs Muir! Gala PF y Adilan ang Wr \ Oho Ho: NER Moroto, Se re: War Rubik 5 -, Fig. 1. ven collecting localities, the easternmost of which, Gilo, is also the type locality of Dendromus ruppi n. sp. / ‘SS ree pkangole "Asofoto 2 a AS Distribution of Dendromus mystacalis lineatus in southern Sudan and northern Uganda. The region S of Torit shows se- Bonner zoologische Beitráge 56 187 Laboni (45 miles S of Torit), ZMUC 1852; Loa (20 miles N of Nimule), MCZ 44790; Lokwi, FMNH 79936 and four further specimens; Magwe, ZMUC 14122; Obbo, FMNH67071, 67078, 98957, USNM 67072, 67073 (not seen). Two localities from W of the Bahr el Jebel: Lado (N of Juba, 5°06’), FMNH 43482, 43483; Yei, FMNH 85382 (possibly D. messorius). Remarks. The following notes refer only to populations outside of Ethiopia, from where the ‘typical’ species was described by HEUGLIN (1863). The description of the species by BOHMANN (1942) was essentially correct, e. g. greatest length of skull GRLE) 19-21 mm, upper molar row (M1-M3) 3.0-3.2 mm; D 5 on hind foot always with a claw. The geographical form closest to Sudan is D. mystacalis lineatus, described from Rhino Camp, on the White Nile in N Uganda. According to BOHMANN (1942) its distribu- tion extends over S Sudan, N and W Uganda and from there to the Kivu Highland. For the Ituri region, this dis- tribution was confirmed by HATT (1940), and for the Kivu region by DIETERLEN (1971) and VERSCHUREN et al. (1983). The form D. acraeus Wroughton, 1909 (type lo- cality: Mt. Elgon, Kenya) is problematic. Studies of D. mystacalis from Kenya and Uganda by DIETERLEN (un- publ.) show that acraeus (WROUGHTON 1909) can hardly be distinguished from /ineatus. BOHMANN (1942) considered acraeus as a distinct form but included it in the species D. ruddi. WROUGHTON (1909) wrote: “Typus acraeus and Typus ruddi belong to a sam- ple in which acraeus represents the extreme with dark dor- sal stripe, and ruddi the extreme without dorsal stripe.” Recently ruddi was included as a synonym in D. messo- rius (MUSSER & CARLETON 2005), a species mostly with- out a dorsal stripe. The type series (USNM) of D. mysta- calis lineatus from Rhino Camp includes a specimen with- out a dorsal stripe, and in samples of acraeus it may al- so be missing, 1.e. it seems that in both forms specimens without a dorsal stripe occur. In several other species of Dendromus the expression of the dorsal stripe can vary, from a complete stripe to no stripe at all. Because the ranges of D. mystacalis and D. messorius overlap, mis- identifications may occur. Description of D. m. lineatus. Dorsally brownish or red- dish-brown. The hairs of the ventral side are white or whitish down to the root. The apical parts have sometimes a yellowish tinge. SETZER’S (1956) description of the dor- Fig. 2. Three skins of Dendromus mystacalis lineatus (SMNS 14737, 14738, 14736) in dorsal and ventral view. The dark mid- dorsal stripe is mostly complete, and the underparts are whitish. 188 Fritz DIETERLEN: Climbing mice of the genus Dendromus in Sudan and Ethopia Table 1. Table 1. Comparison of body and skull measurements of adult D. mystacalis from different regions in NE, E, and S Afri- ca: Ethiopia and Sudan (DIETERLEN unpubl.); Kivu, Congo (DIETERLEN 1971); E Africa (Kenya und Uganda, localities Kaimosi, Sirgoit and Rhino Camp, forms w/ytei, ruddi, acraeus, lineatus) after HOLLISTER (1919); South Africa after DE GRAAF (1981): Africa (without Ethiopia) complete data of specimens given by BOHMANN (1942) under the name D. pumilio (now a synonym of D. mystacalis). HOLLISTER (1919) used as skull length only the condylo-occipital-measure (* in Table 1) which is almost exactly 10% shorter than the greatest skull length (occipito-nasal-length), here correspondingly converted. Skull measurements: GRLE, greatest length; M1-M3, length of upper molar row; INT, interorbital breadth; BRC-W, greatest width of braincase. RTL = rela- tive tail length. Region HB T RTL HF EL WT GRLE M1-M3 INT BRC-W Ethiopia 66.4 93.4 141% 19.3 13.7 — 21.32 3.35 3.23 9.79 58-76 84-108 16-20 11-18 20.3-22.6 3.13-3.60 3.09-3.40 9.37-10.0 n 10 n 10 n 10 n 9 n 6 n 7 n 6 n5 Sudan 60 82 137% 172 11.9 = 19.6 3.14 = _ 53-68 72-9] 16-18 11-14 18.7-20.7 2.843.31 nl pd n 10 n7 ns n4 DR 63.6 11.3 122% 16.5 12.8 8.1 20.0 3415 2.8 9.25 Congo 53-73 63-83 16-18.5 4-12 18.8-21.3 3.0-3.5 2.6-3.4 8.9-9.6 Kivu n 18 n 18 n 18 n 18 nll n 11 nel nll East Africa 63.7 82.4 129% = = = 19,25% 3.03 2.9 9.17 55-70 71-100 17.5-21.1 2.8-3.3 2.5-3.3 8.4-9.8 n 29 n 29 n 29 n 28 n 29 n26 South 63.5 85 134% 16.5 14 8.2 _ — - - Africa 54-80 74-103 15-20 12-16 6-14 n 34 n 34 n 33 n 27 n 17 Africa _ _ = = = = 19.4 3.10 2.87 9.33 17.6-21.6 2.8-3.4 2.5-3.5 8.7-10.0 n 47 n 48 n 71 n 66 sal stripe of eight specimens from South Sudan is correct: „Ihe type specimen of D. p. lineatus has a pronounced mid-dorsal black stripe. None of the specimens in the pres- ent series shows this intense black stripe, but all agree with the remainder of the type series of lineatus in showing on- ly a faint suggestion of this marking. It is quite apparent that Heller selected the most strikingly marked specimen, rather than an average one, to name as the type”. Most skins of the 17 D. mystacalis from South Sudan have the mid-dorsal stripe poorly developed and thus confirm SET- ZER’S description (Fig. 2). In three specimens the stripe is absent. In samples of Ugandan /ineatus some specimens lack a dorsal stripe, but the dorsal colouration is exactly as in typical specimens. Body and skull measurements. Measurements of body and skull (Table 1) show that the Sudanese specimens agree very well in size with other populations of D. m. lin- eatus and also with other forms of D. mystacalis. With the exception of the Ethiopian sample, all (so-called) mysta- calis agree very well in size, appearance and characteris- tics of skull and dentition (Figs. 3, 4; see also DIETERLEN 1971; DIETERLEN & Rupp 1978). Altitudinal distribution. D. mystacalis (except for the Ethiopian form) can be characterized as an inhabitant of the Afrotropical moist savannah, preferring relative open grassy (dry and moist) biotopes. The altitudinal range of their habitats in S Sudan and N Uganda varies between 500 and 1000 m; the northernmost occurrence seems to be limited roughly by the 1400 mm-isohyete of annual pre- cipitation. KINGDON (1974) wrote “It is a low altitude species and is not found much above 2000 m”. This may be true for many regions, but not everywhere. In Ethiopia (typical region of D. mystacalis) the distribution ranges from 1200 to 3800 m, predominantly between 2000 and 3000 m NE and E Congo: Garamba NP 700-1000 m (MISONNE 1963; VERHEYEN & VERSCHUREN 1966): Is- hango, Virunga NP 950 m (VERHEYEN al. 1983); Lwiro (Kivu Highland) ca. 1500-2000 m (DIETERLEN 1971, 1976). Kenya: Taita Hills ca. 1800 m, Kaimosi ca. 1800 m (HOLLISTER 1919). Tanzania: Oldeani ca. 2000-3000 m (DIETERLEN unpubl.). Malawi: Nyika Plateau ca. 2150 m (ANSELL & DOWSETT 1988). Southern Africa (incl. Zim- babwe, Zambia, Mozambique): DE GRAAF (1981; SKIN- NER & SMITHERS 1990). Bonner zoologische Beiträge 56 189 Fig. 3. Skull of a large adult Dendromus mystacalis lineatus (SMNS 14748; greatest length of skull 21.17 mm) from the Ki- vu area, D. R. Congo. Note the rounded braincase, strong inter- orbital region, almost squarish zygomatic arches, and long in- cisive foramina. Fig. 4. Upper right molars of Dendromus mystacalis lineatus (SMNS 14750). Note the large M1 with biserial cusps and a lon- gitudinal valley, separating the labial t3, t6, t9 from the lingual “t0”, 12, t5, and t8. A “t4” is lingually attached to t5; the cusp pattern on M2 is still visible but heavily worn; M3 very small, as typical for the genus. Habitat. In Ethiopia the species is “nearly always asso- ciated with long grass and bushes” (YALDEN et al. 1976; Rupp 1980). NE Congo, Garamba NP: dry savannah, but found there in moist habitats around swamps etc. (VER- HEYEN & VERSCHUREN 1966). Congo (Kivu Highland): moist savannah, but only in relative dry grassy habitat mixed with bush vegetation, possibly due to competition with two other species of Dendromus (DIETERLEN 1971). N Kivu: «biotope caractéristique, végétation élevée, broussailleuse, que l’on trouve sur les bords des marais nombreux quí occupent les fonds de toutes les vallées» (MISONNE 1963). E Africa: “D. mystacalis... has adapted to cultivation quite readily ...” (KINGDON 1974). South- ern Africa: grassland associated with rank vegetation, es- pecially stands of high coarse grasses such as Hyparrhe- nia sp. at 1-2 m height (DE GRAAF (1981; SKINNER & SMITHERS 1990). Nest. “This is the species to which the only positive tree- dwelling records are attached, but the habitats of the dif- ferent forms and apparently of the same form in different locations, are variable” (ROSEVEAR 1969, on D. mysta- calis). For East Africa KINGDON (1974) noted, ”.. it ıs not unusual to find nests in garden shrubs, banana trees and banana bunches, in sweet potato vines, pineapples, palms and in thatched roofs. Their nests may be three metres or more from the ground but are generally lower down in thick herbaceous vegetation.” OWEN (1953) provided the following information on nest sites for Sudanese records of D. mystacalis: “in tree, in banana garden, under stone, swamp near river, tree near river.” The species is certain- ly more common than the small number of specimens in museum collections may suggest, probably because of its small size and weight (10 g). Moreover, because of its climbing habits 1t comes down to the ground where traps are placed only occasionally (DIETERLEN 1971). Dendromus sp. An almost unbelievable observation was reported by HEUGLIN & FITZINGER (1866) and HEUGLIN (1877) but nev- er cited in the scientific literature: HEUGLIN collected spec- imens of Dendromus on the Nile island Argo near Don- gola (19.13 N, 30.27 E) in N Sudan, that is, in the mid- dle of the Nubian desert. The original text (in German) of HEUGLIN & FITZINGER (1866) reads: “Anmerkung. Aus dieser Gattung kommen in Nubien, Ost-Sudan und Abyssinien verschiedene Arten vor. Leider sind Heuglin die beiden auf der Insel Argo bei Dongola und im Belegas-Thale in Abyssinien von ihm eingesammelten in Verlust geraten, bevor er dieselben genau untersuchen und die Art bestimmen konnte.“ HEUGLIN (1877) wrote: „Bisher kannte man nur 190 Fritz DIETERLEN: Climbing mice of the genus Dendromus in Sudan and Ethopia súdafrikanische Vertreter der Gattung der Baummáuse (Dendromys). Mir ist es gelungen, mehrere hierher gehörige Arten in Nordostafrika aufzufinden. Leider sind mehrere der gesammelten Exemplare in Verlust gerathen, namentlich einige, welche wir in Vogelnestern auf der In- sel Argo bei Dongolah erbeuteten, andere im Belegas- Thal...*. In brief, he collected several Dendromus in bird nests on the island of Argo but the specimens were un- fortunately lost before they could be studied properly. HEUGLIN’S (1863) descriptions of two forms from Ethiopia, D. mystacalis and D. pallidus (now a synonym of D. melanotis), is good proof that he perfectly knew these rodents. Therefore there is no reason to question that his specimens from Argo Island were members of the genus Dendromus. HEUGLIN’S (1877) report suggests that he himself participated in the capture of the animals. I became aware of this bibliographic record rather late, around 1995, but had already earlier (DIETERLEN 1971, p. 130) expressed the assumption that specimens of Dendro- mus could have existed along the White Nile in S Sudan during times of a more favourable climate. It is common notion now that Dendromus (and other animals) had the opportunity to disperse from the Upper Nile region in Uganda northwards as early as 10.000 yrs B.P., when the tropical rain forest began to extend northwards, along with the connected savannah belt. During that period the White Nile gained its role as a large river with an enormous cli- matical influence, as it has today (see KENDALL 1969; Liv- INGSTONE 1975; HAMILTON 1982). In the absence of fos- sil evidence or recent records that could document a north- ward dispersion, we can however presume that abundant dry or moist grasland habitat suitable for species of Den- dromus existed in the Bahr-el-Ghazal region. It is not known whether Dendromus still exists on Argo Island. How did these animals reach that remote place? One possibility is by natural dispersal during a longer favourable (moist) Quaternary period (ca. 8000-3000 B.P.; see LIVINGSTONE 1975) in the Sahara. Then its historical occurrence on Argo Island would have been extremely iso- lated. An alternative would be a displacement on a ship during historical times along the White Nile (by more than 2000 km), or from the Ethiopian Highlands or the lower parts of the Blue Nile valley, or downstream the Atbara River. Dendromus melanotis (A. Smith, 1834) Not recorded from Sudan, but the presence of D. melan- otis spectabilis Heller, 1911 cannot be excluded because it occurs at Rhino Camp, Uganda, just about 100 km away from the Sudanese border. The forms spectabilis and ni- grifrons True, 1892 are probably closely related and are both treated as synonyms of D. melanotis by MUSSER & CARLETON (2005). Dendromus messorius (Thomas, 1903) Distributed in W Africa (Ghana to Cameroun), D. R. Con- go, Uganda and Kenya. MUSSER & CARLETON (2005) recorded specimens from S Sudan, two from Torit (US- NM 299 833-834), and another specimen from the for- mer locality Gondokoro in the region of Juba. The form ruddi Wroughton, 1910 is known from Rhino Camp, Uganda. Dendromus messorius prefers moist habitats and was mostly recorded from open areas close to rain forest (HATT 1940; DIETERLEN 1971). The species can be con- founded with D. mystacalis because of its white belly and the frequent absence of a mid-dorsal stripe. Both species may occur sympatrically (HATT 1940). Dendromus ruppi n. sp. (Figs 5, 6, 7) Dendromus mesomelas subsp.: Setzer 1956 Holotype. SMNS 27 572, skin (mounted on cardboard) and skull of adult female; field number 447; collected 23 April 1978 by Hans Rupp. Condition of skin and skull good. Standard measurements (in mm): Head and body length 74 mm, tail 112 mm, hindfoot 19 mm, ear 15 mm, weight 10 g. Paratypes (17). ZMUC 10805, Gilo/ Sudan, leg. J. S. Owen; ZMUC 14121, Gilo, leg. J .S. Owen; ZMUC 14124, Gilo, leg. J. S. Owen; ZMUC uncatalogued, field no. 978, Gilo, leg J. S. Owen; MCZ 45256, Gilo, leg. J. S. Owen; MCZ 45265, Gilo, leg. J. S. Owen; SMNS 27235, Gilo, leg. G. Nikolaus, field no. 386; SMNS 27236, Gilo, leg. G. Nikolaus, field no 387; SMNS 27570, Gilo, leg. H. Rupp, field no. 388; SMNS 27571, Gilo, leg. H. Rupp, field no. 408; SMNS 27572, Gilo, leg. H. Rupp, field no. 447; SMNS 27573, Gilo, leg. H. Rupp, field no. 508; SMNS 27574, Gilo, leg. H. Rupp, field no. 540; SMNS 27575, Gilo, leg. H. Rupp, field no. 541; SMNS 30086 Gilo, leg. G. Nikolaus, field no. 1503; SMNS 30087, Gilo, leg. G. Nikolaus, field no. 1518; SMNS 30088 Gilo, leg. G. Nikolaus, field no. 1519. Type locality. Gilo, Imatong Mts., East Equatoria, South Sudan; altitude ca. 1800-1900 m. Two habitats were not- ed by Rupp and Nikolaus: “near the potato farm” and “in grassy biotopes around a swamp”. Diagnosis. A species of Dendromus resembling D. mys- tacalis, but clearly larger in head and body length and in 191 Bonner zoologisch Bee) aD SHG x GUE Ms on AG |S 5 ASUS ABl20 | | somnter E,NIOLAUS AO G 13% as DENDROMUS vat Y STACALIS LO. AQ > L LO, SÜD- SUDAN or 42 Sommer @ NIKOLAUS - Gew 45 Art Denelromus ny chacalis ka PO meR CAME can 2 9 Eloy Tatar MMe, Ohr AG $~Grola., Denclromus mystacal’s . 6% loz ¡0 y un 2 IEA da ar S 107 03 = 3 Glo, Tone Fang He, ow Ab In: = SIR 1124/20 js Hur a : Y 1 nDendromns impstacalis «2 as a > ES | powm23 447 ana (23 N 6. Mleolans m) a Shas A E i Denolromas OS [9 dorm 22.4, 4978 Gear. grr Santo | Fundort Gr le Tenn Fong Me, RE RE GE wear? | is ont Mileolans ze Fig. 5. Skins of 6 specimens of Dendromus ruppi n. sp. in dorsal and ventral view. From left to right: SMNS 27235, 27236, 27574, 27575, 30086, 30087 (all paratypes). 192 Fritz DIETERLEN: Climbing mice of the genus Dendromus in Sudan and Ethopia Table 2. Body and skull measurements of the specimens of Dendromus ruppi n. sp. collected in the area of Gilo. Mus. No. age HB TL HFL EL ZMUC 10805 — 82 91 20 14 ZMUC 14124 — 74 104 20 13 ZMUC 14121 — 79 106 20 17 ZMUC (978) — 77 95 20 3 MCZ 45256 2 69 98 18 14 MCZ 45265 3 76 107 19 13 SMNS 27235 3 75 110 20 15 SMNS 27236 2 74 108 20 15 SMNS 27570 - 72 98 19 15 SMNS 275715 74 110 19 14 SMNS 27572 3 74 112 19 15 SMNS 27573 — 63 98 19 14 SMNS 27574 4 69 109 21 16 SMNS 27575 3 70 99 20 16 SMNS 30086 2 81 115 22 18 SMNS 30087 2 69 94 20 16 absolute tail length: D. mystacalis 82.0 mm (Sudan), 93.4 mm (Ethiopia) vs. D. ruppi 103.4 mm. Underparts pure- ly white, median dark stripe on dorsum and head present in all specimens studied. A tuft of white hairs is present at the ear basis. Distribution. At present only known from the region of Gilo in the Imatong Mountains, Sudan. WT (g) GRLE MI1-M3_ INT BRC-W - 0522: - - - - 22.1 - ~ - c. 23.0 = - = 12 23.3 3.63 3.4 11.0 15 22.1 3:53 3.3 10.7 11 _ - - - 10 — 3.38 - - 10 22.4 3.47 3.4 10.3 8 - 3.56 - - 11 - 3.54 - - 10 - 3.64 - — 15 23.9 3.61 3.5 11.1 10 22.4 3.50 3.3 10.1 Description. A small Dendromus resembling D. mysta- calis, but clearly larger and with a much longer tail (Fig. 5). Characterized by white underparts, the hairs white from base to tip. Underparts well delimited from the typical brownish pelage on head and dorsum. The blackish mid- dorsal stripe is variable in length but never absent; nor- mally extending from the neck back to the middle dorsum or even to the base of the tail; forward from the neck the stripe (the colour now changes to dark brown) may reach Fig. 6. Two skulls of Dendromus ruppi n. sp.in dorsal and ventral view. Left, SMNS 27572 (holotype), a specimen of medium size (22.4 mm); right, SMNS 30086, the largest specimen (23.9 mm). Note the well rounded braincase and the shape of the zygo- matic arches compared to D. mystacalis (Fig. 3). Bonner zoologische Beitráge 56 193 Table 3. A comparison of body and skull measurements of Dendromus ruppi n. sp., D. mystacalis from the South Sudanese low- land, and the Ethiopian D. mystacalis. Mean, range, and sample size given. RTL = relative tail length. WT (g) GRLE M1-M3 BRC-W Species HB TE RTL HFL EL INT D. ruppi (Gilo) 73.6 103.4 140% 19.8 14.8 11.2 22:7 3.54 3.38 10.62 63-82 91-115 18-22 13-18 8-15 22.1-23.9 3.38-3.64 3.3-3.5 10.0-11.1 n 16 n 16 n 16 n 16 n 10 ns n9 n5 n5 D. mystacalis 60 82 137% 17.2 11.9 _ 19.6 3.14 _ = (Sudan) 53-68 72-91 16-18 11-14 18.7-20.7 2.84-3.31 nll nll n 10 n 7 ns n4 D. mystacalis 66.4 93.4 141% 17.3 13.7 = 21.52 3.35 3.23 9.79 (Ethiopia) 58-76 84-108 16-20 11-18 20.3-22.6 3.13-3.63 3.09-3.40 9.37-10.50 n 10 n 10 n 10 n 9 n 6 17 nó n5 Fig. 7. Upper right molars of Dendromus ruppi n. sp. (SMNS 30086). Cusp pattern on M2 starting to wear down; M3 typical- ly small. its end between the eyes. A black hair spot on the heels, as typical for the genus. The hind feet are reddish brown. The short hairs on the ears are also reddish brown. There is a small but distinct spot of white hairs at the base of the ears. The upper side of the tail is pigmented dark but bears short light bristles. Mammary formula 2 — 2 = 8. D 5 of hind foot bearing a short claw. The skull (Fig. 6) has the typical characteristics of the genus: rounded braincase, narrow zygomatic plate and a distinct masseteric knob at its lower anterior corner. An- terior edge of supraorbital bow smooth (not chambered). Upper incisors longitudinally grooved. M1-length about 57% of upper molar row, M2 ca. 35%, M3 minute. Mo- lars (Fig. 7) rather narrow, breadth of M1, M2, M3 clear- ly larger than in mystacalis (see below). The posterior part of M1 behind its prominent t8 relatively broad in appear- ance (but similar to the same area of D. mystacalis). In most specimens t9 is worn down on the labial side and appears crater-like. Measurements. Tables 2, 3. Ecological and biological data. Three out of seven spec- imens in the SMNS were collected in a swamp near Gi- lo at c.1800 m, and four in the area of the (so-called) “po- tato farm” above Gilo (c.1900 m). The two MCZ speci- mens came from a “nest in olive tree” (SETZER 1956). The SMNS sample comprises six males and five females, col- lected between April 1978 and August 1979. A female caught in April 1978 had four embryos (13 mm in diam- eter); four males caught between April and July had tes- ticle diameters of 3, 5, 5, and 6 mm, respectively. Comparisons. Dendromus mystacalis lineatus in Sudan is a lowland species, with its highest altitudinal record in Katire at c. 1000 m, on the foothills of the Imatong Mts. 194 Fritz DIETERLEN: Climbing mice of the genus Dendromus in Sudan and Ethopia Table 4. Measurements of body and skull of D. mystacalis from Ethiopia. Age class 1 sem.-young adult, 2 young adult, 3 adult, 4 adult/old adult, 5 old adult. HB T Coll.No. age class BMNH 28.1.11.143 3 71 87 18 13 BMNH 128.1.11.144 - 61 94 18 BMNH 28.1.11.145 3 67 84 17 18 BMNH 28.1.11.146 1 58 89 16 14 BMNH 1934.2.24.88 — 58 87 16 13 BMNH 59.660 - 65 95 16 13 BMNH 70.655 4 76 105 17 12 BMNH 70.656 2 64 96 16 11 SMNS 23720 2 71 108 19.5 - SMNS 23732 4 73 89 18 14 Mean range n - 66.4 93.4 17.3 13.7 58-76 84-108 16-20 11-18 n 10 n 10 n 10 n 9 Other populations (outside Ethiopia) are also known from lower altitudes (BOHMANN 1942; DIETERLEN 1971; KING- DON 1974; ANSELL & DOWSETT 1988). The typical D. mys- tacalis from the Ethiopian Highlands, however, is exclu- sively a high mountain animal. D. ruppi ıs also a high- land species, known from the Imatong Mts. at altitudes between 1800 and 1900 m. On the neighbouring Mt. Kinyetti (3187 m, the highest peak in Sudan Republic) an occurrence at even higher altitudes may be possible up to about 2500 m. D. ruppi and (non-Ethiopian) D. mystacalis share the white pelage of the underside and further characters, but D. ruppi is considerably larger in external, cranial, and dental measurements (length of upper molar row, Table 5). The dorsum of D. ruppi is darker than in D. mystacalis and always shows a mid-dorsal stripe. There are further differences in the breadth of the molars between D. rup- pi (n= 9) and D. mystacalis (n= 5). M1 from “tl” to t6: ruppi 1.14 mm (1.02-1.21) vs. mystacalis 1.02 mm (0.98-1.05). M2: ruppi 0.97 mm (0.86-1.04) vs. mysta- calis 0.83 mm (0.80-0.86). M3: ruppi 0.49 mm (0.47-0.53), no data available for mystacalis. GRLE WT (g) BRC-W — 21:71 3.60 3.19 9.37 15 — = x = - 21.53 3.30 3.09 9.70 — 20.27 3.24 3:17 - - 22.31 3.42 3.40 9.78 - 20.68 3.13 3.24 9.63 13 = 3.43 = = = 22.61 3.33 3.28 10.50 - 21.52 3:35 3.23 9.79 20.3-22.6 3.1-3.6 3.1-3.4 9.4-10.5 n 6 ne? n6 n5 Comments. SETZER (1956) discussed two specimens of Dendromus from Gilo/Imatong Mts., South Sudan, in the MCZ: “These two specimens are somewhat darker than D. mesomelas percivali (now a synonym of D. insignis, according to MUSSER & CARLETON 2005) from Mount Gargues, British East Africa. The external measurements agree rather well with percivali, but the skulls of the Gi- lo specimens, even though adult, are markedly smaller... It is apparent that these animals from the Imatong Moun- tains are different from any of the surrounding kinds, but owing to the broken condition of the skulls and there be- ing only the two specimens I feel it best to identify them only at the specific level.” SETZER (1956) did not know that J. S. Owen, the collector of the MCZ specimens, had already sent another four specimens from the same local- ity to the ZMUC at Copenhagen, which Setzer could not consider in his paper. He also failed to recognize that the specimens had entirely white underparts and therefore could not belong to D. mesomelas. In the latter species the hairs of the underside are brownish-grey with an always dark basis, only on the throat and in the anal region they can be whitish (BOHMANN 1942; ROSEVEAR 1969; DIE- TERLEN 1971). Contrary to Setzer’s statement, one of the Bonner zoologische Beiträge 56 195 skulls of the specimens at Harvard (MCZ 45 256) was not damaged (measurements in Table 4) and the other (MCZ 45 265) had already been repaired by MCZ staff when I measured it in 1988 (Table 4). The occurrence of D. me- somelas (now D. insignis) in the mountains of South Su- dan could be expected, because D. insignis was already known then from high altitudes of neighbouring Ethiopia, Kenya, and Uganda. Besides the Imatong Mts., mountains neighbouring to the east (Dongotona and Didinga Mts.) could also be popu- lated by D. ruppi, at least above 1800 m. For the present, however, D. ruppi must be regarded as a species endem- ic to the Imatong Mts. which became isolated as a result of climatic changes during a drier Pleistocene postglacial/pluvial phase in Equatorial Africa (Rupp 1980; LIVINGSTONE 1975). Etymology. I dedicate this new species to my deceased friend and colleague Hans G. Rupp. Hans died on 15 June 1979 at the age of 32, after several weeks of severe ill- ness in a hospital in Nairobi/Kenya, to where he had been transported from South Sudan. As a student of zoology at the University of Tubingen,Germany, he had begun ex- tensive fieldwork for his doctoral thesis on rodents and their entoparasites in southern Sudan 17 months before. In the early 1970s he undertook a number of research trav- els to Ethiopia and published the results about systemat- ics, distribution and ecology of Ethiopian rodents (DIE- TERLEN & Rupp 1976, 1978; Rupp 1980). 3.2. Species of Dendromus in Ethiopia Four species of climbing mice are known within the ter- ritory of Ethiopia: Dendromus mystacalis (Heuglin, 1863) Holotype. SMNS 1055; a specimen collected by Heuglin in 1863 and sent to the Stuttgarter Naturalienkabinett (now SMNS) where it was registered under no. 1055 in the (Accession-) Catalogue for mammals, founded in 1837. The well-preserved specimen was mounted in a climbing posture on a short vertical branch, anchored in a wooden pedestal (Fig. 8). No skull is preserved. The in- scription on the label reads: “1055 Typus! male Dendro- mus mystacalis Hgl. Eifaz, Abyss. v. Hgl. 1863“. Based on its size and the measurements given by HEUGLIN (1863) the specimen was subadult. After 144 years of preserva- tion the reddish-brown dorsal pelage has become dull and the (originally?) weakly developed mid-dorsal stripe dis- appeared. However, the blackish-grey basal part of the hairs remains unchanged. HEUGLIN’s original description (in German) reads: rie I IR ALA G PY fr ce Fig. 8. | Mounted holotype (SMNS 1055) of Dendromus my- stacalis (HEUGLIN, 1863). „Oberseite und Außenseite der Füße zart rostig ockerfar- ben; Unterseite ein eckig nach den Zügeln einspringen- des Feld, das von den Halsseiten ausgeht und bis unter und vor das Auge reicht, sowie Innentheile der Behaarung der Füße rein weiß, beide genannte Farben streng von ein- ander geschieden, auf der Rückenmitte schwache Andeu- tung eines graulichen Längsstreifes. Schnurren schwärz- lich; hinter der Ohrbasis ein hell ockerfarbiger Fleck; Schwanz lang, schlank, mit sehr zarten Härchen ziemlich dicht besetzt; Nasenkuppe, Nägel und Zehenballen fleisch- farbig. Auge mittelgroß, hervorstehend, braun, längs der Außenseite der oberen Schneidezähne eine starke Furche. Körperlänge 2” 5”” (= 65 mm, given in the old German measurement Zoll). Schwanzlänge 3” (= 75 mm). Ohr- höhe stark 15" (=> 15 mm). Das Ohr zu beiden Seiten etwas um- und eingebogen; ausgebreitet so breit als hoch.Wir erbeuteten nur ein einziges Exemplar dieser zier- lichen Art, und zwar im Horst eines Raubadlers, wo das Thierchen wohl 40 Fup über dem Boden, seine Behausung aufgeschlagen hatte. Im Magen fand sich ein grüner Pflan- zenschleim. Vorkommen in der Ebene von Eifag in Cen- tral-Abessinien, auf 6000 Fuß Meereshóhe.* 196 Fritz DIETERLEN: Climbing mice of the genus Dendromus in Sudan and Ethopia Table 5. Comparison of body and skull measurements summarized from published collections of D. mystacalis of Sudan (this pa- per); Congo (DIETERLEN 1971); East Africa (HOLLISTER 1919); South Africa (DE GRAAFF 1981), and Africa (globally, BOHMANN 1942) , and those from the highlands of Ethiopia (this paper) (Data given by VERSCHUREN, VAN DER STRAETEN & VERHEYEN 1983 from Congo and Rwanda are not included but confirm the summarized measurements of Africa). Measurement Africa Ethiopia HB 63.5 mm (53-80) 66.4 mm (58-76) n 92 n 10 T 82.0 mm (63-100) 93.4 mm (84-108) n 92 n 10 HF 16.6 mm (15-20) 17.3 mm (16-20) n 61 n 10 E 13.0 mm (10-17) 13.7 mm (11-18) n 52 n 9 WT 8.15 ¢ (4-14) = n 35 GRLE 19.5 mm (17.5-21.6) 21.52 mm (20.3-22.6) n 88 n 6 M1-M3 3.10 mm (2.8-3.5) 3.55 mm (3.13-3.63) n 91 n7 INT 2.87 mm (2.5-3.5) 3.23 mm (3.09-3.40) n lll n6 BRC-W 9.30 mm (8.4-10.0) 9.79 mm (9.39-10.50) n 103 n5 Comparison of the holotype with further specimens collected. Since the type seems to have never been stud- ied or compared, and Heuglin’s description was barely consulted, a comparison with further mystacalís specimens from Ethiopia and other parts of Africa has probably not been undertaken. Annotations as to literature, distribution and collections can be found in the “Catalogue of the Mammals of Ethiopia” (YALDEN et al. 1976, 1996) and in papers with isolated data or single unpublished records. Yet, no critical comparisons of the Ethiopian mystacalis with all other forms designated as mystacalis seems to have taken place. Material of Ethiopian D. mystacalis studied. BMNH 1934.2.24.88, Bab Bar Sir, Wuha River, 3 miles N of Lake Tana, Cheeseman Collection; BMNH 59.660, Sidamo (07.00 N/ 36.30 E), coll. F. K. Allison; BMNH 70. 655, Mouth of Didessa River (10.05 N/35.38 E), coll. Great Ab- bai Expedition 1968; BMNH 70. 656, Junction of Nile and Guder River, Blue Nile Gorge (09.50 N/ 37.41E), same Expedition; BMNH 28.1.11.142, 28.1.11.143, 28.1.11.144, 28.1.11.145, all from Dangila, S of Lake Tana (11.17 N/ 36.56 E); BMNH 28.1.11.146, Kola village, Domkan Riv- er (10.42 n/36.31 E), all by Cheeseman Collection; SMNS 1055, Eifag (Eifaz) (12.03 N/37.46 E), T. von Heuglin (holotype); SMNS 23720, Chencha (06.16 N/37.40 E), H. Rupp; SMNS 23732, Dorsey (Dorse) (06.13 N/37.40 E), H. Rupp. Description. Generally little colour variation among the forms of D. mystacalis. The colour of the upper parts of Ethiopian specimens is very similar compared to other forms: a brownish- or ochre-grey (also reddish tinge) over the whole back and head. Only the terminal part (2-3 mm) of the pelage hairs 1s coloured as described, the basal part of the dorsal hairs is uniformly dark grey. The black mid- dorsal stripe is variable (Fig. 9): in four specimens the stripe is complete from neck to root of tail, in three oth- er specimens the stripe is thin and not visible throughout its length, and in four there is no mid-dorsal stripe at all; see also YALDEN et al. (1976). No geographical variation in this character could be observed. The underside is pure white from chin to belly. Like in all species of the genus there is a dark spot of hairs just above the heels (DIE- TERLEN 1971). The hairs of the dorsal surface of the hind foot are somewhat reddish-brown as are the short hairs on the ears. The skin of the short-bristled tail is pigmented dark on the upper side and whitish below. Localities and altitudes (Fig. 10). Altitudes mostly as giv- en by collectors or/and authors. Alemaya, Lake 2100 m (09.21 N, 42.01 E); Axum, S of, 2100 m (14.08 N, 38.45 E); Bab Bahr, 3 miles N Lake Tana, 1800 m (12.16 N, 37.07 E); near Boralugu, 3000 m; Chencha, 9 km N, 2700 m (06.16 N, 31.40 E); Dangila, 2000-2100 m (11.17 N, 36.56 E); Debre Tabor, 2500 m (11.50 N, 38.02 E); Didessa River (mouth), 1200 m (10.08 N, 35.38 E); Bonner zoologische Beitráge 56 197 Fig. 9. Ethiopia. Skin of a Dendromus mystacalis (SMNS 23732) from Didessa River (near Guma), 1900 m; Domkan (Dibken), 2000 m; Dorse (Dorsey), S, 2600-3000 m (06.13 N, 37.40 E); Eifag (Eifaz, Ifag), 2000 m (12.03 N, 37.46 E); Gud- er River, junction of Nile, 1800 m (09.50 N, 37.41 E); Harenna Forest, 2000 m; Katcha, S Mt. Batu, 2400 m (06.42 N, 39.44 E); Kebre Mengist, 70 km NW. 1680 m (06.23 N, 38.35 E); Kolay (Golay), Domkan River, 1600 m (10.42 N, 36.31 E); Ladjo, NW Mt. Goba, 3850 m (07.08 N, 39.33 E); Masslo, c. 40 km SW Mt. Goba (06.42 N, 39.50 E); Menagasha Forest, c. 2800 m (08.55 N, 38.37 E); Mt. Badda, W (07.50 N, 39.45 E); (Mt.) Goba, 2800 m (07.01 N, 39.59 E); Sidamo, Arussi, 2500 m (07.40 N, 39.45 E); Simien Mts. National Park, 3100-3600 m (c. 13.15 N, 37.50 E). The altitudinal distribution varies be- tween 1200 and 3850 m, with most records between 2000 and 3000 m. The collecting localities are scattered over a vast region encompassing large parts of the western and central highlands (YALDEN et al. 1976). D. mystacalis oc- curs in the vegetational zone of Woina Dega (ca. 1500-2300 m), including the (so-called) Highland Zone and the lower Montane Zone. Above Woina Dega the species ascends up to 2300-3300 m, reaching parts of the upper Montane and lower Temperate Zone (Rupp 1980). Habitat. “This species has been recorded from a wide range of altitudes (900-3200 m) but is nearly always as- sociated with long grass and bushes.” (YALDEN et al. 1976). In the Menagesha Forest, BEKELE (1996) found mystacalis ın bush vegetation (dominant plant Carissa edulis). Preferred habitats in the Simien Mountains Na- tional Park are Giant Heath (Erica arborea) forests with thick undergrowth (MULLER 1977; MUHMENTHALER 1999; WEHRLI 1999). Notes on biotopes and nests of D. mysta- calis are as follows: Heuglin’s type came from a nest 40 ft. high in a tree with an eagle’s aerie. Three specimens of the Cheeseman collection (at Dangila) were taken at considerable height in trees: “in a straw bee hive, 30 feet from ground”, “same grass nest in tree in high grassland”, “same tree as in...”. Similar observations were made in other regions of Africa. In East Africa the species often frequents disused weaver and bishopbird nests. In the Kivu region, however, nests of D. cf. mystacalis were never found in trees, but in the absence of trees and shrubs in dry grassy biotopes with Melinis minutiflora and other Gramineae; nests were found between stems of grass. D. mystacalis was rarely seen in moist grassy biotopes (DI- ETERLEN 1971). Comparisons. A comparison of body and skull measure- ments of D. mystacalis samples (Tables 1, 4, 5) from var- ious regions of Africa, but excluding Ethiopian popula- tions, revealed surprising results. Mean values for each character (Table 5) were compared with the relatively small Ethiopian sample. In most measurements a distinct difference exists between the true Ethiopian mystacalis and the remaining populations, particularly in the relative tail length, length of skull, and other skull measurements. Fig. 10. Distribution of Dendromus mystacalis in Ethiopia. 198 Fritz DIETERLEN: Climbing mice of the genus Dendromus in Sudan and Ethopia A further difference is altitudinal distribution: Ethiopian mystacalis have a strictly montane distribution, while the forms which occur outside Ethiopia occur principally be- low 2000 m. These results question the conspecificy of all the described forms within D. mystacalis, an issue which should be addressed in the future, including genet- ic data. The description of HEUGLIN (1863) and the char- acters of the type specimen apply only to the Ethiopian populations. All the similar but clearly smaller non- Ethiopian forms, considered at present as part of D. mys- tacalis, may belong to one or more other species.The sta- tus of all the populations of D. mystacalis outside of Ethiopia should be reconsidered. The names Dendromus ansorgei Thomas & Wroughton, 1905 (Angola) and D. Jamesoni Wroughton, 1909 (South Africa) are candidates for a possible lowland species. Dendromus melanotis (A. Smith, 1834) YALDEN et al. (1976, 1996) listed four specimens from three localities at altitudes between 1200 and 2500 m. An- other specimen (AMNH 81119) was recorded by Muss- ER & CARLETON (2005). Another three specimens not yet published were collected by H. Rupp near Chencha (2400-2700 m): two adults (SMNS 23721, 23722), one subadult (SMNS 23723). Measurements of the adults are: HB 64, 78 mm; T 85, 94 mm; HF 19, 19 mm; EL 15, 15 mm; GRLE 22.45, 22.51 mm; M1-M3 3.36, 3.51 mm; BRC-W 10.51, 11.04 mm. No material of the form pallidus, described by HEUGLIN (1863) and listed as a synonym melanotis by MUSSER & CARLETON (2005), was available for study. Heuglin wrote “Lippengegend und die Unterseite rein weiß“ (area of the lips and underparts pure white), but without mentioning the grey basis of the white hairs, typical for melanotis, and without comparing it with his type specimen of D. mvs- tacalis. Dendromus insignis (Thomas, 1903) The taxon insignis, formerly included in D. mesomelas by BOHMANN (1942) and MISONNE (1974), is considerably larger than the South African species D. mesomelas. Os- GOOD (1936) was right to describe his record from the E slope of Mt. Albasso, Ethiopia, as D. insignis abyssinicus, now known by several more specimens from the high- lands. Three specimens in the BMNH (72.1294, 72.1295, 76.113, leg. Largen et al. 1971) recorded from “Bale Din- shu” above 3000 m, and labeled as D. mystacalis, are un- mistakably D. insignis (Dieterlen, unpubl.). Two further adult specimens from the same region were collected by H. Rupp and G. NIKOLAUS in 1971 and 1976 (SMNS 23724, 23983). Measurements: HB 68, 75 mm: T 103, 101 mm; HF 22/23, 21/22; EL 17, 12 mm; WI = 10% 46GR- LE -, 22.08 mm; M1-M3 3.52, 3.31 mm; BRC-W -, 10.94 mm. Dendromus lovati (De Winton, 1899) Endemic to Ethiopia. A distinctive small species, very dif- ferent from all congeners (HB 68-90 mm, T 72-80 mm, WT 12-16 g). Not very common, at least not in trapping yields. Altitudinal distribution 2500 to 3550 m. Preferred habitats are grassland and giant heath (Erica arborea) forests (YALDEN et al. 1976, 1996; MÜLLER 1997; MUH- MENTHALER 1999). Material studied. One specimen from Ankober (SMNS 23927). leg. G. Nikolaus. Acknowledgements. I express my sincere thanks to Dr. Pauli- na Jenkins for her kind help during my repeated visits of the British Museum of Natural History, London (BMNH). I am grateful to several curators for assistance in the following mu- seums and for the loan of important specimens: Dr. Hans Baa- goe, Zoological Museum of the University of Kopenhagen (ZMUC): Dr. Maria Rutzmoser, Museum of Comparative Zo- ology (MCZ) at Harvard University, Cambridge (Mass. ); the late Dr. Henry W. Setzer (in 1976) and Dr. Michael Carleton, Unit- ed States National Museum (USNM), Smithsonian Institution, Washington D.C., Dr. Rainer Hutterer and Dr. Gustav Peters, Zo- ologisches Forschungsmuseum Koenig (ZFMK), Bonn. Some of these visits were supported by the Deutsche Forschungsge- meinschaft (DFG), Bonn. In 2006 my research as a visitor in the collections of the BMNH was supported by SYNTHESYS (the European Union-funded Integrated Infrastructure Initiative grant). My sincere thanks to both of these funding organisations. Dipl. Biol. Kathrin Marquart (Stuttgart) and Uwe Vaartjes (Bonn) kindly assisted in the preparation of figures. Zusammenfassung. Die in Athiopien und Sudan vorkommen- den Arten der Gattung Dendromus (Rodentia: Dendromurinae) werden revidiert. Die zu Dendromus mystacalis (Heuglin, 1863) gerechneten Formen der Klettermáuse aus Sudan und anderen Regionen Afrikas werden mit der typischen Population aus Athiopien verglichen. Athiopische D. mystacalis unterscheiden sich von anderen afrikanischen Populationen in ihrer Morpho- logie und im Lebensraum, während alle Populationen außerhalb Äthiopiens sich in Größe, Erscheinung gleichen und in ähnli- chen Lebensräumen leben. Eine neue Art, Dendromus ruppi, wird nach Material aus den Imatong Bergen im Süd-Sudan be- schrieben. Sie ist eine relativ große Bergform, die vermutlich während klimatischer Schwankungen in einer postpluvialen Pha- se des Pleistozäns isoliert wurde. Die neue Art ist vermutlich en- demisch für die Imatong Berge. Bonner zoologische Beiträge 56 199 REFERENCES ANSELL, W. F. H. & DOwSETT, R. J. (1988): Mammals of Malawi: An annotated checklist and atlas. 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Author’s address: Dr. Fritz DIETERLEN, Staatliches Mu- seum für Naturkunde Stuttgart, Rosenstein 1, D-70191 Stuttgart, Germany; E-Mail: fritzdieterlen@hotmail.com Received: 05.01.2008 Accepted: 30.05.2009 | Bonner zoologische Beiträge Band 56 Heft 3 Seiten 201-208 | Bonn, September 2009 A large new species of Sylvisorex (Mammalia: Soricidae) from Nigeria and the first record of Sylvisorex ollula from the country Rainer HUTTERER & Christian MONTERMANN Zoologisches Forschungsmuseum Alexander Koenig, Bonn, Germany Abstract. A new species of shrew, Sylvisorex corbeti n. sp., is described from SE Nigeria. The only specimen known was previously assigned to Sy/visorex ollula, but is considerably larger, has a longer tail, elongated hindfeet and a very large skull and thus represents the largest species of the genus. From the Obudu Mountains in SE Nigeria a new speci- men is recorded which agrees well with typical S. ollula from lowland forest of southern Cameroon, and which repre- sents the first species record from Nigeria. Key words. Africa, Nigeria, Mambilla Plateau, Sy/visorex, new species. 1. INTRODUCTION The shrews of Nigeria were reviewed by HUTTERER & HAPPOLD (1983) who recorded 23 species in three genera, including Sylvisorex Thomas, 1904, at that time known from the country only by Sylvisorex megalura (Jentink, 1888), a species sometimes included in the genus Suncus Ehrenberg, 1832 (HUTTERER 2005). HUTTERER et al. (1992) added three further species from mountain forests in SE Nigeria, Sylvisorex camerunensis Heim de Balsac, 1968, Sylvisorex ollula Thomas, 1913, and Crocidura attila Doll- man, 1915. Sylvisorex ollula is the largest of the 14 species of the genus known so far (HUTTERER 2005; HUT- TERER et al. 2009; MUKINZI et al. 2009). It was described from lowland forest near the River Ja in SE Cameroon (THOMAS 1904) and is known also from Gabon (BROSSET 1988; GOODMAN & HUTTERER 2004), Equatorial Guinea (Lasso et al. 1996), Central African Republic (Ray & HUTTERER 1996), Republic of Congo (QUEROUIL et al. 2003), and Democratic Republic of Congo (DIETERLEN & HEIM DE BALSAC 1979). Records from Cameroon are rare and concentrated in the south of the country (HEIM DE BAL- SAC 1968; GOODMAN et al. 2001). To the west, the species 1s known from around Mount Cameroon (HEIM DE BAL- SAC 1959, 1968) and from the Korup National Park (HUT- TERER & SCHLITTER 1996). The record from SE Nigeria therefore appears to be isolated. However, more specimens have become available in recent years, both from Cameroon and Nigeria, and we are now in a position to re-evaluate the systematics of the populations at the north- western edge of the range of the S. ollula group. 2. MATERIAL AND METHODS We studied new material from Nigeria and Cameroon obtained by G. Nikolaus and M. Bare. from 1992 to 2006 (see below). These specimens are deposited in the Zoologisches Forschungsmuseum Alexander Koenig, Bonn (ZFMK); other material studied is deposited in the Estacion Biologico Donana, Sevilla (CET) and the Natural History Museum, London (BMNH). Skull measurements were taken with an electronic calliper. All measurements are given in millimetres (mm), and body mass in grams (2). The terminology of cranial and dental structures follows MEESTER (1963) and JENKINS (1984); external and cranial measurements are the same as previously defined by STANLEY et al. (2005) and KERBIS PETERHANS et al. (2008). The terminology of the skeletal elements of the limbs follows WOODMAN & MORGAN (2005). Digital X-rays were made with a Faxitron LX60 machine. The taxonomy of shrews follows HUTTERER (2005), except for the unsettled allocation of Suncus megalura. For the purpose of this paper we include S. megalura ın the genus Sylvisorex. Comparative material of S. ollula. Cameroon: ZFMK 63.190, camp V, above Mueli, northern side of Mount Cameroon, 600 m; ZFMK 63.533, Nyasoso, N Mount Cameroon; ZFMK 99.683, E Nkongsamba, Mount Nlon- ako, camp near Nguengue, 1200 m, cloud forest; ZFMK 2008.220, Cameroon, Reg. Mamfe, Mukwecha/ Araebisu, 165 m; ZFMK 2008.226, SE Cameroon, Campo Region, Nkoelon, 75 m. Gabon: Equatorial Guinea: CET 736, 202 Rainer HUTTERER & Christian MONTERMANN: A large new species of Sylvisorex from Nigeria Monte Alen (Lasso et al. 1996); Nigeria: ZFMK 95.70, SW Obudu Mountains, Boje Forest (06.15 N, 08.55 E), 290 m. Fig. 1. Flat study skins of a Sylvisorex ollula (ZFMK 95.70) from Nigeria and of the holotype of Sylvisorex corbeti n. sp. (ZFMK 88.91). Natural size. 3. RESULTS 3.1. A large forest shrew from Nigeria Sylvisorex corbeti n. sp. (Figs 1, 2, 4, 5) Holotype and type locality. Skin mounted on cardboard and skull of a young adult female (ZFMK 88.91), collect- ed by G. Nikolaus on 23.03.1988 (field no. GN 60) in for- est swamp near Chappa Waddi, (07° 01° N, 11° 41’ E), 1 900 m a.s.l., Gotel Mountains, SE Nigeria. The sex of the specimen was previously given as “male” by HUTTERER et al. (1992), but the collector determined it as female. Measurements of holotype. Body mass 30 g, head and body length 100 mm, tail 64 mm, hindfoot length 17 mm without and 19 mm including claws, ear conch 10 mm. Diagnosis. A very large and dark blackish brown species of Sy/visorex with a total length of 164 mm. Tail of medi- um length (64 % of head and body length), with no long bristle hairs. Hindfoot long and narrow, metacarpals and metatarsals elongated. Skull large and robust, with a long and wide inter-orbital region, a broad infra-orbital bridge, and a long toothrow. Upper P4 with a large parastyl. Description. Sv/visorex corbeti n. sp. is the largest species of the genus. The head is large, as compared to S. ollula (Fig. 1). The colour of the fur is uniformly blackish brown. The hair bases on the dorsum are plumbeous, and the tips blackish brown. No difference in colour exists between dorsum, venter, and limbs. Hairs on dorsum are about 3- 4 mm in length. Facial vibrissae reach 27 mm in length. Ear conch has a medium size, is round and pocketed, and covered by very short hairs. Hind foot very long and nar- row, with elongated digits and short claws (Fig. 2). Ven- tral inner surface of hind foot with folds but not covered by small granule-like bumps. Tail of medium length, colour uniform, no long bristle hairs present. Skull (Fig. 4) very large with a large hexagonal braincase. Dorsal profile (Fig. 5) straight, rostrum long and robust. Interorbital constriction relatively broad (21.2 % of condy- lo-incisive length, versus 20.7 % in S. ollula), its outer margins running almost parallel. Infra-orbital bridge wide, lachrymal foramen large. Maxillary wide compared to S. ollula, zygomatic process of maxillary pointed. Superior and inferior articular facets large and robust. Anterior palate with three incisive foramina; one pair between the upper unicuspids, and a single one behind. Upper teeth: First upper incisor very large, with a long an- terior hook and a less prominent talon (Fig. 5). First up- per unicuspid very large, second, third and fourth unicus- pids much smaller, the third being a little larger than the Bonner zoologische Beiträge 56 203 Fig. 2. X-rays of a hindfoot of Sylvisorex ollula (ZFMK 2008.226, top) and of Sylvisorex corbeti n. sp., (holotype ZFMK 88.91, bottom); scale is 12.8 mm. Note the difference in length of the metacarpals. Table 1. External measurements (mm) and weight (g) of Sylvisorex corbeti n. sp. and of some S. ollula from Nigeria and Came- roon. Species Origin TL HB Tail HF E WT Tail/HB S. corbeti n. sp., Nigeria 164 100 64 17419 10 30 64 % ZFMK 88.91 S. ollula, Nigeria 142 90 32 14/15 8 17 58% ZFMK 95.70 S. ollula, NW Cameroon 142 92 50 14/16 8 19 54% ZFMK 63.190 S. ollula, NW Cameroon 141 84 57 1517 8 20 68 % ZFMK 69.533 S. ollula, NW Cameroon 131 84 47 14/16 8 21 56 % ZFMK 99.683 S. ollula, NW Cameroon 141 88 53 14/15 8 16 60 % ZFMK 2008.220 S. ollula, SE Cameroon 139 83 56 16/17 8 16 67% ZFMK 2008.226 204 Rainer HUTTERER & Christian MONTERMANN : A large new species of Sylvisorex from Nigeria Fig. 3. Skull of Sy/visorex ollula (CET 736) from Equatorial Guinea in dorsal and ventral view. Total length of skull 24.62 mm. second and fourth. P4 large, with a prominent parastyle and a large paracone (or metacone, according to MEESTER 1963). The talon of P4 ıs longer than in $. ollula, extend- ing closer to M1 than in S.ollula. M1-2 are large and square-shaped in occlusal view, M3 is short (0. 69 x 1.56 mm in the holotype), but not as reduced as in $. ollula (Fig. 4). Mandible: Robust with a long ramus, a low coronoid process, and a wide angular process. COR 54.7 % of LTR (59.6 % in S. ollula). Foramen mentale below posterior root of lower p4. Condyle robust and rectangular, about as wide as high. Lower teeth: Tip of first incisor curved upwards, with two denticulations on cutting blade. Lower p4 as long as wide (occlusal view). Lower molars long and slender in occlusal view; third lower molar without talonid basin. Comparisons. S. corbeti n. sp. is larger than any other species in the genus, as understood here. It can only be compared with $. ollula, which it resembles externally. In absolute external measurements, $. corbeti n. sp. is larg- er and has a longer tail than specimens of S. ollula from Nigeria or Cameroon (Table 1), although the relative tail length is similar. Specimens from Gabon and Equatorial Guinea (GOODMAN & HUTTERER 2004, Lasso et al. 1996) Bonner zoologische Beitráge 56 205 Fig. 4. Skull of Sylvisorex corbeti n.sp. (holotype ZFMK 88.91) from SE Nigeria. Total length of skull 26.48 mm. are smaller also. The fore- and hindfeet of the new species are also considerably longer, due to a prolongation of skeletal elements such as metacarpals and metatarsals (Fig. 6). The skull of the holotype of 5. ollula (THOMAS 1913) was figured by JENKINS (1984), and another specimen from Equatorial Guinea is shown in Fig. 3. $. corbeti n. sp. (Figs 4, 5) is distinguished by size (Table 2), but also by a num- ber of cranial and dental details mentioned above. The largest fossil species, S. olduvaiensis Butler & Greenwood, 1979, is considerably smaller (BUTLER & GREENWOOD 1979); it has a mean coronoid height of 4.5 mm (5.94 mm in the new species). Habitat. The holotype was collected in a forest swamp at 1900 m. It shares this habitat with rodents such as Oto- mys occidentalis Dieterlen & Van der Straeten, 1992 and Praomys obscurus Hutterer & Dieterlen, 1992. Etymology. We dedicate this species to Gordon B. Cor- bet, former curator of mammals at The Natural History 206 Rainer HUTTERER & Christian MONTERMANN: A large new species of Sylvisorex from Nigeria Museum, London. He has served the scientific commu- nity for decades by hosting visitors from all around the world at the British Museum mammal collections. Besides writing textbooks on mammals of the Palaearctic and In- domalayan Regions and the World, he has demonstrated his interest in African mammals by careful revisions of hedgehogs, hyraxes, and elephant shrews. He also de- scribed the now probably extinct Nigerian pygmy hip- popotamus (CORBET 1969). 3.2. A new record of Sylvisorex ollula from Nigeria The presumed record of S. ollula from Nigeria (HUTTER- ER et al. 1992) is no longer valid, after our study revealed that the voucher specimen represents a different species described here as new. However, Sy/visorex ollula does occur in Nigeria, as a new yet unrecorded specimen demonstrates. During a survey of small mammals of the Boje Forest at 290 m in the SW Obudu Mountains (06.15 N, 08.55 E), G. Nikolaus collected a male of S. ollula on 13 January 1995. The specimen (ZFMK 95.70, Fig. 5) is much smaller than S. corbeti n. sp. and agrees well with typical S. ollula in size (Table 1) and shape of skull (Fig. 5). 4. DISCUSSION The discovery of Sy/visorex corbeti n. sp. further enriches the morphological diversity of the genus, as previously summarized by HUTTERER (1985) and HUTTERER & SCHLITTER (1996). With a body mass of 30 g it marks the upper limit of the genus. The new species shows a unique combination of characters of a terrestrial forest shrew (large body, tail of medium length, ear conch of moderate size) with those of scansorial shrews (elongated limbs). The digits of the hindfoot (expressed by the length of metatarsal III) are even longer than in Sy/visorex megalura (Fig. 6), a species known for its adaptation for climbing (VoGEL 1974). We know nothing about the lifestyle of the new species, except that it lives in forest swamp at 1900 m. It is part of a guild of small mammal species confined to the forested parts of the Mambilla Plateau shared by Nigeria and Cameroon (HUTTERER et al. 1992). The true content of the genus Sy/visorex is still not known. New species with surprising adaptations have been discovered in Central Africa (MUKINZI et al. 2009; HUTTERER et al. 2009) and add to the diversity of the genus. We included two further undescribed species from D. R. Congo in Fig. 6, to be described in the future. If these and others are included, the number of species of Sylvisorex is close to 20. Genetic studies suggest that $. megalura may be related to Suncus (QUEROUIL et al. 2001), Fig. 5. Lateral views of the maxillary portion of the skulls of (top to bottom) Sy/visorex corbeti n. sp. (holotype) and S.ollu- la (ZFMK 95.70, SE Nigeria), ZFMK 69.533 (NW Cameroon), ZFMK 2008.220 (NW Cameroon), and CET 736 (Equatorial Guinea). Scale 10 mm. and other authors (DUBEY at al. 2008) propose the inclusion of all African Suncus and Sylvisorex species into one genus. A paraphyly of Sylvisorex was already considered by HEIM DE BALsAC € LAMOTTE (1957). Further problems are posed by the morphologically unique genera Ruwenzorisorex Hutterer, 1986 and Scutisorex Thomas, 1913, both of which are genetically close to Sylvisorex. No conclusive phylogeny is available at this moment, but the study of African forest shrews remains an exciting enterprise. Bonner zoologische Beitráge 56 207 Table 2. Cranio-dental measurements of Svlvisorex corbeti n. sp. and S. ollula from Nigeria and Cameroon. Species CI PL UTR P4-M3 IO MB GW PGL HCC LTR COR S. corbeti n. sp. 26.48 11,59 11.47 6.18 5.61 7.63 11.94 8.13 6.18 10.86 5.94 ZFMK 88.91 S. ollula, SE Nigeria 24.26 10.37 10.47 5.61 4.86 726 11.02 7.40 5.98 9.84 5.96 ZFMK 95.70 S. ollula, NW Cameroon 24.79 10.43 10.76 5.88 5.34 7.60 11.14 7.67 6.19 10.19 6.28 ZFMK 63.190 S. ollula, NW Cameroon 24.75 - 10.61 10.89 5.80 4.98 744 — 7.54 5.99 10.32 5.96 ZFMK 69.533 S. ollula, NW Cameroon 25.28 11.28 11.01 5.80 4.97 FAT 11.18. 7,72 6.21 10.35 3.92 ZFMK 99.683 5. ollula, NW Cameroon 24.10 10.20 10.83 5.97 o 746 11.36 7.61 5.85 10.76 6.42 ZFMK 2008.220 S. ollula, SE Cameroon 24.48 9.73 10.87 5.87 5.16 730 11.15 7.46 6.17 10.45 5.97 ZFMK 2008.226 S. ollula, Equ. Guinea 24.62 10.40 10.39 5.69 5.14 728 11.49 7.66 6.33 9.86 6.21 @E1736 17,00 16,00 15,00 14,00 13,00 12,00 11,00 10,00 9,00 8,00 7,00 3,00 total length of hind foot [mm] 4,00 5,00 length of metatarsal of digit Ill [mm] 6,00 7,00 Fig. 6. Relation between length of the metatarsal of digit III and total length of hind foot, as taken from digital X-ray images. | S. akaibei Mukinzi, Hutterer & Barriere, 2009, 2 S. johnstoni (Dobson, 1888), 3 S. konganensis Ray & Hutterer, 1996, 4 S. vulca- norum Hutterer & Verheyen, 1985, 5 S. granti Thomas, 1907, 6 S. n. sp. A, 7 5. morio (Gray, 1862), 8 S. camerunensis Heim de Balsac, 1968, 9 S. lunaris Thomas, 1906, 10 S. isabellae Heim de Balsac, 1968, 11 S. ollula, 12 S. n. sp. B, 13 S. megalura, 14 $. corbeti n. sp. Acknowledgements. We are grateful to Gerhard Nikolaus and Michael Barej who collected new material in Nigeria and Cameroon, and to Jean-Claude Mukinzi who collected relevant new material in the D. R. Congo. We also thank Paulina D. Jenkins for access to the collections of the British Museum, Carlos Lasso for access to his material at the Estacion Biologico Donana, and Dirk Rohwedder and Wolfgang Bischoff for their assistance with the photographs and X-rays. Gustav Peters kindly corrected the manuscript. Zusammenfassung. Aus dem súdóstlichen Hochland von Ni- geria wird eine neue Spitzmausart als Sy/visorex corbeti n. sp. beschrieben. Das der Beschreibung zu Grunde liegende Tier war zuvor als Sylvisorex ollula bestimmt worden, ist aber deutlich größer, besitzt einen längeren Schwanz, verlängerte Hinterfüße und einen sehr großen Schädel und repräsentiert damit die größ- te Art der Gattung. 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DUBEY, S., SALAMIN, N., RUEDI, M., BARRIERE, P., COLYN, M. & VOGEL, P. (2008): Biogeographic origin and radiation of the Old World crocidurine shrews (Mammalia: Soricidae) inferred from mitochondrial and nuclear genes. Molecular Phylogenet- ics and Evolution 48: 953 —963. EISENTRAUT, M. (1973): Die Wirbeltierfauna von Fernando Poo und Westkamerun unter besonderer Bedeutung der pleistozä- nen Klimaschwankungen für die heutige Faunenverteilung. Bonner zoologische Monographien 3: | -428. GOODMAN, S. M. & HUTTERER, R. (2004): A report on the shrews (Mammalia: Soricidae) of Monts Doudou, Gabon: Elevation- al distribution and preliminary insights into their ecology. Memoirs of the California Academy of Sciences 28: 93 —105. GOODMAN, S. M., HUTTERER, R. & NGNEGUEU, P. R. (2001): A report on the community of shrews (Mammalia: Soricidae) oc- curring in the Minkébé Forest, northeastern Gabon. Mamma- lian Biology 66: 22 —34. HEIM DE BALSAC, H. (1959): Nouvelles données sur la fauna so- ricidienne du Cameroun. Bonner zoologische Beiträge 26: 94 99, HEIM DE BALSAC, H. (1968): Nouvelles données sur la fauna so- ricidienne du Cameroun. Bonner zoologische Beiträge 26: 94-99. HEIM DE BaLsac, H. & LAMOTTE, M. (1957): Evolution et phylogénie des soricidés africains 2. La lignée Sy/visorex- Suncus-Crocidura. Mammalia 21: 15-49. HUTTERER, R. (1985): Anatomical adaptations of shrews. Mammal Review 15: 43-55. HUTTERER, R. (2005): Order Soricomorpha. In: Wilson, D. E. & Reeder, D. A. (eds.), Mammal species of the world: A taxonomic and geographic reference. Third edition. Johns Hopkins University Press, Baltımore. HUTTERER, R., DIETERLEN, F. & NIKOLAUS, G. (1992): Small mammals from forest islands of eastern Nigeria and adjacent Cameroon, with systematical and biogeographical notes. Bon- ner zoologische Beiträge 43: 393-414. HUTTERER, R. & HAPPOLD, D. C. D. (1983); The shrews of Nige- ria (Mammalia: Soricidae). Bonner zoologische Monographien 18: 1-79. HUTTERER, R., RIEGERT, J. & SEDLÁCEK, O. (2009): A tiny new species of Sy/visorex (Mammalia: Soricidae) from the Bamen- da Highlands, Cameroon. Bonner zoologische Beitráge 56: 151-157. HUTTERER, R. & SCHLITTER, D.A. (1996): Shrews of Korup Na- tional Park, Cameroon, with the description of a new Sylvi- sorex (Mammalia: Soricidae). Pp. 57-66, In: Contributions in Mammalogy: A Memorial Volume Honoring Dr. J. Knox Jones, Jr. Museum of Texas Tech University, il + 315 pp. JENKINS, P.D. (1984): Description of a new species of Sylvisorex (Insectivora: Soricidae) from Tanzania. Bulletin of the British Museum of Natural History (Zoology) 47: 65—76. KERBIS PETERHANS, J.C., HUTTERER, R., KALIBA, P. & MAZIBUKO, L. (2008): First record of Myosorex (Mammalia: Soricidae) from Malawi with description as a new species, Myosorex gnoskei. Journal of East African Natural History 97: 19-32. Lasso, C., HUTTERER, R. & RIAL, A. (1996): Records of shrews (Soricidae) from Equatorial Guinea, especially from Monte Alen National Park. Mammalia 60: 69-76. MEESTER, J. (1963): A systematic revision of the shrew genus Crocidura in southern Africa. Transvaal Museum Memoir 13: 1-127. MUKINZI, I., HUTTERER, R. & BARRIERE, P. (2009): A new species of Sy/visorex (Mammalia: Soricidae) from lowland forests north of Kisangani, Democratic Republic of Congo. Mam- malia 73: 130-134. QUEROUIL, S., HUTTERER, R., BARRIERE, P., COLYN, M., KERBIS PETERHANS, J. C. & VERHEYEN, E. (2001): Phylogeny and evo- lution of African shrews (Mammalia: Soricidae) inferred from 16s rRNA sequences. Molecular Phylogenetics and Evolution 20: 185-195. QUEROUIL, S., VERHEYEN, E., DILLEN, M. & COLYN, M. (2003): Patterns of diversification in two African forest shrews: Sylvi- sorex johnstoni and Sylvisorex ollula (Soricidae, Insectivora) in relation to paleoenvironmental changes. Molecular Phylo- genetics and Evolution 28: 24-37. Ray, J.C. & HUTTERER, R. (1996): Structure of a shrew commu- nity in the Central African Republic based on the analysis of carnivore scats, with the description of a new Sylvisorex (Mammalia: Soricidae). Ecotropica 1: 85-97. THOMAS, M. R. O. (1913): On African bats and shrews. Annals and Magazine of Natural History (8) 11: 314 -321. VOGEL, P. (1974): Note sur le comportement arboricole de Sy/vi- sorex megalura (Soricidae, Insectivora). Mammalia 38: 171-176. WOODMAN, N. & MORGAN, J. J. P. (2005): Skeletal morpholo- gy of the forefoot in shrews (Mammalia: Soricidae) of the genus Cryptotis, as revealed by digital X-rays. Journal of Mor- phology 266: 60-73. Authors’ addresses: Dr. Rainer HUTTERER, Zoologisches Forschungsmuseum Alexander Koenig, Section of Mammals, Adenauerallee 160, D-53113 Bonn, Germany, e-mail: r.hutterer.zfmk@uni-bonn.de; Dipl.Biol. Christian MONTERMANN, Brantestrasse 6, D-56746 Hohenleimbach, Germany. Received: 15.08.2009 Accepted: 30.08.2009 Buchbesprechung 209 GRIMMBERGER, Eckhard & RUDLOFF, Klaus, unter Mitar- beit von KERN, Christian (2009). Atlas der Säugetiere Eu- ropas, Nordafrikas und Vorderasiens. Natur und Tier Ver- lag, Münster. 496 S., 437 Karten, 1094 Abbildungen. IS- BN 978-3-86659-090-8. Preis 98,00 €. Auf dieses Werk haben Insider schon lange gewartet. Der Ehrgeiz, alle Säugetierarten Europas, Nordafrikas und Vorderasiens in farbigen Lebendfotos zu dokumentieren, führte die beiden Verfasser schon seit vielen Jahren in zahlreiche Länder Europas, Nordafrikas und Asiens. Eck- hard Grimmberger hat seine fotografische Meisterschaft bereits früher mit einem Buch über die Fledermäuse Eu- ropas (SCHOBER & GRIMMBERGER 1998), das in vier Sprachen übersetzt wurde, unter Beweis gestellt. Klaus Rudloff hat als Kurator im Tierpark Berlin Zugang zu vie- len seltenen Säugetieren gehabt und diese im Foto doku- mentiert. Was nicht durch eigene Tätigkeit im Foto zu bekommen war, ergänzten viele Zoologen aus der ganzen Welt. Das vorliegende Buch erweitert den geografischen Rah- men über Europa hinaus und schließt die nordafrikanis- chen Länder von Marokko bis Ägypten und die vorderasi- atischen Länder von Jordanien bis Aserbaidschan ein. Möglich wurde dies auch durch das taxonomische Ref- erenzwerk „Mammal Species of the World“ (WILSON & REEDER 2005), das einen leichten Zugang zur Nomen- klatur und Verbreitung aller Säugetierarten erlaubt. Im Fahrtwind dieses Werkes sprossen seit 2005 etliche Säugetierführer aus dem Boden, von denen der vor- liegende einer der originellsten ist. Das Buch beginnt mit einleitenden Kapiteln (Einführung, Prinzipien der zoologischen Systematik, Die Welt der Säugetiere, Säugetierschutz, Zur Benutzung des Buches), die für Laien verständlich sind, aber auch Studenten der Zoologie noch genügend Stoff vermitteln könnten. Den Hauptteil des Buches bilden mit über 400 Seiten Artkapi- tel von mehr als 400 Säugetierarten, die der Systematik der bereits genannten Weltliste von Wilson & Reeder fol- gen. Jedes Artkapitel enthält neben dem wis- senschaftlichen Namen einen Trivialnamen in deutscher, englischer, französischer, russischer und spanischer Sprache, eine kurze Beschreibung des Tieres, sodann Angaben zu Verbreitung, Lebensraum und Lebensweise, Schutzstatus und weiterführender Literatur, begleitet von einer farbigen Verbreitungskarte und einem oder mehreren Farbfotos. Diese Fotos sind durchweg von hoher Qualität und bilden einen Schatz, wıe er bisher in dieser Art nicht verfügbar war. Viele seltene Säugetierarten sieht der Leser hier zum ersten Mal, so z. B. Meriones grandis, Gerbil- lis hoogstraali, Mesocricetus raddei, Microtus anatolicus und andere. Die Vielfalt der Nagetiere wird erst durch Lebendaufnahmen richtig erfahrbar. Die Schönheit und Vielfalt der Pferdespringer, Springmäuse, Rennmäuse, Hamster oder Blindmäuse ist überwältigend. Das gilt aber auch für die anderen Gruppen, seien es Igel, Spitzmäuse, Maulwürfe, Fledermäuse, Hasenartige, Huftiere, oder Kar- nivoren, je nach Vorliebe. Im Buch nicht berücksichtigt sind die Walartigen. Originell sind auch die Anhänge, die die Feldbestimmung erleichtern sollen. Sie zeigen Schwänze von Springnagern und Rennmäusen, Hinterfuß-Sohlenschwielen von Nagetieren, Zahn- und Schädelmerkmale von einigen Kleinsäugern, sowie die Spiegel von Hirschen. Alle An- hänge zeigen Detailansichten in Farbfotos. Ein Liter- aturverzeichnis und ein Register beschließen diesen wun- derbaren Atlas, der uneingeschränkt empfohlen werden kann. Das Buch ist für ein breites Leserpublikum von In- teresse, und auch der Spezialist wird Bilder und Informa- tionen finden, die für ıhn neu sind. Eine Ausgabe in en- glischer Sprache würde den Leserkreis noch um ein vielfaches erweitern. SCHOBER, W. & GRIMMBERGER, E. (1998): Die Fleder- mäuse Europas. 2. Auflage. Ulmer, Stuttgart. WILSON, D. E. & REEDER, D. M., eds. (2005), Mammal species of the world: A taxonomic and geographic ref- erence. Third edition, 2 volumes. The Johns Hopkins University, Baltimore. Rainer HUTTERER Zoologisches Forschungsmuseum Alexander Koenig, Bonn 210 Buchbesprechung TEMPLE, Helen J. & CUTTELOD, Annabelle (compilors) (2009). The Status and Distribution of Mediterranean Mammals. Gland, Switzerland and Cambridge, UK, IUCN. vii + 32 pp. ISBN 978-2-8317-1163-8. Available from IUCN, Publications Services, 28 Rue Mauverney, 1196 Gland, Switzerland. This booklet presents the results of a Mediterranean Mam- mal Red List workshop held in Malaga, Spain, in 2007. This workshop was embedded in the Mediterranean Bio- diversity Assessment and in Global Mammal Assessment activities of the IUCN, in cooperation with experts from around the world. The aim of this workshop was to as- sess the taxonomy, distribution, and conservation status of all mammal species living in the 31 countries borde- ring the Mediterranean Basin from Albania to Turkey, and including the Atlantic Islands and Western Sahara. A to- tal of 297 species was assessed, excluding species of mar- ginal occurrence in the region. 16.5% of the Mediterranean mammals are threatened with extinction at the regional scale, and a further 8% are Near Threatened. By comparison, 56% of dolphins and whales, and 5% of birds are threatened with extinction. More than one-quarter (27%) of the Mediterranean mam- mals have declining population, and 31% are stable. The mammals of the Magreb region were included in the as- sessment, and this region holds a large number of ende- mic species, unique to the Mediterranean and found nowhere else. In the Sahara, species richness is relative- ly low, but a high proportion of Saharan species are threa- tened. The booklet includes many more interesting data and maps showing the geographic distribution of species richness and endemic species richness in the Mediterranean Regi- on. An appendix lists all mammal species considered and their Red List status. Copies of this useful summary can be obtained from IUCN. Rainer HUTTERER Zoologisches Forschungsmuseum Alexander Koenig, Bonn Bonner zoologische Beiträge Instructions to Authors Bonner zoologische Beiträge publishes original papers, reviews, and scientific notes covering the whole field of zoology, with em- phasis on biodiversity, biogeography, history of zoology, morphol- ogy, phylogenetics, and taxonomy. The material must not be sub- mitted for publication elsewhere. All manuscripts will be peer-ref- ereed. Final acceptance of items for publication is the responsibil- ity ofthe corresponding editor dealing with the respective taxon (see list of editors). Manuscripts not compatible with the following guidelines will be returned. 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ILL 3 9088 01533 1 2 Special Issue: New African Mammals HUTTERER, Rainer & PETERS, Gustav: 131 Editorial: New species and records of mammals from Africa GOODMAN, Steven M.; RAHERIARISENA, Martin & JANSA, Sharon A.: 133 A new species of Eliurus Milne Edwards, 1885 (Rodentia: Nesomyinae) from the Reserve Spéciale d’Ankarana, northern Madagascar HUTTERER, Rainer; RIEGERT, Jan & SEDLACEK, OndFej: 151 A tiny new species of Sy/visorex (Mammalia: Soricidae) from the Bamenda Highlands, Cameroon DENys, Christiane; Missoup, Alain Didier; TCHIENGUE, Barthelemy; ACHOUNDONG, Gaston; 159 EKOBO, Atanga; BILONG Bilong, CHARLES Felix; LEMBE, Dieudonné MASSOMA & NICOLAS, Violaine: Altitudinal distribution of and anthropogenic influence on small mammal assemblages on Mount Kupe, SW Cameroon KERBIS PETERHANS, Julian; STANLEY, William T.; HUTTERER, Rainer; DEMOS, Terrence C. 175 & AGWANDA, Bernard: A new species of Surdisorex Thomas, 1906 (Mammalia, Soricidae) from western Kenya DIETERLEN, Fritz: 185 Climbing mice of the genus Dendromus (Nesomyidae, Dendromurinae) in Sudan and Ethiopia, with the description of a new species HUTTERER, Rainer & MONTERMANN, Christian: Fi ; 201 A large new species of Sy/visorex (Mammalia: Soricidae) from Nigeria and the first record of Sylvisorex ollula from the country Buchbesprechungen / Book Reviews GRIMMBERGER, Eckhard & RUDLOFF, Klaus, unter Mitarbeit von KERN, Christian (2009) 209 Atlas der Säugetiere Europas, Nordafrikas und Vorderasiens (R. HUTTERER) TEMPLE, Helen J. & CUTTELOD, Annabelle (compilors) (2009) 210 The Status and Distribution of Mediterranean Mammals (R. HUTTERER) Titelbild/ Cover illustration: Eliurus carletoni Goodman, Raheriarisena & Jansa (Mammalia: Rodentia), a new endemic rodent from Madagascar (see contribution of GOODMAN et al., pp. 133-149) | De Agamis Bonner zoologische Beiträge On Agamid Lizards Proceedings of ‘DEAGAMIS — 15! International Symposium on Agamid Lizards’ held in the Zoologisches Forschungsmuseum Alexander Koenig in Bonn, February 22-24, 2008 Guest Editors: Philipp Wagner, Thomas M. Wilms & Wolfgang Böhme Herausgegeben vom Zoologischen Forschungsmuseum Alexander Koenig, Bonn +f Leibniz Gemeinschaft « NRW. (2007 109) Bonner zoologische Beiträge werden publiziert im Eigenverlag Zoologisches Forschungsmuseum Alexander Koenig, Adenauer- allee 160, D-53113 Bonn (Germany). Die Zeitschrift erscheint mit vier Heften im Jahr, zum Preis von 11,50 € je Heft bzw. 46,- € je Band incl. Versand. Korrespon- denz betreffend Abonement, Kauf oder Tausch bitten wir zu richten an die Bibliothek, Zoologisches Forschungsmuseum Alex- ander Koenig, Adenauerallee 160, D-53113 Bonn (Germany); E-Mail: d.steinebach.zfmk@uni-bonn.de. © Zoologisches Forschungsmuseum Alexander Koenig (ZFMK), Bonn, Germany. ISSN 0006-7172 Satz: Eva-Maria Levermann, Kaiserstr. 151, D-53113 Bonn (Germany); E-Mail: emlevermann@netcologne.de. Druck: JFoCARTHAUS, Bonn, Germany. Bonner zoologische Beiträge Editorial Board (in brackets editorial competence) Dr. Fabian HERDER (editor-in-chief, Ichthyologie) Tel.: +49 228- 9122 255, Fax: +49 228-9122 212; E-Mail: f.herder.zfmk(@uni-bonn.de. Dr. Netta DORCHIN (Diptera), Tel.: +49 228-9122 292, Fax: +49 228-9122 212; E-Mail: n.dorchin.zfmk@uni-bonn.de. Dr. Renate VAN DEN ELZEN (Vertebrata, except Mammalia), Tel.: +49 228-9122 231, Fax: +49 228-9122 212; E-Mail: r.elzen.zfmk(wuni-bonn.de. Dr. Bernhard A. HUBER (Invertebrata, except Insecta), Tel.: +49 228-9122 294, Fax: +49 228-9122 212; E-Mail: b.huber.zfmk(@uni-bonn.de. Dr. Gustav PETERS, (Mammalia, bioacustics), Tel.: +49 228-9122 262, Fax: +49 228-9122 212; E-Mail: g.peters.zfmk(wuni-bonn.de. Dr. Bradley S. SINCLAIR (language advisor), Entomology Laboratory - CFIA, K.W. Neatby Bldg., C.E.F. 960 Carling Ave., Ottawa, ON, Canada KIA 0C6; E-Mail: sinclairb@inspection.ge.ca. Dr. Dieter STUNING (Insecta, except Coleoptera and Diptera), Tel.: +49 228-9122 220, Fax: +49 228-9122 212; E-Mail: d.stuening.zfmk@uni-bonn.de. Editorial office: Adenauerallee 160, D-53113 Bonn, Germany. Advisory Board Prof. Dr. Miguel Angel ALONZO-ZARAZAGA, Museo nacional, Ciencias Naturales, E-28006 Madrid; E-Mail: zarazaga@@mnen.csis.es. Prof. Dr. Ulrike Aspock, Naturhistorisches Museum, 2. Zoologische Abteilung (Insekten), Postfach 417, A-1014 Wien; E-Mail: ulrike.aspoeck(@nhm-wien.ac.at. Prof. Dr. Paolo Aupisio, Universita di Roma “La Sapienza”, Dipl. Biol. Anim e dell’ Uomo (Zool.), Viale dell’ Universita 32, 1-00185 Roma, Te.: +39 6-49914744; E-Mail: paolo.audisio@@uniromal.it. Prof. Dr. Aaron M. BAUER, Villanova University, Department of Biology, 800 Lancaster Avenue, Villanova, PA 19085-1699, USA, Tel: +1-610-519-4857, Fax: +1-610-519-7863; E-Mail: aaron.bauer(@villanova.edu. Dr. Jürgen HAFFER, Tommesweg 60, D-45149 Essen, Tel.: +49 201-710426; E-Mail: j.haffer@web.de. Dr. Jeremy D. HoLLoway, The Natural History Museum, Department of Entomology, Cromwell Road, London, SW7 5BD, U.K.; E-Mail: j.holloway@nmh.ac.uk. Dr. Marion KOTRBA, Zoologische Staatssammlung, Diptera, Miinchhausenstr. 21, D-81247 München, Tel.: +49 89-8107 147, Fax: +49 89-8107 300; E-Mail: marion.kotrba@zsm.mwn.de. Prof. Dr. Boris KRYSTUFEK, Slovenian Museum of Natural His- tory, P. O. Box 290, SI-1001 Ljubljana; E-Mail: boris.krystufek@zrs.upr.si. Pro. Dr. Sven O. KULLANDER, Swedish Museum of Natural His- tory, Department of Vertrebrate Zoology, P. O. Box 50007, SE-104 05 Stockholm; E-Mail: sven.kullander@nrm.se. Prof. Dr. Steven PERRY, Rheinische Friedrich-Wilhelms-Univer- sitát, Institut fúr Zoologie, Poppelsdorfer Schloss, D-53115 Bonn, Tel: +49 228-73 3807; E-Mail: perry@uni-bonn.de. Dr. Wolfgang SCHAWALLER, Staatliches Museum fúr Naturkunde, Rosenstein 1, D-70191 Stuttgart, Germany, Tel.: +49 711-8936 221, Fax: +49 711-8936 100; E-Mail: schawaller.smns@naturkundemuseum-bw.de. Dr. W. David Sissom, Dept. of Life, Earth and Environmental Sciences, W. Texas A. & M. University, WTAMU Box 60808, Canyon, Texas 79016, USA; E-Mail: dsissom@wtamu.edu. Dr. Miguel VENCEs, Technische Universität Carolo-Wilhelmi- na, Zool. Inst., Abt. Evolutionsbiol., Mendelssohnstr. 4, D-38106 Braunschweig, Tel.: +49 531-391 3231, Fax: +49 531-391 3222; E-Mail: m.vences@tu-braunschweig.de. Prof. Dr. Heike WÄGELE, Rheinische Friedrich-Wilhelms-Uni- versität, Institut für Evolutionsbiologie und Ökologie, D-53121 Bonn, Tel.: +49 228 73 5159, Fax: +49 234-322 4114; E-Mail: hwaegele@evolution.uni-bonn.de. Dr. Erich WEBER, Eberhard-Karls-Universität, Zoologische Schausammlung, Sigwartstr. 3, D-72076 Tübingen, Germany; E-Mail: erich.weber@uni-tuebingen.de. | PREFACE The ıguanıan lizard family Agamidae exists in more than 350 species which are grouped in about 45 genera. Agamid lizards are distributed through the Mediterranean, subtropical and tropical regions of the Old World with the ex- ception of Madagascar. They occupy a great variety of biotopes ranging from deserts through steppes and savannas to tropical rain forest. The adaptations to specific habitats have led to an extraordinary morphological and ecological di- versity. Agamids may be wide-ranging or confined to biodiversity hotspots, oth- ers are relictual endemics to small areas. Many of them, however, are under an- thropogenic pressure, either by habitat loss or by direct persecution, thus being important for the conservation of biodiversity on a global scale. New, intensive studies with multiple methods including molecular genetic approach- es have shown that the taxonomic diversity of agamids, in Africa as well as in SE Asia, 1s still considerably underestimated. Also some phylogenetic hypotheses and concepts of the last decades have been challenged by new, current research. It was therefore overdue that the herpetologists working on various aspects of agamid sys- tematics and biology in various parts of the world were brought together in order to present their results and ideas to each other and to create a new network of coopera- tive research on agamids (in Latin “de agamis””). It was the idea and initiative of the Herpetology Section of the Zoologisches Forschungsmuseum Alexander Koenig (ZFMK), in particular of the young researcher Philipp WAGNER who devoted a part of his PhD thesis to these fascinating lizards, to in- = vite colleagues from many countries to a first international symposium “DeAgamis”, and = the resonance was very good. A part of the scientific outcome of this symposium is col- = lected in the present issue of Bonner zoologische Beiträge. I hope this issue will stimulate colleagues to attend a second symposium DeAgamis which I am inviting to St. Petersburg in August 2010. St. Petersburg, October 2009 Prof. Dr. NATALIA B. ANANJEVA EDITORIAL De Agamis The Zoologisches Forschungsmuseum Alexander Koenig (ZFMK) has established a long tradition in organizing and hosting international symposia. Most of these dealt with tropical zoology and often focussed on Africa and on vertebrates. Some special symposia were devoted to herpetology, the first of these to the reptiles of the Canary Islands, three further ones to all aspects of monitor lizard research, and the last one to the world’s widest-distributed lizard, the lacertid Zootoca vivipara. In this tradition, one of us (Philipp WAGNER) developed the idea of a First Internation- al Symposium on Agamid Lizards (his favourite squamate group) which was held at ZFMK on February 22-24, 2008 under the name “DeAgamis”. The idea to bring the world’s leading specialists on agamid lizards to Bonn and to de- velop new cooperation projects had a very positive echo. 50 researchers from 9 coun- tries (Australia, Czech Republic, England, Germany, The Netherlands, Russia, Switzer- land, the USA and Vietnam) attended, and a total of 20 talks and posters which covered a broad variety of topics on nearly all agamid groups were presented to the participants. Because not all contributors were able to submit their presentations to the present volume, their abstracts are summarized in an appendix in order to inform the readers who did at- 1 tend the meeting, and about the topics presented and discussed. 6 We wish to thank the Deutsche Forschungsgemeinschaft (DFG) for providing the funds that enabled us to invite speakers from abroad and even overseas. Moreover, we are indebted to our motivated and effective student crew, especially to Michael BAREJ, Monique HÓLTING and Julia WURSTNER. 214 Editorial Partcipants of DeAgamis in Bonn, February 23rd, 2008. Finally, we are happy to inform the readers that there will be a second International Symposium on Agamid Lizards (DeAgamis II) to be organized by our friend and colleague Prof. Dr. Natalya B. ANANJEVA in St. Petersburg in 2010 which lets us hope that our first symposium DeA gamis might evolve into a series of meetings. Bonn, October 2009 — PHILIPP WAGNER, THOMAS WILMS & WOLFGANG BÓHME Bonner zoologische Beiträge Band 56 Heft 4 Seiten 215-223 Bonn, November 2009 Studies on African Agama V. On the origin of Lacerta agama Linnaeus, 1758 (Squamata: Agamidae) Philipp WAGNER!*, Thomas M. WiLms?, Aaron BAUER? & Wolfgang BÖHME! ! Zoologisches Forschungsmuseum A. Koenig, Adenauerallee 160, D-53113 Bonn, Germany philipp.wagner.zfmk@uni-bonn.de; w.boehme@uni-bonn.de 2 Zoologischer Garten Frankfurt, Bernhard-Grzimek-Allee 1, D-60316 Frankfurt thomas.wilms@stadt-frankfurt.de 3 Villanova University, 800 Lancaster Avenue, Villanova, Pennsylvania 19085, US “corresponding author Abstract. Herein we present our strategy to preserve the nomenclatural stability of the widespread and common Afrotropical lizard Agama agama (Linnaeus, 1758). We recognized several proposed syntypes, belonging to a va- riety of species, 1.e., 4gama agama, Agama atra and Tropidurus plica. But we have shown that these specimens do not belong to the type material. However, if they were included in the type material the selection of the e.g. American syntype, Tropidurus plica — a tropidurine iguanid, as lectotype would result in taxonomic chaos, as Aga- ma agama is an African species and includes in its synonymy the type species of the type genus of the family Agamidae. The illustrated types in SEBA (1734) are recognized by us as a variety of agamid lizards with a type lo- cality encompassing the New and the Old World and not simply ‘America’, as given by LINNAEUS. Morphologically, Agama agama is highly variable making it impossible to assign the syntype that agrees with the current concept of Agama agama to a geographic population, particularly as the only character, viz. the colour pat- tern, is no longer discernible in this specimen. This endangers the stability of the known subspecies of Agama aga- ma. We therefore designate a neotype for Lacerta agama Linnaeus, 1758 herein. Keywords. Nomenclature; taxonomy; neotype; Squamata; Agamidae; Agama agama; African rainbow lizard; type locality; Africa. INTRODUCTION The genus Agama Daudin, 1802 is one of the most wide- ly distributed lizard genera within Africa, although the re- lationships between the species in the genus are still poor- ly known. Agama is a highly taxonomically complex genus and most of the taxa are very similar and highly variable in pholidosis and only identifiable by the nuptial colouration of males (MCLACHLAN 1981, JACOBSEN 1992, WAGNER 2007). Therefore, it is often not possible to as- sign a single specimen with colouration lacking to a par- ticular species or subspecies. This situation causes prob- lems in the determination of type material to assess the validity of the different taxa. Thus, in many species and subspecies validity is doubtful (e.g., Agama bocourti Rochebrune, 1884; Agama cornii Scortecci, 1928; A. pic- ticauda Peters, 1877; A. congica Peters, 1877). Past gener- ic reviews have only been attempted for some species groups (BOULENGER & POWER 1921, GRANDISON 1956, 1968) and most workers have considered it point- less to designate lecto- or neotypes to stabilize the nomen- clature of the taxa. As an example, GRANDISON (1968) des- ignated a lectotype of Agama benueensis Monard, 1951 (today recognized as a subspecies of Agama doriae Boulenger, 1885; see Moopy & BOHME 1984), thereby sta- bilizing another Agama species because several individ- uals of the type series were identified as 4. paragama Grandison, 1968. However, the situation has changed and several authors are now working on a broad scale phyloge- ny of the entire genus (WAGNER & LEACHE, own data). For this analysis it is essential to clarify the taxonomic status of Agama agama (Linnaeus, 1758) because many taxa are assigned to this species as synonyms or subspecies. Even in recent years several former subspecies of 4. agama have TAB. CVII BN Y a Fig. 1. Plate 107 published in Thesaurus I by Sera (1734) showing three lizards described as “Salamandra Americana re parte Lacertum referens, amphibia; mas”. , posterio- Bonner zoologische Beiträge 56 217 14. LACERT A cauda tereti longa, pedibus pentada- étylis, dorfo antice denticulato, collo capite- que pone aculeato. Salamandra americ. Lacerte emula altera. Seb. thef.1. p.170. 1.107. f.3. Salamandra amer. am- phibia. Seb. rhef-1. p. 169. 2.107. f-1,2. CAPUT ovatum, fquamis anteriora verfus imbri- catum, pone juxta aures aculeis inxqualibus fpinofum, quod fingulare in hac fpecie. x um Fig. 2. Description of ‘Lacerta amphibia’ in the *Amoenitates academic” by LINNAEUS (1749, Vol. 1, p. 288). been recognized as species. BÓHME et al. (2005) have el- evated the Agama lionotus Boulenger, 1896 complex (Agama lionotus lionotus Boulenger, 1896, Agama l. el- gonis Lönnberg, 1922, Agama I. dodomae Loveridge, 1923, Agama l. ufipae Loveridge, 1932) leaving the true Agama agama to full species rank, with the following sev- eral nominal subspecies of its own: Agama agama aga- ma Linnaeus, 1758; Agama a. africana Hallowell, 1844 (type locality: Liberia); Agama a. boensis Monard, 1940 (type locality: Madina Boé, Guinea-Bissau) and Agama a. mucosoensis Hellmich, 1957 (type locality: Mucoso near Dondo, Angola). 4gama a. savattieri Rochebrune, 1884 [type localities: Casamance (Senegal?), Mélacorée (Guinea?), Albréda (The Gambia), Bathurst (FBanjul, The Gambia] has been synonymized with 4. agama africana by GRANDISON (1968). Even the status of these subspecies remains unclear and is dependent on the definition of A. agama. Therefore, it is essential to consult the original description by LINNAEUS (1758), the type specimens and previous descriptions of the species. The results of this search are herein present- ed. 1. PRE-LINNAEEN (BEFORE 1758) NAMES ‘Salamandra americana Seba, 1734’ In his Thesaurus SEBA (1734) published a plate (thes. I, p. 170, t. 107) depicting three lizards (see fig. 1) which he described as “Salamandra Americana, posteriore parte Lacertum referens, amphibia; mas’. The Thesaurus is published in two separate editions, one in French and Latin, the other one in Dutch and Latin. We herein rec- ognize the Dutch edition because this would be the text that probably most accurately reflects SEBA’s own lan- guage. In the Dutch part of the accompanying text he de- scribed the species shown in figure one of the plate as the LACERTA cauda tereti longa, peaibus pentataclyhs , dorfo an- tice denticularo, collo capiteque pone acıleatıs. Amen. Acad. ¿.p.288 Sy, nat. 36. MER. Salamandia americana amphibia. Seb.thefr 1.269. n 207.2. Salamandra amer. lacerte emula altera. Seb. thef.t. P«170. £.107-f-}. Habitat in AMERICA. amphibia. Fig. 3. ‘Lacerta amphibia’ published in the ‘Museum Adolphi Friderici’ (LINNAEUS 1754, p. 44) with the first mention in of the locality ‘Habitat in America’. ‘West-Indische Salamander’ (= West Indian salamander), half salamander, half lizard. The specimens shown in fig- ures two and three of the same plate (herein fig. 1) were also recognized as Salamandra americana but the figure captions were ‘Wyfie van den voorgaanden Salamander’ (= the female of the above mentioned salamander) (fig- ure two) and ‘Ander soort van West-Indischen Salaman- der als de voregi’ (= another species as the previous one of the west Indian salamander) (figure three). All the subsequent descriptions of the species refer to this plate and the text mentioned. The name ‘Salamander americana” is mentioned later by many authors (see be- low). However, confusions among agamids, geckos and sala- manders were common during the 18th and early 19th cen- turies. As an example, a gecko, today known as Hemi- dactylus platyurus, was described in an agamid genus as Stellio platvurus by SCHNEIDER (1792). In the same work SCHNEIDER (1792) also described Stellio fimbriatus from Madagascar, today known again as a member of the geck- onid genus Uroplatus. Also Trapelus savignyi AUDOUIN, 1827, described in an agamid genus but in a gecko sec- tion of the text is today known as synonym of the gecko Stenodactylus sthenodactylus (Lichtenstein, 1823) (WAG- NER & CROCHET 2009). ‘Lacerta amphibia Linnaeus, 1749’ In the ‘Amoenitates academice’, LINNAEUS’ first impor- tant contributions as collected theses in seven volumes on systematics, he (LINNAEUS 1749, Vol. 1, p. 288; see fig. 2) described the lizard which was shown in SEBA (1734) as ‘Lacerta cauda tereti longa, pedibus pentadactylis, dor- so antice denticulato, collo capiteque pone aculeato” lat- er known as ‘Lacerta amphibia’ and synonymised the 218 Philipp WAGNER et al.: Studies on African Agama V. On the origin of Lacerta agama Linnaeus, 1758 Agsma, 30. L, cauda tereti longa, aculeato. Amun. «cad. 1. p. 288, Muf, Ad. Er. Y. p. 44 Seb. mel. 1. 2. 107, fo Y, Habitat im America. Color corporis pallidus. Abdomen minimo firiatum, collo' fupra capiteque poftice 22/3: Fig. 4. Description of Lacerta agama in the nomenclaturally binding 10'h edition of LINNAEUS 1758 (p. 207). name “Salamandra americana’. In naming his species LIN- NAEUS referred to the plate published by SEBA (1734, plate 107; see fig. | herein) and obviously followed the sugges- tions of SEBA (1734), considering this species half sala- mander and half lizard. But LINNAEUS (1749) also referred to the differences between the specimens shown in the fig- ures in SEBA (1734) as he synonymised figure three as Salamandra americana and figures one and two as Salamandra americana amphibia. Several years later, LIN- NAEUS (1754, p. 44) published his ‘Museum Adolphi Frid- erici”. Herein, he added a type locality as “Habitat in America’ and again he referred to two taxa (Salamandra americana amphibia, fig. 1 £ 2; Salamandra americana lacerate aemula, fig. 3). 2. LINNAEUS (1758) Lacerta agama Linnaeus, 1758 In the 10th and currently still binding edition of his guid- ing work, LINNAEUS (1758, p. 207) again re-named “Lac- erta amphibia’ to the currently valid Lacerta agama Lin- naeus, 1758 (see fig. 4). In contrast to the descriptions be- fore, he referred all the three images of plate 107 in SE- BA (1734) to this monotypic species and mentioned again ‘America’ as type locality. In his description, LINNAEUS (1758) did not refer to other images or vouchers of a mu- seum collection. Therefore, the individuals which are the basis of the illustrations in SEBA (1734) are the syntypes, because the ‘International Code of Zoological Nomencla- ture’ (ICZN 1999) does not recognize “iconotypes” and no other types are mentioned. However, QUENSEL, director of the Swedish Academy of Science from 1799 to 1806, also mentioned three specimens (QUENSEL 1802, QUENSEL in ANDERSSON 1900) from the collection of the Museum Adolphi Friderici in Drottningholm and noted ‘that they are identical with Linnaeus’s Lacerta amphibia’ (ANDER- SSON 1900, p. 11). However, the use of ‘identical’ here on- ly means identical to species not identical type specimens. ANDERSSON (1900) examined these specimens and iden- tified two as ‘true Agama colonorum Daudin, 1802’ and the last as ‘Uraniscodon plica Linnaeus, 1758’. A recent examination by us of the three specimens (Museum Adol- phi Friderici: NRM 107, 108, 112; see fig. 5 herein) re- vealed that they represent three different valid species: Agama cf. agama (LINNAEUS, 1758), Agama atra Daudin, 1802 and Tropidurus plica (Linnaeus, 1758). But two of these specimens are without doubt not identical with the individuals illustrated in SEBA (1734). As it is obvious, the lizards shown in figures one and two clearly have seg- mented tails, which are not only lacking in the three mu- seum vouchers, but also in the genus Agama in general. These segmented tails, which are arranged in whorls, are typical for the species of the genera Acanthocercus Fitzinger, 1843 and Laudakia Gray, 1845 (both Agami- dae) occurring in Africa, Asia and Europe. Only the spec- imen of figure three shows a tail typical for Agama, but also for the South American genus Tropidurus. These dif- ferences were also recognized by DUMERIL € BIBRON (1837) who referred the specimens shown in figures one and two of plate 107 in SEBA (1734) to Stellio vulgaris Sonnini & Latreille, 1802 (today known as Laudakia stel- lio vulgaris) and erroneously synonymised Lacerta am- phibia with Agama colonorum, as they only referred fig- ure three to Agama colonorum. However, it is obvious by comparing the NRM specimens with the remaining image in SEBA (1734) that it is not iden- tical with one of the specimens. The atra specimen has a very different colouration, whereas the plica one has the mouth closed, but similar in scalations to the figured spec- imen. The cf. agama specimen is not identical in head sca- lation and proportion. Consequently, the NRM specimens are not identical with the lizards shown in SEBA (1734) and are not type speci- mens of Lacerta agama Linnaeus, 1758. As a nomenclat- ural stabilization of this name is indispensable for further research, a neotype is designated by us (see below). However, even if e.g. the most similar NRM voucher (the T. plica-specimen) to the remaining image were the name- bearing type of Lacerta agama, it would require suppres- sion because it would destabilise two well known species, two genera and two families. Thus Tropidurus plica (as currently known) would become Lacerta agama. However, Agama colonorum, a synonym of Lacerta agama today known as Agama agama, is al- so the type species of the genus Agama. Therefore, Tropidurus plica and closely related taxa like 7. lumaria and 7. umbra would become members of the genus Aga- ma. Bonner zoologische Beiträge 56 219 Further, the genus Agama is the type genus of the lizard family Agamidae. This would mean that these Old World lizards would lose their name to a New World genus, which would cause a taxonomic chaos and 1s therefore ob- viously unacceptable. According to article 75.6 ofthe ICZN this approach should not be taken into consideration because it endangers the stability of the well known species 7. plica, the genus Aga- ma and the family Agamidae. Remarks on the identity of the NRM specimens As shown above, two of the specimens are not identical with Agama agama as they are identified as 4. atra and Tropidurus plica. The remaining voucher (fig. 5c) is a true Agama species, and probably Agama agama itself. It ıs obvious from figure 5c that the specimen is in good con- Fig. 5. Formerly supposed syntypes of Lacerta agama Linna- eus, 1758 in the Museum Adolphi Friderici (NRM 107, 108, 112). A= Tropidurus plica (Linnaeus, 1758); B= Agama atra Daudin, 1802; C= Agama cf. agama (Linnaeus, 1758). dition but completely lacking colouration. As previously noted, species of the genus Agama, and especially Aga- ma agama, are characterized by a high variability of scale counts (BOULENGER & POWER 1921, THYS VAN DEN AU- DENAERDE 1963, GRANDISON 1968, MOODY & BÖHME 1984, BÖHME et al. 2005) and therefore nearly all taxa are determined by aspects of colouration of adult males (McLACHLAN 1981, JACOBSEN 1992, WAGNER 2007). Thus, it is not possible to assign this voucher to a specif- ic Agama agama population or subspecies and even not clearly to Agama agama itself. Remarks on the erroneous type locality ‘America’ & the nomen Agama Today it is obvious that the type locality “in America’ is erroneous and does not correspond with the actual distri- bution of the species, genus and family. Surprinsingly, Agama colonorum Daudin, 1802, the first available syn- onym of Agama agama, was also described from ‘? Amerique méridionale”, ‘Cuba’ and “Jamaique’. DAUDIN (1802) suggested this name for Lacerta agama Linnaeus, 1758 and mentioned several names (e.g. Lacerta agama; Iguana cordylina Laurenti, 1768; ‘Salamandra americana Seba, 1734”; Lacertus major Sloane; Guana lizard Brown; Blue lizard Edwards; L’agame Daubenton; L’agame ou caméléon de Mexique Stedman) as synonyms. Addition- ally, DAUDIN (1802) also mentioned aspects of the linguis- tic usage of the name ‘agama’ and mentioned that the name is used for a variety of different lizard species in South America. However, the source of the name ‘agama’ is from the Ewe language of the Kwa group, spoken in Togo, Benin and Ghana (BAUMANN 1936, WESTERMANN 1954). DUMERIL & BIBRON (1837) were the first who rec- ognized that the type locality is in error and mentioned a distribution of the species “sur la cóte de Guinée et au Senegal’ (= at the coast of Guinea and in Senegal). However, the consideration of the type locality as “in er- ror’, the simplest solution, does not solve the problem. Ac- cording to article 76 of the code, the ‘type locality of a nominal species-group taxon 1s the geographical place of capture, collection or observation of the name-bearing type; if there are syntypes and no lectotype has been des- ignated, the type locality encompasses the localities of all of them’. As the syntypes are not available, the type lo- cality can only be concluded from the identified lizard gen- era shown on the plate in SEBA (1734). MERTENS (1938) also recognized the problem and restrict- ed it to ‘Cameroon’, but without the designation of a lec- to- or neotype. According to article 76 of the code, MERTENS’ (1938) restriction was invalid and the type lo- cality thus still not clarified. ho 159) o Philipp WAGNER et al.: Studies on African 4gama V. On the origin of Lacerta agama Linnaeus, 1758 Fig. 6a. Neotype of Lacerta agama Linnaeus, 1758 (ZFMK 15222): a = image of the living neotype; b = complete lateral, dorsal (arrow indicates the place of scales used for SEM images, see fig. 6c), ventral body views. Fig. 6b. Neotype of Lacerta agama Linnaeus, 1758 (ZFMK 15222): details of head scalation. Bonner zoologische Beiträge 56 [597 159) Fig. 6c. Neotype of Lacerta agama Linnaeus, 1758 (ZFMK 15222): SEM images of dorsal scales. Location of the scales is indi- cated by an arrow in fig. 6a. Description of Lacerta agama Linnaeus, 1758 Agama agama (Linnaeus, 1758) 1734 Salamandra americana Seba, Naturalium Thesauri, thes. 1, Tome 1, Pl. 107, fig. 1-3. 1749 Lacerta amphibia Linnaeus, Amoenitates academi- ce, vol. 1: 288. 1754 Lacerta amphibia Linnaeus, Museum Adolphi Fri- derici: 44. 1758 Lacerta agama Linnaeus, Syst. Nat., Ed. 10, 1: 207. 1802 Agama colonorum Daudin, Hist. Nat. gen. Part. Rept. 3: 358. 1831 Agama occipitalis Gray, in Griffith, Animal King- dom Cuvier, 9 Synops. Spec.: 56. 1844 Tropidolepis africanus Hallowell, Proc. Acad. Nat. Sci. Philadelphia 1844: 171. Type locality: Liberia. 1877 Agama colonorum var. congica Peters, M. Ber. K. preuß. Akad. Wiss. Berlin. 1877: 612. 1877 Agama picticauda Peters, M. Ber. K. preuß. Akad. Wiss. Berlin. 1877: 612. 1884 Agama agama savattieri Rochebrune, (synonym of A. a. africana fide Grandison 1968) Faune Sénégambie, Rept.: 89. 1896 Agama smithii Boulenger, (fide Largen & Spawls 2006), Proc. zool. Soc. London 1896: 213. 1940 Agama boensis Monard, Arq. Mus. Bocage, Lisboa 11: 155. Type locality: Madina Boé, Guinea. 1957 Agama agama mucosoensis Hellmich, Veröff. Zo- ol. Staatsamml. München 5: 44. Type locality: Don- do, Angola. Neotype. ZFMK 15222: Mokolo, Margui-Wandala Province, northern Cameroon, leg. W. BÖHME & W. HARTWIG, 14.11.1974 (figs 6a—c). Diagnosis. A large species of the genus. The nuptial male of Agama agama is characterized by a yellow head, a red throat and a tri-coloured (yellow, red and black) tail. The subordinate males, females, and adolescents possess an olive-green head and an olive-green to brown body with yellow dots, ocelli and stripes. 222 Philipp WAGNER et al.: Studies on African Agama V. On the origin of Lacerta agama Linnaeus, 1758 Description of the neotype of Lacerta agama Linnaeus, 1758 Adult male, measuring 290.11 mm in length (snout-vent length 128.93 mm, tail 161.18 mm, head length 33.49 mm, head width 23.61 mm, head height 14.07 mm). Nostril slightly above the canthus rostralis, pierced in the middle of a large vaulted nasal scale, directed obliquely posterodorsally. Between the nostrils is a longitudinal, finely keeled shield, followed by finely keeled to smooth, regularly arranged scales in the interorbital region and on the remaining upper side of head. Supraocular scales are smooth. Parietal shield more or less pentagonal, parietal organ slightly visible. Nine supralabial scales on the left, eight on the right side, nine infralabial scales on both sides. The scales originating from both sides of the parietal midline have their imbrications anteriorly directed; the keels in the upper temporal region have their mucrones at the anterior margins. Ear-opening large, about the same size as the eye; tympanum visible; anterior mar- gin being composed of spiny, mucronate scales. Four spiny tufts on the left side and five on the right, one before the ear opening, one above, one to two below and one to two behind. Nuchal crest low, consisting of 13 weakly differ- entiated lanceolate scales. Gular scales flat, juxtaposed and becoming smaller towards the gular fold. Dorsal scales keeled, becoming larger from neck and tail to the middle of the body; in 58 scale rows around midbody. Scales on tail arranged in weakly but nonetheless distinctly differ- entiated whorls that consist of three scale rings each, dor- sally strongly, ventrally weakly keeled to smooth. Ventral scales smooth, slightly to strongly imbricate at their pos- terior margin; in 80 rows between throat and vent. One row of 10 preanal pores. Scales on forelimb with strong- ly keeled scales, on the upper arm twice as large as the dorsals, becoming smaller towards the underside and the manus. 3" and 4 finger longest, 4h slightly longer, dig- ital length decreasing 2-5-1, subdigital lamellae keeled. Hindlimbs also covered by distinctly keeled scales, on the upper tights twice as large as the dorsals, becoming larg- er towards the lower tights and smaller towards the un- derside. Colouration in life. Head and nape reddish, limbs and body blue, base of tail white, followed by red and then black. Remarks. The rank of the two subspecies boensis and mu- cosoensis 1s questionable. GRANDISON (1968) mentioned boensis as a large-scaled Agama species and therefore some vouchers from Guinea could be identical with that taxon and in this consequence boensis should be elevat- ed to species rank (WAGNER & INEICH unpubl. data). 4ga- ma a. mucosoensis, with its striking blue and yellow colouration is as different to A. a. agama as A. planiceps, A. lionotus and many more well diagnosed species of the genus. Therefore it is very doubtful that mucosoensis is a subspecies of agama. Acknowledgements. We are grateful to Pierre Andre Crochet for his advises using the International Code on Zoological Nomenclature and to Sven Kullander for using the images of the NRM specimens. Cornelius de Haan and Arie van der Pas trans- lated the Dutch part of the text in Seba. REFERENCES ANDERSSON, L. G. (1900): Catalogue of Linnean type-specimens of Linnaeus’s Reptilia in the Royal Museum in Stockhol. Bi- hang till K. Svenska Vet.-Akad. Handlingar 26: 1-29. BAUMANN, H. (1936): Schópfung und Urzeit des Menschen im Mythos der afrikanischen Völker. Dietrich Reimer & Andrews Steiner Verlag, Berlin, 435 pp. BÖHME, W., WAGNER, P., MALONZA, P., LÖTTERS, S. & J. KÖH- LER (2005): A new species ofthe Agama agama group (Squa- mata: Agamidae) from western Kenya, East Africa, with com- ments on Agama lionotus Boulenger, 1896. Russian Journal Herpetology 12: 83-90. BOULENGER, G. A. & J. H. POWER (1921): A revision of the South African agamas allied to Agama hispida and A. atra. Trans- actions Royal Society South Africa 9: 229-287. Daupm, F. M. (1802): Histoire Naturelle, générale et parti- culieredes reptiles, ouvrage faisant suite, a l’histoire naturelle, generale et particuliere composée par Leclerc de Buffon, et redigée par C. S. Sonnini, vol. 3. F. Dufart, Paris. Dumerit, A. M. C. & G. BIBRON (1837): Erpétologie Générale; ou, Histoire Naturelle Complete des Reptiles. Vol. 4. Libr. En- cyclopédique Roret, Paris, 570 pp. GRANDISON, G. C. (1956): On a collection of lizards from West Africa. Mémoires de I’ institut francais d' Afrique noir 18: 224-245. GRANDISON, G. C. (1968): Nigerian lizards of the genus Agama (Sauria: Agamidae). Bulletin of the British Museum of Nat- ural History of Zoology 17: 67-90. JACOBSEN, N. H. G. (1992): The status of Agama aculeata ar- mata Peters, 1854 (Reptilia: Agamidae). Journal of the Her- petological Association of Africa 41: 30-34. LINNAEUS, C. (1749): Amoenitates academice seu dissertationes varie physic, medice botanice antehac seorsim edite nunc collectee et auctee cum tabulis eneis. Vol. 1, Holmiz, Lipsiz, 563 pp, tab. I-XVIL. LINNAEUS, C. (1754): Adolphi Friderici Regis Svecorum, Gotho- rum, Vandalorumquae &c.in quo Animalia Rariora Imprimis et Exotica: Quadrupedia, Aves, Amphibia, Pisces, Insecta, Ver- mes Describuntur et Determinatur, Latine et Suetice cum Iconibus. Typ. Regia. Holmiz, Stockholm, I-XXX, 1-96 pp. + 7 pp. LINNAEUS, C. (1758): Systema nature per regna tria natura, se- cundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Edito decima. Tomus I. Lauren- ti Salvi, Holmia, Stockholm, 823 pp. MCLACHLAN, G. R. (1981): Taxonomy of Agama hispida (Sauria: Agamidae) in southern Africa. Cimbebasia A 5: 219-227. MERTENS, R. (1938): Herpetologische Ergebnisse einer Reise nach Kamerun. Abhandlungen der senckenbergschen natur- forschenden Gesellschaft 442: 1-52, Bonner zoologische Beiträge 56 Moopy, S. & W. BOHME (1984): Merkmalsvariationen und ta- xonomische Stellung von 4gama doriae Boulenger, 1885 und Agama benueensis Monard, 1951 (Reptilia: Agamidae) aus dem Sudangúrtel Afrikas. Bonner Zoologische Beitráge 35: 107-128. QUENSEL, C. (1802): Catalogue of the collections of the Royal Museum. Stockholm. SCHNEIDER, J. G. (1792): Amphibiorum Physiologiae Specimen Alterum Historiam et Species Generis Stellionum seu Geckonum Sistens. Traiecti ad Viadrum (Frankfurt / Oder), C. L. F. Aitzi 2: 54 pp. SEBA, A. (1734): Locupletissimi Rerum Naturalium Thesauri Ac- curata Descriptio, et Iconibus Artificiosissimis Expressio, per Universam Physices Historiam. Opus, cui, in hoc Rerum Ge- nere, Nullum par Exstitit. Ex Toto Terrarum Orbe Collegit, Di- gessit, Descripsit, et Depingendum Curavit Albertus Seba, Et- zela Oostfrisius, Academize Caesarex Leopoldino Caroline [59] ho Ww Naturee Curiosorum Collega Xenocrates dictus; Societatis Re- giz Anglicane, et Instituti Bononiensis, sodalis. Tomus I, Jans- sonio-Waesbergios & J. Wetstenium & Gul. Smith, Amster- dam, 33, 178 pp, 111 pls. THYS VAN DEN AUDENAERDE, D. F. E. (1963): Les Agamidae du Congo: les especes et leur distribution géographique. Revue zoologique et botanique Africain 68: 203-250. WAGNER, P. (2007): Studies in African Agama |: On the taxo- nomic status of Agama lionotus usambarae Barbour & Loveridge, 1928 (Squamata, Agamidae). Herpetozoa 20 (1/2): 69-73. WAGNER, P. & P.-A. CROCHET (2009): The status of the nomina Trapelus savienyi Audouin, 1827 and Agama savignii DUMERIL & BIBRON, 1837 and the valid nomen of the Savi- gny’s Agama (Sauria: Agamidae). Zootaxa 2209: 57-64. WESTERMANN, D. (1954): Wórterbuch der Ewe-Sprache. Aka- demie Verlag, Berlin, 795 pp. Bonner zoologische Beiträge Band 56 Heft 4 Seiten 225-228 Bonn, November 2009 A record of Asian agama of the genus Calotes Cuvier, 1817 (Squamata: Agamidae) in Kenya Martin SANDERA 1:2 & Zuzana STAROSTOVA ! ! Department of Zoology, Faculty of Science, Charles University in Prague, Viniéna 7, CZ-128 44 Prague 2, Czech Republic; 2 Museum of Nature Bohemian Paradise, Prachov 37, CZ-506 01 Jiéin, Czech Republic; E-Mail: m.sandera@seznam.cz Abstract. Agamas of the genus Calotes Cuvier, 1817 occur in South and Southeast Asia. The garden lizard, Calotes ver- sicolor (Daudin, 1802), was introduced or dragged to several places and islands in Asia, Indian Ocean and USA (Flori- da). The record of this species in Africa was regardless unexpected. We demonstrate the first record of Calotes versicol- or in the African continent. Besides recognizing this species from the photograph, we were also able to identify it by us- ing DNA sequencing. Key words. Calotes versicolor, invasive species, African herpetofauna, distribution. INTRODUCTION Agamas of the genus Calotes Cuvier, 1817 occur in South and Southeast Asia (Moopy 1980, MANTHEY & SCHUS- TER 1996). The garden lizard, Calotes versicolor (Daudin, 1802), is widespread in South and Southeast Asia. Its area of distribution extends from Southeastern Iran and Afghanistan to Nepal, India, Sri Lanka, Myanmar, Indo- China, Southern China, Malaysia and Sumatra (MANTHEY & SCHUSTER 1996, RADDER 2006). This species probably colonized Sri Lanka thanks to human settlers and became more abundant in areas with extremely disturbed vegeta- tion (ERDELEN 1984). There are many regions or places in Asia and in the Indi- an Ocean where Calotes versicolor has been introduced or dragged. These are e.g.: Sumatra (Roon 1915), Réu- nion and Mauritius (VINSON & VINSON 1969), Celebes (ERDELEN 1978), the Maledives (HASEN DiDI 1993), Oman (SEUFER et al. 1999) and the Seychelles (MATYoT 2004). Finally, the garden lizard has also been introduced to Flori- da in the USA (ENGE € Krysko 2004). This species is adaptable and has a potential to be an invasive species with negative ecological impacts in the areas where it has been dragged (MATYoT 2004; RADDER 2006). RECORD, METHODS AND RESULTS Locality: Mombasa, railway station (GPS: S 04%03"27" EJ08923937 7, 20 m.a.s.]). The first find was made on March 11, 2007 about 11 p.m. The individual, a female, was sitting on a metal contain- er approx. 80 cm above the ground. The agama was caught, photographed (Fig. 1 and 2), and a sample for a DNA analysis was taken using buccal swabs. Another observation was done during a short excursion on March 21, 2007 (about 6 p.m.). As previously, a sin- gle individual was sitting on a heap of old metal railway ties (Fig. 3) approx. 100 m from the first finding. Theo- retically, this individual might have been the same as the first one. Its capture was unsuccessful and further identi- fication was therefore impossible. The animal hid in the heap of ties. However, ıt was probably a female. The railway station in Mombasa is situated close to the centre of the city. The territory of the station is fenced and guarded. The area of the station is several hectares large and its space 1s fragmented by used and unused buildings, wagons, containers, railroad ties, beams, planks, etc. The railway connects the railway station in Mombasa with the industrial port (Kilindini Harbour) distant by about one kilometer. The Mombasa railway station and port serve as a transship point for container transfer. We also observed a second agama species at the Mom- basa railway station. It was a member of the 4gama li- onotus complex. According to the referred occurrence (SPAWLS et al. 2004, BOHME et al. 2005) it could be A. /. 226 Martin SANDERA & Zuzana STAROSTOVA: A record of Asian agama of the genus Calotes Cuvier, 1817 Fig. 1. Calotes versicolor from the Mombasa railway station. (11. 1112007; photo M. Lazurková). lionotus. We saw males, females and offspring. We have also observed one gecko Hemidactylus cf. platvcephalus and one specimen of Hemidactylus sp. DNA was extracted from the buccal swab using Nucle- oSpin Tissue kit (Macherey-Nagel) following the manu- facturer’s protocol for buccal swabs. Fragments of mito- chondrial cytochrome b gene and nuclear oocyte matura- tion factor mos (c-mos) gene were amplified by poly- merase chain reaction using primers for cytochrome b from BURBRINK et al. (2000) and primers designed by SAINT et al. (1998) for c-mos. The sequencing was done with the same primers in the Abi Prism 3100 Avant Ge- netic Analyzer using the Abi Prism BigDye Terminator v 3.1 Ready Reaction Cycle Sequencing Kit (Applied Biosystems) in the sequencing laboratory of Charles Uni- versity in Prague, Czech Republic. The obtained partial 420 bp sequence of c-mos gene was 99% identical with the sequence of c-mos gene of Calotes versicolor from GenBank (Acc. No. AF137525). We were unsuccessful in amplification of the whole mitochondrial gene for cy- tochrome b; however, two unambiguously sequenced frag- ments of this gene (245 bp and 364 bp) were identical with the cytochrome b sequence of Calotes versicolor from GenBank (Acc. No. AB183287) in 82 % and 84 %, re- spectively. DISCUSSION Based on the comparison of DNA from our sample and sequences published in GenBank, we can, with high con- fidence, confirm finding individuals of Calotes versicol- or in Kenya. The lower correspondence in sequence com- position in the cytochrome b gene might be explained by a faster rate of evolution of this fragment and the existence of multiple cryptic species in the Calotes versicolor group, as described previously by ZuG et al. (2006), who how- ever used another fragment of mitochondial DNA. The finding of one or two individuals of Calotes versi- color, although within a short time period, suggests a re- cent introduction. With regard to the proximity of the Kilindini Harbour the introduction was probably accom- plished through shipping. The question is from which country the animals originate. Thanks to the lower corre- spondence in sequence composition in the cytochrome b gene mentioned above we can exclude Myanmar. The specimens from Mombasa presumably look like the spec- imens from islands Réunion and Mauritius (W. DENZER, Oxford, pers. comm. 2008). Further faunistic research in Mombasa, namely at the rail- way station and the port, would be useful for a surveil- Bonner zoologische Beiträge 56 227 Fig. 2. Calotes versicolor from the Mombasa railway station. (11. 111.2007; photo H. Sanderovä). Fig. 3. Calotes versicolor from the Mombasa railway station. (21. 11.2007; photo M. Sandera). lance of the potential population of the Calotes agamas in Kenya. Their impact on the local herpetofauna (name- ly on other lizards) and on invertebrate fauna should al- so be investigated. The garden lizard could represent an invasive species in Kenya with negative ecological im- pacts. DIONG et al. (1994) have pointed out to the fact that in Singapore it has somewhere displaced the native Bron- chocela cristatella. MAUREMOOTOO et al. (2003) include C. versicolor in the list of introduced vertebrates with a significant impact on native biodiversity in Mauritius, namely geckos, and consumes native invertebrates. VIN- SON (1968) assumed that C. versicolor may have been re- sponsible for the disappearance or decrease of phasmids in Réunion and Mauritius. Similar findings of introduced or dragged Calotes versi- color from other countries can also be expected in other places beside Africa. Acknowledgements. We thank H. Sanderová and M. Lazurková for providing us with photographs. The travel to Kenya was sup- ported from the Museum of Nature Bohemian Paradise, and DNA analysis together with the symposium De Agamis costs were covered by the Research Plan of the Ministry of Educa- tion, Youth and Sports of the Czech Republic 0021620828. REFERENCES BÖHME, W., WAGNER, P., MALONZA, P., LÖTTERS, S. & J. KÖH- LER (2005): A New Species of the 4gama agama Group (Squa- mata: Agamidae) from Western Kenya. Russian Journal of Herpetology 12 (2): 142-150. BURBRINK, F. T., Lawson, R. & J. B. SLOWINSKI (2000): Mito- chondial DNA phylogeography of the polytypic North Amer- ican rat snake (Elaphe obsoleta): A critique of subspecies con- cept. Evolution 54: 2107-2118. DIONG, C. H., CHou, L. M. & K. K. P. Lim (1994): Calotes ver- sicolor, the changeable lizard. Nature Malaysiana 19: 46-54. ENGE, K. M. & K. L. Krysko (2004): A new exotic species in Florida, the bloodsucker lizard, Calotes versicolor (Daudin, 1802) (Sauria: Agamidae). Biological Science 67 (3): 226-230. 228 Martin SANDERA & Zuzana STAROSTOVÁ: A record of Asian agama of the genus Calotes Cuvier, 1817 ERDELEN, W. (1978): Distributional patterns of the genus Calotes (Sauria, Agamidae) in Sri Lanka. Loris 14 (6): 350-356. ERDELEN, W. (1984): The genus Calotes (Sauria, Agamidae) in Sri Lanka: distribution patterns. Journal of Biogeography 11: 515-525. Hasen Dıpı, N. T. (1993): Observation on the nesting of a gar- den lizard (Calotes versicolor) in the Maldives. Hamadryad 18: 42. MANTHEY, U. & N. SCHUSTER (1996): Agamid Lizards (Engl. ed.). T.F.H., Neptune City, 189 pp. MATYOT, P. (2004): The establishment of the crested tree lizard, Calotes versicolor (Daudin, 1802) (Squamata: Agamidae), in Seychelles. Phelsuma 12: 35-47. MAUREMOOTOO, J. R., LECKRAZ, N. R., PUTTOO, M., BELLOUARD, E., GANESHAN, S. & S. P. BENI MADHU (2003): Invasive alien species of Mauritius. Pp. 12-37 in: MACDONALD, I. A.W., REASER, J. K., BRIGHT, C., NEVILLE, L. E., HOWARD, G. W., MURPHY, S. J. & G. PRESTON (eds.): Invasive alien species in southern Africa: national reports & directory of resources. Global Invasive Species Programme (GISP), Cape Town, South Africa. Moopy, S. M. (1980): Phylogenetic relationships and historical biogeographical relationships of the genera in the family Agamidae (Reptilia: Lacertilia). PhD dissertation. Universi- ty of Michigan. 373 pp. RADDER, R. S. (2006): An overview of geographic variation in the life history traits of the tropical agamid lizard, Calotes ver- sicolor. Current Science 91 (10): 1354-1363. Room, N. de (1915): The reptiles of the Indo-Australian Archi- pelago. E. J. Brill, Leiden. 384 pp. SAINT, K.M., AUSTIN, C.C., DONNELLAN, S.C. & M. N. HUTCHIN- SON (1998): C-mos, a nuclear marker useful for squamate phy- logenetic analysis. Molecular Phylogenetics and Evolution 10: 259-263. SEUFER, H., KOWALSKI, T. & H. J. ZILGER (1999): Herpetologi- sche Impressionen einer Reise in den Oman. Herpetofauna Weinstadt 21: 24-34. SPAWLS, S., HOWELL, K., DREWES, R. C. & J. ASHE (2004): Field Guide to Reptiles of East Africa (reprint). A & C Black, Lon- don. 543 pp. VINSON, J. (1968): Les Blattoptéroides, Orthoptéroides et Der- matoptéroides des Mascareignes. The Mauritius Institution Bulletin 6 (3): 103-118. VINSON, J. & J. M. VINSON (1969): Saurian Fauna of Mascarene Islands. The Mauritius Institution Bulletin 6: 203-320. ZuG, G. R., BRown, H. H. K., SCHULTE, J. A. & J. V. VINDUM (2006): Systematics of the Garden Lizards, Calotes versico- lor Group (Reptilia, Squamata, Agamidae), in Myanmar: Cen- tral Dry Zone Populations. Proceedings of the California Aca- demy of Sciences ser. 4, 57 (2): 35-68. Bonner zoologische Beiträge | Band 56 Heft 4 Seiten 229-238 Bonn, November 2009 Taxonomic and biological study on Calotes ceylonensis Müller, 1887 (Reptilia: Agamidae) of Sri Lanka D. M. S. Suranjan KARUNARATHNA!, A. A. Thasun AMARASINGHE? & Edi STÖCKLB 1 TUCN, Sri Lanka Country Office, 53, Horton Place, Colombo 7, Sri Lanka; E-mail: dmsameera@gmail.com 2 Taprobanica Nature Conservation Society, 146, Kendalanda, Homagama, Sri Lanka; E-mail: aathasun@gmail.com 3 Naturhistorisches Museum Basel, Switzerland; E-mail: eduard.stoeckli@bs.ch Abstract. Calotes ceylonensis Müller, 1887 is an endemic, rare and vulnerable, arboreal agamid lizard species of Sri Lanka, which is found only at the low country dry and intermediate zones below 500 m a.s.l. This work is mainly based on examination of type specimens as well as published literature and our observations of ecological and conditions and threats on Calotes ceylonensis. The analysis of habitat data has shown that this species is widely spread within the well shading riverine-forested areas and poorly in the home gardens. The results of this survey indicate that C. cevlonensis lays about 4-12 eggs from August to October. Hatchlings come out from November to December. These lizards’ natural predators are arboreal Colubrid snakes, Hornbills and Civet cats. The current habitat destruction poses a huge threat to this species. Keywords. Agamidae, Calotes ceylonensis, Taxonomy, Natural history, Ecology, Biology, Behaviour, Sri Lanka. INTRODUCTION There are eighteen species of agamid lizards in Sri Lan- ka and fifteen (83.33%) of them are endemic to the island (DE SILVA 2006; MANAMENDRA-ARACHCHI et al. 2006). These eighteen species are classified under sub family Draconinae (MACEY et al., 2000). According to BAHIR & SURASINGHE (2005), MANAMENDRA-ARACHCHI et al. (2006) and lucnsL & MENR (2007) four species are Crit- ically Endangered; five species Endangered; three species Vulnerable and six Least Concern. The genus has a pit or fold in front of shoulder or none; body is compressed; supra ocular scales generally enlarged; dorsal scales are usually regular; dorsal crest and gular sac are more or less developed; tail is elongated, slender and sub cylindrical (DERANIYAGALA 1953). The genus Calotes extends throughout southern Asia, most of the East Indian Archipelago (TAYLOR 1953).There are seven species in the genus Calotes. Five of them (C. cey- lonensis Múller, 1887; C. liocephalus Gúnther, 1872; C. liolepis Boulenger, 1885; C. nigrilabris Peters, 1860; C. desilvai Bahir & Maduwage, 2005) are endemic to Sri Lanka. The remaining two Calotes species (C. calotes [Linnaeus, 1758]; C. versicolor [Daudin, 1802]) are probably widespread throughout South East Asia. Ac- cording to the published literature Calotes ceylonensis is a largely arboreal species found only from the low coun- try dry and intermediate zones below 500 m a.s.! (Das & DE SıLva 2005; MANAMENDRA-ARACHCHI & LIYANAGE 1994). Its conservation status is Rare and Vulnerable (MANAMENDRA-ARACHCHI & LIYANAGE 1994; JUCNSL & MENR 2007). The information available on this species 1s marginal, and therefore further studies on their behaviour and ecology, which may be very important for the con- servation of the species, are needed. Hence it is essential to gather information on the C. ceylonensis in different ar- eas of the country as a first step towards conservation of this lizard species. MATERIALS AND METHODS The material examined is deposited at the NMB, Naturhis- torisches Museum Basel, Switzerland and WHT, Wildlife Heritage Trust of Sri Lanka, Colombo, Sri Lanka. Diag- noses and descriptions are based on external morpholo- gy. This work is mainly based on examination of type 230 D. M. S. Suranjan KARUNARATHNA: Taxonomic and biological study on Calotes ceylonensis of Sri Lanka Fig. 1. Distribution map of C. ceylonensis (Square, Holotype; Triangle, Paratype; Circle, Other). specimen and WHT specimens as well as published liter- ature and our observations on their ecological and behav- ioural conditions and threats. The locality records for each species include WHT specimen data, published locality records as well as our observations during past decade. We have examined 45 localities (March 2002 to April 2006) Table 1. Measurements of C. cevlonensis hatchlings. throughout 7 districts (Anuradapura: 5 / Monaragala: 13 / Ampara: 8 / Hambantota: 7 / Badulla: 4 / Matale: 3 / Polonnaruwa: 5) (fig. 1). A total of 68 individuals (males: 37; females: 18; juveniles: 13) were recorded during the study period. The collected species were examined, meas- ured and noted down carefully before released back to the same habitats (tab. 1 & 2). The diagnostic keys given by DERANIYAGALA (1953), MANAMENDRA-ARACHCI (1990), MANAMENDRA-ARACHCI (1998), MANAMENDRA-ARACHCHI & LIYANAGE (1994), SMITH (1935) and TAYLOR (1953) were used for species identification. Principal components analysis of preferred habitats was done using Pcord4 and each number of males, females and juveniles against the preferred habitat type were used in this analysis. All the photographs and line drawings displayed with the photographer’s and artist's initials; ES (Edi Stóckli), TA (Thasun Amarasinghe) and SK (Suranjan Karunarathna). All measurements were taken to the nearest 0.1 mm with dial calipers. Scale counts. SUP, Supralabials were counted from the first scale anterior to that at angle of gape, not including the median scale (when present); INF, infralabials were counted from first scale posterior to mental, to angle of gape; DS, dorsal spines were counted from first spine to last of mid-dorsal row; CR, canthus rostralis (counted from rostral scale along scale row passing over nostril to posterior end of supraciliary ridge); MDS, mid dorsal scales (counted from scale behind rostral to posterior mar- gin of the thigh); MBS, mid body scales were counted from center of mid-dorsal row forwards and downwards across ventrals (this count 1s, however, made unreliable by the unequal size and uneven arrangement of the later- al scales); MVS, mid ventral scales were counted from first scale posterior to mental, to last scale anterior to vent: SAT, Spines around tympanum were counted from first spine to last above tympanum. External measurements. SVL, snout-vent length (distance between tip of snout to anterior margin of vent); HL, head length (distance between posterior edge of mandible and tip of snout); HW, head width (maximum width of head); DHL, dorsal head length (distance between posterior edge Measurement given in mm 1 2 3 4 5 6 SVL 34.1 32.8 35.0 34.5 34.6 35.1 TL 65.5 66.8 67.2 64.8 66.1 7 8 9 10 11 Average 33,9 34.8 33.4 84.20 32:9 34.11 66.2 64.5 63.9 66.5 65.8 65.51 Bonner zoologische Beiträge 56 231 Table 2. Measurement of C. cevlonensis Eggs. No. Egg length Egg width 1 16.5 8.6 2 16.8 8.4 3 14.3 ao 4 15.6 8.5 5 13.2 8.7 6 Piel 8.8 7 16.9 7.8 8 171 7.6 9 14.6 8.5 10 18,7 7.8 11 14.8 1.9 12 14.4 8.3 13 135 8.8 14 16.1 8.9 15 16.7 79 16 [6:2 8.5 17 13.8 8.1 18 14.5 8.6 19 15.3 8.4 20 15.9 8.1 21 13.5 8.9 22 17 2.9 23 16.8 8.3 24 16.4 8.7 25 15.8 8.1 26 15.3 8.8 27 13.6 1.9 28 13:5 8.6 29 15.3 8.9 30 16:7 7.8 31 16.2 8.3 32 16.6 8.8 Mean 15.55 8.34 of cephalic bone and tip of snout); NFE, nostril — front eye length (distance between most anterior point of orbit and middle of nostril); UAL, upper-arm length (distance between axilla and angle of elbow); LAL, lower-arm length (distance from elbow to wrist with both upper arm and palm flexed); FL, finger length (distance between tip of claw and the nearest fork); FEL, femur length (distance between groin and knee); TBL, tibia length (distance be- tween knee and heel, with both tibia and tarsus flexed); TL, toe length (distance between tip of claw and nearest fork); AG, axilla-groin length (distance between axilla and groin); SA, snout-axilla length (distance between tip of snout and axilla); TAL, tail length (measured from an- terior margin of vent to tail tip); PAL, palm length (tak- en from posterior most margin of palm and tip of longest finger); FOL, foot length (distance between heel and tip of longest toe, with both foot and tibia flexed); TBW, width of tail base (most distance of the tail base); LOW, inter orbital width (least distance between the upper mar- gins of orbits); ED, eye diameter (horizontal diameter of orbit); SFE, snout-front eye length (distance between an- terior most point of orbit and tip of snout); SBE, snout-back eye length (distance between posterior most point of orbit and tip of snout); SFT, snout-front tympa- num length (distance between anterior most point of tym- panum and tip of snout); TD, Tympanum diameter (least distance between the inner margins of tympanum). RESULTS Calotes ceylonensis Müller, 1887 English Name: Painted-lip Lizard Sinhala Name: Thola — wisithuru Katussa Calotes mystaceus, var. ceylonensis Múller, 1887 (MULLER 1887: 8, 292-293, pl. 3.). Calotes ceylonensis (BOULENGER 1890; DERANIYAGALA 1931; SMITH 1935; TAYLOR 1953). Calotes kelaartii Nevil, 1887 (HALY 1887: 2, 133. [Species described but not named]; NeviL 1887: 2, 134, pl. 5 [Species named but not described]). Calotes saleoides Werner, 1896 (WERNER 1896: 46, 7). Holotype (fig.2). Male (81.9 mm SVL); Cat. no. NMB 3340; Loc. Kumbukan-aar: South East Ceylon (+ Kum- bukkan Oya?: Sri Lanka [“aaru” in Tami [= “Oya” in Sin- hala]); Coll. P. Sarasın & F. Sarasin; Date. 1886. Paratype (in NMB catalogue labeled as Syntype). Male Fig. 2. C. cevlonensis, NMB 3340; Holotype, male; lateral aspect of head (ES). 232 D. M. S. Suranjan KARUNARATHNA: Taxonomic and biological study on Calotes ceylonensis of Sri Lanka Fig.3. C. cevlonensis, NMB 3341; Paratype, male; lateral aspect of head (ES). (67.6 mm SVL); Cat. no. NMB 3341; Loc. North East province: Ceylon (+ Northern Province ? / Eastern Province ?: Sri Lanka); the same coll. & same date (fig. 3). Other materials examined. WHT 7397, male, 59.1 mm, Wasgamuwa; WHT 7514, male, 74.8 mm, Giritale; WHT 1427A, male, 79.6 mm, Wasgamuwa; WHT 1427B, male, 75.0 mm, Wasgamuwa; WHT 1427C, female, 76.7 mm, Wasgamuwa; WHT 1427D, female, 61.6 mm, Wasga- muwa; WHT 1428, male, 80.3 mm, Tatugala-Bulupitiya; WHT 0515, male, 72.9 mm, Pallegama; WHT 1625A, male, 77.2 mm, Wasgamuwa; WHT 1625B, male, 71.9 mm, Wasgamuwa; WHT 0511, male, 82.5 mm, Kon- ketiya-Buttala; WHT 0521, female, 72.1 mm, Konketiya- Buttala; WHT 0522, male, 73.2 mm, Konketiya-Buttala; WHT 0519, male, 73.0 mm, Yala; WHT 0520, male, 73.4, Yala. WHT 1624A, male, 65.1 mm, Wasgamuwa o ER 7 LV AAA Fig. 4. C. ceylonensis, WHT 0511; male; lateral scalation of head (TA). Diagnosis. Two well-separated spines above the tympa- num, the anterior and upper lateral pointing backwards and upwards, the other straight backwards, an oblique fold in front of the shoulder runs across throat covered with small granular scales. Dorsal crest formed of 7 to 13 small spines. In a fully-grown male the head is larger and the base of the tail more swollen. Description. (Based on MÚLLER 1887; Holotype (NMB 3340); Paratype (NMB 3341) and WHT collection). Length of head one and a half times its breadth; snout a little longer than the orbit; forehead concave; cheeks swollen in the adult male; upper head scales unequal, smooth; canthus rostralis and supraciliary edge sharp; two well separated spines above the tympanum, the upper and anterior one mid way between it and the nuchal crest; di- ameter of the tympanum half that of the orbit; Supralabi- als, 10-14; Infralabials, IX-XUI (fig. 4). Body com- pressed; dorsal scales rather small, smooth or faintly keeled, the anterior and upper ones pointing backwards and upwards (fig. 5); the others straight backwards, larg- er than the ventrals, which are strongly keeled and mu- cronate; 53 to 76 scales round the body. No trace of a gu- lar sac; gular scales strongly keeled, as large as the ven- trals; an oblique fold in front of the shoulder runs across throat covered with small granular scales. Dorsal crest Bonner zoologische Beiträge 56 Fig. 6. C. ceylonensis Mature Male (lateral aspect) (SK). formed of 7 to 13 small spines; no dorsal crest. Limbs moderate; third and fourth fingers are sub equal; Relative length of fingers; 1<5<2<4<=3. Fourth toe distinctly longer than third. Relative length of toes; 1<2<5<3<4. The hind limb reaches to the tympanum or beyond. Tail long and slender; in the adult male it is markedly swollen at the base, with large, thick, keeled scales, those of the up- per median row forming a slight serrated ridge. Remarks. (Based on MÜLLER [1887] and observations on living specimens). The body color is changeable from grayish brown to light blue or dark brown, back of the head and anterior part of back pale brownish, divided in to more or less distinct spots by dark transverse bars; hin- der part of back and tail with dark cross bars; upper lip with a pale strongly defined stripe, which extends to be- yond the ear; sides of neck and chest black or white with black reticulations; gular region grayish or blackish; bel- ly pale brown with more or less distinct angular bands (figs. 6 & 7). Scale counts and External measurements (in mm) of Holotype. SUP, 10; INF, 10; DS, 9; CR, 14; MBS, 59; MDS. 78: MVS: 101: SAT, 2: SVL, 81.9; HL, 31.2; HW, 18.6; DHL, 20.6; NFE, 6.7; UAL, 12.3; LAL, 14.7; FLI, 4.3; FLII, 7.0; FLUI, 10.1; FLIV, 9.6; FLV, 6.6; FEL, 20.3; P2247. 11542: TLL 7.5: TLL, 11:6; TLIV,. 14.1; TLV, 9.9; AG, 33.9; SA, 37.7; TAL, 185; PAL, 12.2; FOL, Fig. 7. C. cevlonensis Mature Male (dorsal aspect) (TA). 234 D. M. S. Suranjan KARUNARATHNA: Taxonomic and biological study on Calotes ceylonensis of Sri Lanka 16 = juveniles 14 12 10 8 No. of Individuals ON >» 0 — RF DF WE SA HG Macro Habitat Fig. 8. PC analysis of preferred macro habitats (RF, Riverine Forests; DF, Dry-mixed Forest; WF, Wet-mixed Forest; SA, Sav- annah; HG, Home Gardens). 18 - juveniles No. of Individuals Ls (00) O: N GR 77 TM TB RO WA Micro Habitat Fig. 9. PC analysis of preferred micro habitats (GR, Ground; TT, Tree Top - >10m height; TM, Middle of Tree 5 to 10m height; TB, Base of Tree 1 to 5m height; RO, On Rocks; WA, On Walls). Bonner zoologische Beiträge 56 18:8; SET, 23.5; TD, 2.8. Scale counts and External measurements (in mm) of Paratype. SUP, 11; INF, 11; DS, 8; CR, 14; MBS, 53; MDS, 73; MVS, 84; SAT, 2; SVL, 67.6; HL, 19.9; HW, 11.6; DHL, 16.2; NFE, 4.7; UAL, 9.7; LAL, 11.2; FLI, 4.0; FLII, 6.4; FLIII, 7.5; FLIV, 7.3; FLV, 5.1; FEL, 16.3; IBAI Ost 3:55 TLI, 6.0; TLIN, 10.2; TLIV, 10.6; TLV, 8.6; AG, 27.5; SA, 25.1; TAL, 156; PAL, 10.6; FOL, 17.9; TBW, 6.3; IOW, 9.0; ED, 6.0; SFE, 8.1; SBE, 13.5; Sale ieo: TD: 2:6. Distribution and Habitat. Calotes ceylonensis 18 record- ed only from the semi evergreen monsoon forests, plan- tations and home gardens of the dry and intermediate zones up to 500 m a.s.l. within the well shading riverine- forested areas and poorly in the home gardens (Das & DE SILVA 2005; ERDELEN 1984; MANAMENDRA-ARACHCHI & LIYANAGE 1994). According to our observations a large number of individuals of this species mostly favour shady riverine forests over the other macro habitats (fig. 8) and we also could locate a great number of individuals from top of the trees rather than the other microhabitats (fig. 9). According to our survey the female sex ratio 1s low com- paratively to the males except in Ampara district. The highest number of males was recorded from Monaragala district and least numbers from Matale and Badulla dis- 16 14 tricts and also the number of females, juveniles, hatchlings as well as eggs were low in these two districts (fig. 10 and tab. 3). Reproduction. There have been only few attempts to use hemipenis morphology as a taxonomic tool for agamid lizards although there is a considerable amount of struc- tural diversity (MCCANN 1949, BÖHME 1988). The hemipenis of C. ceylonensis (fig. 11) seems less differen- tiated as compared to C. nigrilabris and €. liocephalus (see also the descriptions and fig. 18 in BÖHME 1988). The pedicel is considerably longer than the head; below the head, it is broadened out in to two shallowly concaved shoulders; there are no spines. The head is quadrangle in shape. It is shallowly divided longitudinally in to four lobes, two being very slightly larger than the others. The surface of the head is reticulatedly pitted, the pits being larger on the outside and diminishing in size towards the divisions between the lobes (fig. 11). The female digs a nest hole in the ground and deposits about 4-12 eggs in August to October on shady places near from tree bases. While laying eggs the females change their body color to black. The eggs are 13.5 mm-17.1 mm long and 7.8 mm-8.9 mm wide. Hatchlings come out from November to December (DERANIYAGALA 1953). Behaviour. This species 1s largely arboreal and active dur- ing the day, widely spread within the well shading river- ine-forested areas and poorly in the home gardens. They 2 u 12 2. 10 = E 8 = 6 So 4 © z 2 0 o xD & > so NS > „Ss © ES SS S Na S = Se SS dl e x N > 9 2 sO S S RS 2s District m Males @ Females @ Juveniles Hatchlings OEggs Fig. 10. No. of individuals recorded in each district. 236 Table 3. Observed locations and individual details of C. ceylonensis. District Monaragala Ampara Hambantota Matale Badulla Polonnaruwa Anuradapura D. M. S. Suranjan KARUNARATHNA: Taxonomic and biological study on Calotes ceylonensis of Sri Lanka Location Pitakumbura Wellavaya Puwakpale Bulupitiya Monaragala Hangala Yakunhela Buttala Nilgala Ulhela Rathugala Pihibiyamukalana Hamapola Hulannuge Buddangala Kumana Galoya Yala Makara Aravila Manakanda Palatupana Kataragama Weeravila Hambantota Akasachihthiya Tissamaharama Yala Maduruoya Giritale Polonnaruwa Wasgomuwa Minneriya Koslanda Udadiyaluma Akkaraseya Nikapotha Hukkumbura Wasgomuwa Narangamuwa Galkulama Padaviya Ritigala Anuradapura Kekirawa OO Ru q h He WH OO RB Re He o > A ] o o => m . . .n - =- +. + + p » pa > + 2.0 [m 8 oo oO No. of Males No. of Females OO 2.00 -=- O KY CO KH CS oo-o-.-..-600 COFFS oO OO nO »-OP»OOoOñrO Sr, oooO o OOO 09000 090 oo ro oo 0 m oO SSH OO 0900 09 O rm ro 0 co KK © S 2 | NN DD SD SS NM Mw DD =) DO Oo 00 0 © © ©°o © oo So ©S°S- OO N N OO CO COO NN NN OC WN No. of Juveniles No. of Eggs SS PS, OA SO 1 OS AOS DO O eat SOO Ore Sea O SS) O ASADO MOS SOMO NOOO 208 On Ol Om No. of Hatchlings Bonner zoologische Beiträge 56 237 Fig. 11. specially feed on butterflies, honeybees and other insects. These lizards’ natural predators are arboreal colubrid snakes, hornbills and civet cats. Being in danger this lizard suddenly climbs into a tree to an average height of 15 m and then glides to another tree, which is | m away till the danger disappears. If there is no other tree to glide, they just jump down to the ground and stay a few minutes with- out moving, and then suddenly run on another tree. Hatch- lings are mostly waiting for their preys on Lantana cam- era trees (an invasive plant to Sri Lanka) while camou- flaging their snout color into pink (the flowers of L. cam- era are also pink). Their prey are insects which feed on L. camera. Conservation status. Rare (MANAMENDRA-ARACHCHI & LIYANAGE 1994), Least Concern (BAHIR & SURASINGHE 2005) and Vulnerable (IUCNSL & MENR 2007). DISCUSSION The holotype of Calotes ceylonensis (NMB 3340) was col- lected from “Kumbukan-aar: South East Ceylon” by P. & F. Sarasin. Today there is known no area or river basin as “Kumbukan-aar” in South East Sri Lanka. However there is a river basin called “Kumbukkan Oya”. In Sinhala “Oya” means a small river/large stream. In Tamil “aaru” is the same meaning for “Oya or River”. There are sev- eral rivers in East and North parts of Sri Lanka named as “aaru” in Tamil. However there is no evidence that name used for “Kumbukkan Oya”. Somehow the Tamil people live in that area used to call this river as “Kumbukkan 1cm C. ceylonensis, WHT 1428; male, 80.3 mm SVL; lateral aspect of left hemipenis (TA). aaru”. Therefore we believe that the person who gave this information to P. & F. Sarasin might have been be a Tamil person. In addition now there is also a considerable pop- ulation of C. cevlonensis inhibiting this area. Therefore we assume the name for holotype locality as “Kumbukkan Oya”, which 1s currently used in Sri Lanka. In addition the paratype of (NMB 3341), was collected from “North East province: Ceylon” by the same collec- tors. Actually nowadays there is no district as “North East province” in Sri Lanka and this province considered as two provinces called Northern Province and Eastern Province. Therefore it is difficult to place the exact location and al- so C. ceylonensis is distributed throughout these two provinces of the country. The ecological and behavioural status of C. ceylonensis has not been investigated up to now. C. ceylonensis is an endemic, rare and vulnerable species. Therefore many published literature on their ecological status will have to be done for the conservation of this species. And also cap- tive breeding methods may be needed for ex-situ conser- vation of these species. A major portion of the primary forests of Sri Lanka has undergone considerable destruc- tion during the past 150 years. As a result: most of the agamid lizards that inhabit in these primary forests now live in degraded or altered habitats. According to DE SIL- VA (1996) C. ceylonensis is distributed in primary forest where a considerable amount of destruction and alteration of the forest by human activities have taken and are still taking place. As a consequence only 5% to 25% of pri- mary or secondary forest / scrub jungle still exist with the 238 D. M. S. Suranjan KARUNARATHNA: Taxonomic and biological study on Calotes ceylonensis of Sri Lanka greater extent being anthropogenic vegetations. C. ceylo- nensis 18 sympatric with Calotes liolepis ın Ritigala, Nil- gala, and the eastern slope of the Knuckles region; with C. versicolor, Otocryptis nigristigma, Lyriocephalus scu- tatus and Sitana ponticeriana ın many habitats through- out the intermediate and dry zones. Acknowledgements. We would like to express our sincere thank to Kelum Manamendra-Arachchi (WHT) and M. M. Bahir who provided valuable comments and helped improve the document. We also thank Rohan Pethiyagoda (WHT), Franz Tiedemann (NMW), Madhava Botegue (TNCS) and Sudath Nanayakkara (WHT), who helped in diverse ways to enrich this work. REFERENCES BAHIR, M. M. & T. D. SURASINGHE (2005): A conservation as- sessment of the agamid lizards of Sri Lanka. In: Yeo, D. C. J., NG, P. K. L. & R. PETHIYAGODA (eds. ): Contributions to bio- diversity exploration and research in Sri Lanka. The Raffles bulletin of zoology, supplement 12: 381-392. BÖHME, W. (1988): Zur Genitalmorphologie der Sauria: Funk- tionelle und Stammesgeschichtliche Aspekte. Bonner zoolo- gische Monographien 27: 1-176. BOULENGER, G. A. (1890): The Fauna of British India, includ- ing Ceylon and Burma: Reptilia and Batrachia. Taylor and Francis, London. xvii + 541. Das, I. & A. DE SILVA (2005): Snakes and other Reptiles of Sri Lanka. New Holland Publishers, UK 144. DERANIYAGALA, P. E. P. (1931): Some Ceylon lizards. Ceylon Journal of Science, section B, 16: 139-180. DERANIYAGALA, P. E. P. (1953): A Colored Atlas of some verte- brates from Ceylon, Tetrapod Reptilia, National Museums of Sri Lanka, Colombo 2: 101. DE SiLva, A. (1996): The Herpetofauna of Sri Lanka: a Brief Re- view. Published by the author. 99. De SILVA, A. (2006): Current status of the Reptiles of Sri Lan- Ka. Pp. 134-163 in: BAMBARADENIYA, C.N.B. (ed.): Fauna of Sri Lanka: Status of Taxonomy, Research and Conservation. IUCN Sri Lanka. ERDELEN, W. (1984): The genus Calotes (Sauria: Agamidae) in Sri Lanka: distribution patterns. Journal of Biogeography 11: 515-525. HALy, A. (1887): Notes on species of Calotes. Taprobanian 2: 133, TUCNSL & MENR (2007): The 2007 Red List of threatened Fauna and Flora of Sri Lanka. Colombo, IUCN Sri Lanka 148 pp. Macey, J. R., SCHULTE II, J. A., LARSON, A., ANANJEVA, N. B., WANG, Y., PETHIYAGODA, R., RASTEGER-POUYANI, N. & T. J. PAPENFUSS (2000): Evaluating Trans-Tethys migration: an ex- ample using Acrodont lizard phylogenetics. Systematic Biol- ogy 49 (2): 233-256. MANAMENDRA-ARACHCHI, K. (1990): A guide to the agamids in Sri Lanka. Young Zoologists’ Association of Sri Lanka. Oc- casional Paper 5: 1-6. MANAMENDRA-ARACHCHI, K. (1998): Let's hear it for the Gar- den Lizards. Sri Lanka Nature 2 (1): 48-62. MANAMENDRA-ARACHCHI, K., DE SILVA, A. & T. AMARASINGHE (2006): Description of a second species of Cophotis (Reptil- ia:Agamidae) from the highlands of Sri Lanka. Lyriocephalus 6 Supplement 1: 1-8. MANAMENDRA-ARACHCHI, K. & S. LIYANAGE (1994): Conserva- tion and distributions of the agamid lizards of Sri Lanka with illustrations of the extant species. Journal of South Asian Na- tural History 1: 77-96. MÜLLER, F. (1887): Fünfter Nachtrag zum Katalog der herpeto- logischen Sammlung des Basler Museums. Verhandlungen der Naturforschenden Gesellschaft in Basel 8: 292-293: pl 3. MCCAnn, C. (1949): The hemipenis in reptiles. Journal of Bom- bay natural History society 46 (2): 347-373. NEVIL, H. (1887): Notes on Calotes in Ceylon. Taprobanian 2: 134, pl. 5. SMITH, M. A. (1935): The fauna of British India including Cey- lon and Burma, Reptilia and Amphibia, II — Sauria. Taylor and Francis, London. xiv + 440 pp, pl. 1. TAYLOR, E. H. (1953): A review of the lizards of Ceylon. Uni- versity of Kansas Science Bulletin 35: 1525-1585. WERNER, F. (1896): Zweiter Beitrag zur Herpetologie der indo- orinentalischen Region. Verhandlungen der zoologischen und botanischen Gesellschaft in Wien 46: 7. Bonn, November 2009 am zoologische Beiträge Band 56 Heft 4 Seiten 239-253 Studies on African Agama VI. Taxonomic status of the West African Agama (Sauria: Agamidae) with prominent tail crests: Agama boulengeri Lataste 1886, Agama insularis Chabanaud, 1918 and Agama cristata Mocquard, 1905 Philipp WAGNER", Ivan INEICH?, Adam D. LEACHE?, Thomas M. WILMs?, Sébastien TRAPES, Wolfgang BÖHME! & Andreas SCHMITZ® ! Zoologisches Forschungsmuseum A. Koenig, Adenauerallee 160, D-53113 Bonn, Germany philipp.wagner.zfmk@uni-bonn.de; w.boehme@uni-bonn.de 2 Museum national d’Histoire naturelle, Departement de Systematique et Evolution (Section Reptiles) UMR 7205 CNRS “Origine, Structure et Evolution de la Biodiversite”, CP n° 30-25 rue Cuvier, F-75231 Paris Cedex 05, France. 3 Genome Center & Department of Evolution and Ecology, University of California, Davis, CA 95616 USA. 4 Zoologischer Garten Frankfurt, Bernhard-Grzimek-Allee 1, D-60316 Frankfurt, Germany > Université de Montpellier II, UMR 5119 Ecolag, IRD-CNRS-UM2-IFREMER, cc093, place Eugene Bataillon, F-34095 Montpellier, France. 6 Museum d "histoire naturelle, | route de Malagnou, CH-1208 Geneva, Switzerland “corresponding author Abstract. This publication reviews the taxonomy of three West African 4gama species, A. boulengeri, A. cristata, and A. insularis, each characterized by a prominent tail crest in adult males. Following the results from morphological and genetic analyses, Agama insularis is recognized as a synonym of the revalidated Agama cristata, whereas this species is clearly distinct from Agama boulengeri. We present a detailed distribution map for these 4gama species, as well as for A. weidholzi. Following the results of recent publications, Agama atra knobeli ıs herein regarded as a full species. Key words. Africa, Guinea, Los Islands, Ile de Roumé; Sauria, Agamidae, Agama cristata, A. insularis, A. boulengeri, A. knobeli, Agama weidholzi. INTRODUCTION Despite an increasing amount of work on the taxonomy and phylogeny of African lizards in the genus Agama (BÖHME et al. 2005, PADIAL 2005, WAGNER 2007, WAGN- ER et al. 2008a, WAGNER et al. 2008b, WAGNER et al. 2009a, WAGNER et al. 2009b) several species are only poorly known, and the validity of some taxa remains ques- tionable (e.g. A. bocourti Rochebrune, 1884; A. bottegi Boulenger, 1897; A. cornii Scortecci, 1928; A. insularis Chabanaud, 1918 and 4. agama boensis Monard, 1940 [re- garded as a synonym of A. weidholzi by GRANDISON 1969]). Many species are only diagnosable based on adult male coloration (LOVERIDGE 1933, THYS VAN DEN AUDE- NAERDE 1963, MCLACHLAN 1981, BÖHME et al. 2005, WAGNER 2007, WAGNER et al. 2008a, WAGNER et al. 2008b), and the lack of appropriate comparative material coupled with inadequate descriptions of the adult coloration in life prevents formal taxonomic evaluations. Three species in West Africa are characterized by possess- ing a prominent tail crest in adult males: 4gama boulen- geri, A. insularis and A. cristata. The taxonomic status of these taxa 1s the focus of this study. Agama insularis, was described by CHABANAUD (1918) on the basis of a juvenile and a subadult female collected by Dr. G. Bouet in 1914 on lle Roumé, which is part of the [les de Los archipelago located in the Atlantic Ocean just several kilometres (ca. 8 km) offshore from Conakry, Guinea. In his description of A. insularis, Chabanaud (1918) only compared his new species with Agama kirkii Fig. 1. The two syntypes of Agama insularis Chabanaud, 1918 (MNHN 1918.041-042). Bonner zoologische Beiträge 56 24] Boulenger, 1884 from southern Africa and stated both as closely related. The only morphological differences not- ed were related to (1) head shapes (more elongate in A. insularis) and (2) body scalation (A. insularis has small- er scales, 120-130 scale rows around midbody versus about 90 in A. kirkii, and A. insularis had dorsal scales that are smaller compared to ventral scales. Based on a larger series of specimens collected from the type locality, which included adult specimens, PARKER (1939) reassessed the validity of A. insularis (material of the IRSNB, Appendix 1). In this publication PARKER (1939) also compared A. insularis with A. boulengeri Lataste, 1886, a morphologically similar species distrib- uted in Mauritania and Mali. He distinguished A. insularis from A. boulengeri by its more strongly carinate scales, especially on the occiput and lower surfaces of the tail; larger nasal, separated from the rostral by a single elon- gate scale; larger number of labial scales, 8—9 versus 11 and the colouration of adult males. Nevertheless, PARK- ER (1939) only described the adult colouration from pre- served specimens. Adult males are brownish black above with light specking, especially on the vertebral region; lower surfaces of the head, chest and abdomen are black- ish-grey; middle of the gular region is jet black. JOGER (1979) described the adult male colouration of living 4. boulengeri as follows: (1) ground colour pale gray-brown with transverse rows of white spots; (2) throat dirty white with longitudinal stripes, parts of the head gray-blue; (3) tail gray-blue and (4) a characteristic half-moon shaped black bar on the anterior margin of the shoulder. LAMBERT & MULLIE (1998) refer, beside the tail crest of the males, to the nuptial colouration of females as the most striking difference between the sexes of A. boulengeri. They de- scribe the males as drab with uniform purple-grey coloured scales dorsally, whereas the females have a bril- liant yellow mid-dorsal band with three approximately bat- shaped transverse bands on a bright orange background. LAURENT (1947) reported a series of Agama insularis from Iles Roumé and a neighboring island, Ile Kassa, and men- tioned that the characters given by PARKER (1939) were in agreement with his specimens. However, LAURENT (1947) described additional sources of morphological vari- ation, including 10 to 13 precloacal pores, 8 to 10 supral- abial and 7 to 10 infralabial scales. Because of differences in the number of scale rows around midbody (Ile Roumé: 115 to 125; Ile Kassa: 143 to 147) he suggested that the Ile Kassa population deserved subspecific status. The taxonomic status of Agama insularis has been changed since these early studies. GUIBE (1954) listed 4. insularis as valid in the type catalogue of the collection of the Muséum national d’histoire naturelle in Paris, al- though WERMUTH (1967) regarded this species as ques- tionable, since it was only known at this time from its type locality. JOGER (1979) reasserted the validity of A. insu- laris; however, Moony (1980) placed A. insularis in syn- onymy of 4. boulengeri, but without giving a reason. Nev- ertheless, BOHME (1985) followed JOGER (1979) and stat- ed that A. insularis and A. boulengeri are both valid species and probably closely related. Finally, BRyGoo (1988) and ULBER & BARTS (1997) treated A. insularis as a valid species, although without offering additional com- ments regarding this decision. Interestingly, another species of Agama from West Africa possessing a prominent tail crest has remained mostly overlooked since its description. Agama cristata was de- scribed by MOCQUARD (1905) from a single specimen col- lected from Bomanesco (Sankaran) in Guinea by M. A. Chevalier, and later donated to the Muséum national d’his- toire naturelle in Paris by M. Maurice de Rothschild. Moc- QUARD (1905) did not compare his new species with any other species and characterized it mainly by the large crest proceeding from the neck to the tail. Later, 4. cristata was apparently regarded as a synonym of A. sankaranica Cha- banaud, 1918 by GUIBÉ (1954) and WERMUTH (1967), pre- sumably because both taxa share the same type locality (A. sankaranica: Moussaia, Sankaran, Guinea; A. crista- ta: Bomanesco, Sankaran, Guinea). Later, BRYGOO (1988) confirmed the validity of A. cristata in his type catalogue presumably because of the tail crest, but again he was not followed by ULMER & Barts (1997). Despite the presence of a prominent tail crest in adult males shared between Agama cristata, A. boulengeri and the island endemic species, Agama insularis, a compari- son of these species remains to be conducted. In contrast to the previous three species, 4. sankaranica and Agama weidholzi Wettstein, 1932 are small and solitary-living species. The latter was included in the comparison, be- cause it is endemic to this region and so far only known from Senegal (e.g. WETTSTEIN 1932), Gambia (BOHME 2005), Mali (e.g. GRANDISON 1969) and Guinea-Bissau (MONARD 1940). Additionally, MONARD (1940) included specimens of A. weidholzi and A. sankaranica in the type series of A. boensis. (GRANDISON 1969, BOHME 2005) and for that reason a close relation to A. sankaranica could be possible. Our aim is therefore to compare the species of Agama with prominent tail crests in adult males (Agama cristata, A. boulengeri and the island endemic species, Agama insu- laris) to clarify their taxonomic status, distribution, and phylogenetic relationships. We also include other species of West African Agama in our study that lack prominent tail crests in adult males, but that are hypothesized to be close relatives because they occur in the same areas. 242 Philipp WAGNER et al.: Studies on African Agama VI. Taxonomic status of the West African Agama Fig. 2. The holotypes of a) Agama sankaranica (MNHN 1901.0395) and b) 4gama cristata (MNHN 1901.0394). Bonner zoologische Beiträge 56 243 MATERIAL & METHODS The voucher specimens examined in this study are acces- sioned in the following natural history collections: Col- lection of Jean Frangois Trape, deposited in Institut de Recherche pour le Developpement (IRD) Dakar, Senegal (TR); Institut royal des Sciences naturelles de Belgique (IRSNB); Museum national d’histoire naturelle de Paris, France (MNHN; Museum für Naturkunde, Berlin, Ger- many (ZMB) and Zoologisches Forschungsmuseum Alexander Koenig, Bonn, Germany. The type specimens of Agama cristata and A. insularis were included, how- ever, the type specimens of A. insularis are juveniles and therefore topotypical material was also used for this tax- on. Since several studies have now shown that DNA barcod- ing, especially when using the mitochondrial 16S rRNA gene, is a reliable tool in reptile or amphibian taxonomy (e.g. VENCES et al. 2005; BwoNG et al. 2009) molecular data were collected to calculate a simple neighbour-join- ing phylogeny (fig. 4) and to analyse the sequence vari- ation between species. A portion of the mitochondrial 16S rRNA gene of Agama insularis (ZFMK 88247; GenBank accession number: GU133326) from the type locality, as well as Agama cristata (TR555; GenBank accession num- ber: GU133325) from Guinea, Agama boulengeri (MNHN; Gen-Bank accession number: GU133324) from Mauritania, Agama weidholzi (ZFMK 75001; GenBank accession number: GU133328) from Gambia, Agama sankaranica (ZFMK 84992; GenBank accession number: GU133327) from an unknown locality and Agama aga- ma (ZFMK 15222 [neotype]; GenBank accession num- ber: GU133323) were sequenced. Acanthocercus atricol- lis (ZFMK 41748: GenBank accession number: GU133322) was used as outgroup. DNA was extracted using QuiAmp tissue extraction kits (Quiagen) or a modified Chelex-Protocol (WALSH et al. 1991, SCHMITZ 2003). The primers 16sar-L (light chain; 5’— CGC CTG TTT ATC AAA AAC AT - 3”) and 16sbr- H (heavy chain; 5” - CCG GTC TGA ACT CAG ATC ACG T — 3’) of PALUMBI et al. (1991) were used to am- plify a portion of the mitochondrial 16S ribosomal RNA gene. PCR cycling procedure was as described in SCHMITZ et al. (2005). PCR products were purified using Quiaquick purification kits (Quiagen). Sequences were ob- tained using an automatic sequencer (ABI 377). Sequences were aligned using ClustalX (THOMPSON et al. 1997; de- fault parameters) and manually checked using the origi- nal chromatograph data in the program BioEdit (HALL 1999). A preliminary phylogenetic tree was calculated in the program Paup* 4.0b10 (SworrorD 2002) using the Neighbor-joining algorithm (NJ) and 20000 pseudo-repli- cates to estimate node support. This program was also used to compute the uncorrected pairwise distances for all se- quences. The following measurements and scale counts were used to compare the different species in statistical analysis (for selected characters see table 1): Snout-vent length (SVL): measured from mental scale to cloaca; tail length (TL): measured from cloaca to tip of tail; Tail crest length (TcL): length of the tail crest from midpoint of pelvic region to tip of crest; head length (HL): measured from jugale to rostral scale; head height (HH): measured at the jugal-pos- torbital region; head width (HW): measured across the ju- gal-postorbital region just anterior to the external audito- ry meatus; Midbody scales (MS): scale rows around mid- body; tail crest scales (TcS): number of tail crest scales, counted from midpoint of pelvic region to tip of crest; dor- sal scales (DS): number of midline longitudinal dorsal scales, counted from midpoint of pectoral region to mid- point of pelvic region; cloacal pores (CP). Excel 2000 and SPSS (10.0) software packages were used to run statistical analyses. Hierarchical Cluster analysis and Principal Component Analysis (PCA) were used to evaluate the morphological data and to explore the phe- netic relationships between the taxa examined. RESULTS & DISUSSION Morphology. Significant differences were found when the effect of body sizes was removed from analyses (see fig. 3), but there is still a high level of overlap among species in all PC’s. Best results were found in PC 1 and 2 (PC I= 41.379% of variance; PC 2= 25.825% of variance; PC 3= 24.098% of variance; PC 4= 8.699% of variance) and are visualized in fig. 3. Agama insularis has a significantly larger relative head height, head width and head length than A. boulengeri, but lower average of mid-body scale rows and significantly smaller relative head length than A. cristata. Agama cristata differs significantly in a high- er average of mid-body scale rows and larger relative head length from both 4. insularis and A. boulengeri. Agama boulengeri differs significantly in smaller relative head height, head width and head length from both 4. insularis and A. cristata. Males have significantly lower tail-crest- scales and tail-crest-length than either 4. insularis or A. cristata. Differences in pholidosis between A. boulengeri and A. cristata are low and mostly overlapping (see table 1), which is unsurprising within the genus Agama. But in average, A. cristata has a higher count of scale rows around midbody than A. boulengeri. Also the differences in colouration between 4. boulengeri and A. cristata are low. Small differences in colouration were found between Agama boulengeri on one hand and A. cristata on the oth- er hand. Especially the colouration of the throat in adult 244 Philipp WAGNER et al.: Studies on African Agama VI. Taxonomic status of the West African Agama A 2.000004 PA 1.00000 4 o de A en = = = 5 Damon +. c 5 + o 0 = hd ° fo} A + 2 000007 u 2 id fs) £ hh a + -2,00000 4, : a : -2,00000 -1.00000 0.00000 1.00000 Principal component 2 Fig. 3. Projection of the first and second principal component from a PCA run on 16 individuals assigned to + = Agama bou- lengeri, &= Agama insularis @ = Agama cristata. males is distinct. In A. boulengeri the throat is dirty white with longitudinal stripes, whereas in A. cristata the basal parts of the throat are dark bluish-black. Genetics. As obvious from fig. 4, Agama boulengeri is very distinct from the two other known species with tail crests (A. insularis; A. cristata), and it is found in a basal position to all other Agama species included in this study (NJ: 70). In contrast, the two other species, A. insularis and A. cristata form a maximally supported clade (NJ: 100) that is the sister clade of A. weidholzi (NJ: 84) with this subclade significantly separated from its sister clade containing 4. agama and A. sankaranica as sister taxa, though this latter subclade receives hardly any support (NJ: 56). This low support comes as no surprise as other studies show clearly that 4. agama and A. sankaranica are not very closely related (LEACHE et al. 2009). Table 2 summarizes the uncorrected p-distances for 496 bp of the 16S rRNA gene. Agama boulengeri shows the highest genetic differences to all other species with val- ues ranging from 12.1%-15.9% thus showing that it is not closely related to any of the other included Agama species, and is far removed from the two other taxa with tail crests (A. insularis: 15.5% A. cristata: 14.1%). Thus the genet- ic data clearly confirm the specific distinctness of A. cristata/A. insularis from A. boulengeri. Although genetic interspecific differences range at mod- erately high values ranging from 8.0%-11.9%, the genet- ic distance between A. insularis and A. cristata is low at only ca. 0.7% (corresponding to only differing basepairs). The low level of sequence divergence observed between A. cristata and A. insularis is consistent with the hypoth- esis that these taxa represent a single species. Agama boulengeri is distinct in pholidosis, colouration and genetics from A. insularis and A. cristata. But no signif- icant differences in morphology and genetics were found between A. cristata and A. insularis. It is important to note that this result is based on an examination of type speci- mens, topotypical material, and freshly collected materi- al (see appendix). Therefore we consider Agama insularis Chabanaud, 1918 as a junior synonym of the revalidated Agama cristata Mocquard, 1905. Agama cristata MOCQUARD, 1905 1905 Agama cristata Mocquard, Bull. Mus. Hist. Nat. 11: 288. 1918 Agama insularis Chabanaud, Bull. Mus. Hist. Nat., 3: 2, 3. Terra typica: “ile Rooma (groupe des ¡les de Las)” (= Roumé Island, Los Islands), Guinea; Syn- types. MHNP 1918.41, 1918.42 1939 Agama insularis — Parker, Mém. Mus. Roy. Hist. Nat. Belg., Ser. 2, 15239. 1947 Agama insularis — Laurent, Bull. Mus. Roy. Hist. Nat. Belg., 23 (16): 5. 1954 Agama insularis — Guibé, Catalogue des Types de Lézards du Museum national d Histoire naturelle: 26. 1954 Agama cristata — Guibé, Catalogue des Types de Lé- zards du Muséum national d’Histoire naturelle: 26. 1967 Agama cristata — Wermuth, Das Tierreich, 86: 11. 1967 Agama insularis — Wermuth, Das Tierreich, 86: 16. 1979 Agama insularis — Joger, Salamandra 15: 36. 1982 Agama insularis — Welch, Herpetology of Africa: 48. 1985 Agama insularis — Böhme, Proc. Intl. Symp. Afr. Vertebr., Bonn, 1985: 471. 1988 Agama insularis — Brygoo, Bull. Mus. Nat. Hist. Na- turelle 10, supplement 3, 1—-56. 1997 Agama insularis — Ulber & Barts, Herprint Interna- tional, Bredell, South Africa, 418 pp. 2008 Agama maria nom. nov. Barabanov, Russ. J. Herp. 15: 206. 2009 Agama cristata — Wagner & Böhme, Russ. J. Herp. 16: 161-162. Holotype. MNHN 1901.0394. Terra typica. Bomanesco, Sankaran, Guinea. Comment on Nomenclature. BARABANOV (2008) recent- ly proposed a nomen novum for Agama cristata Moc- quard, 1905 because he recognized this taxon as a primary junior homonym of Agama cristata Merrem, 1820, recent- ly known as the iguanid lizard Corytophanes cristatus. However, WAGNER & BÖHME (2009) did not accept this suggestion because in accordance to article 23.9.5 of the ICZN (1999) the author failed to refer the case to the Com- Bonner zoologische Beiträge 56 245 Acanthocercus atricollis 0.1 Agama boulengeri Agama agama - Neotype Agama sankaranica Agama weidholzi Agama cristata Agama insularis Fig. 4. Neighbor-joining tree based on 496 bp of the mitochondrial 16S rRNA gene. Values above the nodes represent bootstrap (20000 pseudo-replicates) values in percent for the neighbor-joining analysis. Significantly supported values are in bold. Values below 50% are not shown. mission of Zoological Nomenclature which would be nec- essary because both taxa are not congeneric since 1827, where BOIE (in SCHLEGEL 1827) connects Merrem’s crista- ta as type species with his newly described genus Cory- thophanes. Diagnosis. A large species of Agama (total length of adult male up to 340 mm), which is characterized by its bluish- black colouration on the throat and the large nuchal and tail crests of adult males. The species differs from nearly all other Agama species in having a large tail crest, usually reaching the last third of the tail. Only three other species, A. boulengeri, A. kirkii and A. knobeli, have such a prominent tail crest. Agama cristata differs in detail from: (1) A. boulengeri in (a) higher numbers of scales rows around midbody (97 to 123 in 4. boulengeri and 111 to 147 [latter value fide LAURENT 1947] in A. cristata); in 246 Table 1. Comparison of measurements and pholidosis. Agama boulengeri Agama cristata Philipp WAGNER et al.: Studies on African Agama VI. Taxonomic status of the West African Agama Agama weidholzi* Agama gracilimembris* average (min—max), n average (min—max), n SVL 88.4 (62.5-129.8), 22 92.0 (69.0-123.0), 16 54-65 44-57 TL 185.0 (119.2-236.2), 12 149.3 (98.8-215.0), 12 - - HH 11.0 (8.1-16.3), 22 12.3 (8.6-17.6), 16 = - HW 16.1 (12.0-24.0), 22 17.5 (11.1-23.8), 16 - = HL 23.8 (17.0-33.6), 22 25.3 (20.1-33.3), 16 - - SAM 112.9 (97-123), 22 121.4 (111-136), 13 68-82 70-85 TCS 71.3 (62-78), 10 77.5 (75-80), 6 - = PP 11.3 (8-14), 12 11.4 (10-13), 7 - - SVL= snout-vent length; TL= tail length; HH= head height; HW= head width; HL= head length; SAM= scale rows around midbody; TCS= tail crest scales; PP= precloacal pores. All measurements in mm;.*= after GRANDISON 1969 Table 2. Uncorrected p-distances for 496 bp of the mitochondrial 16SrRNA sequences used in this study. Taxa 1 1 Acanthocercus atricollis ZFMK 41748 - 2 Agama boulengeri MNHN 0.2135 3 Agama agama ZFMK 15222 0.2029 4 Agama cristata TR555 0.1937 =) Agama insularis ZFMK 88247 0.2022 6 Agama sankaranica ZFMK 84992 0.1929 7 Agama weidholzi ZFMK 75001 0.2117 (b) having smaller relative head height, head width and head length; and (c) males of A. boulengeri have a low- er number of tail-crest-scales and a lower length of the tail- crest. (d) male coloration (larger ocelli, lower parts of the throat and gular fold deep bluish-black). (2) A. kirkii in (a) having smaller body scales; (b) dorsal scales are smaller than ventral scales; and (c) in having no narrow blue and white banded tail (d) a geographic dis- tribution confined to northern parts of southern Africa (Malawi, Zambia, Zimbabwe, Mozambique, Botswana). (3) A. knobeli is herein recognized as a valid species, be- cause we recognized striking dissimilarities in morphol- ogy between A. knobeli and the former nominate form A. atra: Agama atra is lacking the prominent large tail crest and has more spinose scales as A. knobeli, but further re- search on the relationships of these two species is in need. 2 3 4 3 6 7 0.1348 — 0.1405 0.1040 — 0.1553 0.1123 0.0072 2= 0.1209 - 0.0803 0.1036. 0.119277 0.1590 0.1069 0.0892 = 0.1002 0.1068 However, A. knobeli differs from A. cristata in (a) hav- ing larger body scales, (b) dorsal scales in same size as ventral scales and (c) in having a pale vertebral stripe in nuptial coloured adult males, and (d) a geographic distri- bution restricted to Namibia. Colouration in alcohol. The holotype of A. cristata is uni- form brown. Syntypes of A. insularis are grey to brown in different shades, but typical ocelli of Agama females and juveniles are visible. One subadult female specimen (TR 2353) from the island Ile de Roumé (Los islands) is uniform brown with scattered orange stripes and bars on the body and shows dark stripes on a white throat. Even juveniles from the same locality show dark framed ocel- li on the body and white dots on the head. In both juve- niles (MNHN 2008.0023, TR 2352) a lateral orange stripe is obvious. Bonner zoologische Beiträge 56 247 Fig. 5. Images of the following Agama species (in life): A= ‘Agama insularis’ from island Roumé (photo by Dr. Guy Kremer). B= Agama cristata, male, from Kindia, Pastoria, Guinea (photo by Ivan Ineich). C= Agama cristata, male, from Chutes de Kinkon, Guinea (photo by Ivan Ineich). D= 4gama cristata, fe- male, from Chutes de Kinkon, Guinea (photo by Ivan Ineich). E= Agama boulengeri, male, from Mauritania (photo by Hemmo Nickel). F= Agama boulengeri, female, from Mauritania (photo by Hemmo Nickel). G= Agama boulengeri, male, from Chütes de Felou, Mali (photo by Ulrich Joger). H= Agama weidholzi, male, from Mali (photo by Ulrich Joger). I= Agama cristata, male, from Kindia, Pastoria, Guinea (tail crest in detail) (photo by Ivan Ineich). 248 Philipp WAGNER et al.: Studies on African 4gama VI. Taxonomic status of the West African Agama Mauritania 14 CANS Triana eas See te A ela > iz mm Fig. 6. Distribution map of Agama boulengeri, A. cristata and A. weidholzi. Agama boulengeri: MAURITANIA: 1= Choúm; 2= Ben Amira; 3= Aggui; 4= Guelta Molomhar; 5= Chinguetti; 6= Guelta Hamdoun: 7= Kanoal; 8= Terjit; 9= Zerga Mountains; 10= Tintáne; 11= Irıji; 12= Tidjikja; 13= Tichit; 14= Guelta Fanar; 14= Guelta Mat- mata; 15= Achram; 16= Ain El Ghaire; 17= Bou Bleiine; 18= Kifa; 19= Bougarı; 19a= Guelta Oumm Lebare; 19b= Guelta Me- traucha; 19c= Oumm El Khez; 20= Ayoún El Atroús; 21= between Timbedgha and Ayoún El Atrous; 22= Mbout; 23= Sélibabi (for details see Padial 2006). MALI: 24= Medine (type locality of A. boulengeri); 25= Chutes du Félou (Joger 1981). Agama cristata: GUINEA: 26= Moussaia, Sankaran (type locality of A. cristata); 27= Ile Rooma, Iles de Los (type locality of A. in- sularis), 28= Chutes de Kinkon near Pita; 29= Kindia. Agama weidholzi: THE GAMBIA: 30= Bwiam. SENEGAL: 31= Boughari, Ca- samance; 32= between Tiara and Mantiankani; 33= 13 km Southwest of Kolda; 34= Tabadienke, 30 km South of Diallakoto; 35= 20 km S of Médina Gounas; 36= 12 km West of Kounkané; 37= 5 km East of Darsalam (Niokolo-Koba National Park). GUINEA- BISSAU: 38= Madina do Boé, Pitche. MALI: 39= 9 km North of Fatao (14°24’N, 9°29’ W): 40= 20 km West of Kita (13°03’N, 9°42’ W); 41= 5 km East of Kita (13°03’N, 9°25’ W); 42= between Négala and Kassaro (12°55’N, 8°40’ W); 43= Négala (12°52’N, 8°27’W) Colouration in life (see fig. 5). Males. Body brownish, scattered with dark framed white to creamy ocelli, usual- ly bigger ocelli form distinct rows. Lips creamy to bluish- creamy. Head and neck brownish with a pale stripe un- derneath the eye. Lower parts of the throat and gular fold deep bluish-black with strips running to the chin. Poste- rior part of the neck, body, parts of the hindlimb and tail sometimes speckled with white and dark scales. Some- times a pale vertebral band between the limbs is obvious. Tail at the base pale speckled black, downwards brown- ish. Belly and underside of the limbs creamish to dark grey. Females. Body brownish, scattered with dark framed white to creamy ocelli, when pregnant with dark orange to red coloured bands on the lateral body sides between the limbs. Sometimes lateral parts of the body white fol- lowed by a darker band and pale brownish on vertebral parts. Head and neck brownish with pale to yellow dots on the upper head and a pale stripe underneath the eye. Lips creamy to white. Fig. 7. Mite pocket-like structure (indicated by an arrow) at a specimen of Agama cristata from Kindia, Pastoria in Guinea. Bonner zoologische Beiträge 56 249 Fig. 8. Comparison of tail regenerates with x ray images (point of fracture indicated by arrows): A) original tail of Agama bou- lengeri (ZMB 22922). B, C) long tail regenerate of A. boulengeri (ZMB 55884). D, E) short tail regenerate of A. boulengeri (ZMB 55885). F, G) short tail regenerate of A. boulengeri (ZMB 55887) i) Un Philipp WAGNER et al.: Studies on African 4gama VI. Taxonomic status of the West African Agama Fig. 9. A remarkable bifurcated tail in a specimen of Agama lionotus lionotus (Kenya: Kitui District, NMK L/1252). Bonner zoologische Beiträge 56 2 Juveniles. Juveniles are only known from preserved vouchers. Distribution. A. cristata ıs so far only known from Guinea. Beside the type localities of the two taxa, the species 1s also recorded from the hydroelectrical power station at Kinkon water falls (11°2°54.9”N 12227” 1.8”W; about 820 m a.s.1.) and the Pastoria station (former Insti- tute Pasteur) (10°5’36.5”N 12°50’21”W; about 388 m a.s.l.). Agama boulengeri ıs endemic to the oriental region from Senegal, western Mali and Mauritania (GENIEZ et al. 2004), which is a similar distribution pattern as in Taren- tola parvicarinata JOGER, 1980 (Sauria: Phyllodactylidae) (Joger 1980). JOGER (1979) regarded the species as endem- ic for the Variscian Fold Mountains between the Adrar Mountains in Mauritania and northern Senegal (see fig. 6). The limits of the geographic distribution of Agama cristata are less clear, and 1t is therefore more difficult to discuss their speciation and biogeography. However, the Senegal River, which separates A. boulengeri from A. cristata may have functioned to keep these lineages sep- arated. Relationships. Morphologically, 4. cristata is very sim- ilar to A. boulengeri. But preliminary phylogenetic results indicate a sister taxa relation between A. cristata and A. weidholzi, with A. boulengeri placed as the sister taxon to all species analyzed (fig. 4). These results do not cor- relate with morphology, because A. weidholzi is a small, ground dwelling species without any tail crest, and is sim- ilar in morphology to the west and central African species A. gracilimembris. GRANDISON (1969) compared series of the two species and stated both as closely related. GARTSHORE (1985) also mentioned similarities in the colouration of gravid females between 4. gracilimembris, A. doriae and A. sankaranica. There are also similarities between 4. weidholzi, A. gracilimembris and A. sankaran- ica in behavioural aspects. All these are solitary species that lack social interactions outside of the breeding sea- son (GARTSHORE 1985), and males occupy home ranges but do not defend territories. Unfortunately, 4. gracilimembris could not be included in the phylogenetic analysis because of a lack of fresh DNA samples of this rare species. Biology. Agama cristata, like A. boulengeri, is a rupi- colous species living on rocks in syntopy with Trachylepis quinquetaeniata (Sauria: Scincidae). Also Trachylepis affı- nis was recorded in the area nearby. In males, a mite pock- et behind the tympanum is sometimes obvious (see fig. Un 7). These mite pocket-like structures are rare in agamıd lizards. ARNOLD (1986) described nuchal pockets from iguanian lizards whereas BERTRAND € Mopry (2004) re- ported the first pocket-like structures in an Agama lizard (Agama caudospinosa). In this case, the newly described mite Prervgosoma livingstonei was found heavily concen- trated in a skin-fold in front of the forelimb. Here, we re- port a similar pocket-like structure in front of the forelimb of an Agama cristata specimen (fig. 7, indicated by an ar- row), which is similar to the pocket shown in BERTRAND & Mopry (2004). However, these species are not close- ly related, and we predict that these structures are more common in Agama lizards than they are currently believed to be. Remarks on caudal anatomy. Some specimens of Aga- ma cristata and A. boulengeri (e.g. a subadult female; TR 2353) show remarkable tail regeneration (fig. 8). Complete caudal regeneration in Agamid lizards is relatively rare, and regenerated tails are normally short and possess a knob-like end (e.g. Agama lionotus, see fig. 9). This is probably because Agamidae, like Chamaeleonidae and Platynota, lack a cartilaginous stratum or plane (ANANJE- VA 1985, ANANJEVA & DAnovV 1991). As an example, the total length of a specimen of A. cristata (TR 2352) is 89.5 mm; with a tail length (TL) in total of 68.8 mm and a re- generate length (RL) of 36.2 mm (RL/TL= 0.53). There- fore, the regenerate measures a bit more than half of the length of the tail. The total length of the regenerate is longer than the documented bifurcated tail of Laudakia caucasia (ANANJEVA & DANOV 1991) with a length of 23.1 mm. This phenomenon is also visible in 4. boulengeri. Three specimens of the ZMB collection show a relatively long, slender and not knobbly regenerated tail. ZMB 55885 has a TL of 45.8 mm with a RL of 19.7 mm (RL/TL= 0.43). ZMB 55887 has a TL of 75.2 mm with a RL of 16.8 mm (RL/TL= 0.22), whereas ZMB 55884 has a TL of 109.6 mm with a RL of 46.4 mm (RL/TL= 0.42) (see fig. 8). Also a specimen of the species Agama lionotus lionotus Boulenger, 1896 from East Africa (Kenya: Kitui District, NMK L/1252) shows a remarkable bifurcated tail, but here, in difference to the bifurcated tail reported by ANAN- JEVA & DANOV (1991) from Laudakia caucasia, only one tail tip is a regenerate (see fig. 9). Acknowledgements. We are thankful to G. Lenglet of the In- stitute royal des Sciences naturelles de Belgique (IRSNB) for the loan of important material. 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LEACHE, A. D., CHONG, R. A., PAPENFUSS, T. J., WAGNER, P., BÖHME, W., SCHMITZ, A., RÖDEL, M.-O., LEBRETON, M., In- EICH, I., CHIRIO, L., BAUER, A., ENIANG, E. A. & S. BAHA EL Din (2009): Phylogeny of the genus Agama based on mito- chondrial DNA sequence data. Bonner Zoologische Beiträge 56: 273-278. LOVERIDGE, A. (1933): Reports on the scientific results of an ex- pedition to the Southwestern highlands of Tanganyika terri- tory, VII, Herpetology. Bulletin of the Museum of Compara- tive Zoology at Harvard College 74: 195-416. MCLACHLAN, G. R. (1981): Taxonomy of Agama hispida (Sau- ria: Agamidae) in southern Africa. Cimbebasia ser. a 5 (6): 219-227. MOCQUARD, M. F. (1905): Note préliminaire sur une collection de reptiles et de batraciens offerte au Muséum par M. Mau- rice de Rothschild. Bulletin du Museum nationale d’Histoire naturelle Paris 11: 285-290. MONARD, A. (1940): Résultats de la mission scientifique du Dr. MONARD en Guinée Portugaise (1937-1938). VIII Reptiles. Arq. Mus. Bocage, Lisboa 11: 147-180. Moopy, S. M. (1980): Phylogenetic and Historical Biogeograph- ical Relationships of the Genera in the Family Agamidae (Rep- tilia: Lacertilia). Unpublished Ph.D. Thesis, Univ. of Michi- gan, 373 pp. PADIAL, J. M. (2005): A new species of Agama (Sauria: Agami- dae) from Mauritania. Herpetological Journal 15 (1): 2735. PALUMBI, S. R., MARTIN, A., ROMANO, S., MCMILLAN, W. O., STICE, L. & G. GRABOWSKI (1991): The simple fool’s guide to PCR. Department of Zoology and Kewalo Marine Labo- ratory, Hawaii, 47 pp. PARKER, H. W. (1939): Résultats scientifiques des croisieres du Navire-École Belge “Mercator”. V. Reptilia et Amphibia. Mém. Mus. Roy. Hist. Nat. Belg., Ser. 2, 15: 85-90. SCHLEGEL, H. (1827): Notice sur l’Erpetologie de l’ile de Java par M. Boie. Bull. Sci. Nat. Géol. 9: 233-240. SCHMITZ, A. (2003): Taxonomic and phylogenetic studies on scincid lizards (Reptilia: Seincidae). Unpubl. PhD thesis, Uni- versity of Bonn, 262 pp. SCHMITZ, A., INEICH, I. & L. CHIRIO (2005): Molecular review of the genus Panaspis sensu lato in Cameroon, with special reference to the status of the proposed subgenera. Zootaxa, 863: 1-28. SWOFFORD, D. L. (2002): PAUP*: Phylogenetic analysis using Parsimony (*and other methods), version 4.0b10. Sunderland, MA: Sinauer Associates. THOMPSON, J. D., GIBSON, T. J., PLEWNIAK, F., JEANMOUGIN, F. & D. G. HIGGINS (1997): The ClustalX windows interface: flexible strategies for multiple sequence alignment aided by quality analysis tools. Nucleic Acids Research 24: 4876-4882. THYS VAN DEN AUDENAERDE, D. F. E. (1963): Les Agamidae du Congo: les especies et leur distribution géographique. Revue de Zoologie et de Botanique Africaines 68: 203-250. ULBER, T. M. & M. Barts (1997): Catalogue of valid species and synonyms Volume 4 (Agamidae/Uromastycidae). Herprint International, Bredell, South Africa, 418 pp. VENCES, M., M. THOMAS, M., VAN DER MEIJDEN, A., CHIARI, A. & D. R. VIEITES (2005): Comparative performance of the 16S tRNA gene in DNA barcoding of amphibians. Frontiers in Zoology, 2: 5. Bonner zoologische Beiträge 56 253 WAGNER, P. (2007): Studies in African Agama I: On the taxo- nomic status of Agama lionotus usambarae Barbour & Loveridge, 1928 (Squamata, Agamidae). Herpetozoa 20: 69-73. WAGNER, P. & W. BOHME (2009): On the status in nomenclature of Agama cristata Mocquard, 1905. A reply to Andrei Bara- banov. Russian Journal of Herpetology 16 (2): 161-162. WAGNER, P., BURMANN, A. & W. BOHME (2008a): Studies on African Agama II. Resurrection of Agama agama kaimosae Loveridge, 1935 (Squamata: Agamidae) from synonymy and its elevation to species rank. Russian Journal of Herpetology 15: 17. WAGNER, P., KRAUSE, P., BURMANN, A. & W. BOHME (2008b): Studies on African Agama III. Resurrection of Agama aga- ma turuensis LOVERIDGE, 1932 (Squamata: Agamidae) from synonymy and its elevation to species rank. Salamandra 44: 35-42. Appendix. Material examined Agama boulengeri: MALI. Kayes: Chútes de Félou (ZFMK 20058, 22176 — 180, 25481 — 485). MAURITANIA. Adrar: Terjit (ZFMK 76917); 20km N Atar (ZFMK 79729); Ayoun-el Atrons (ZFMK 83825 — 827); Guelta Metraucha (ZFMK 76862 — 864, 76868); Galoula (=Passe de Galoula) (ZMB 32583, ZMB 55882 — 888, ZMB 32584). Agama cristata: GUINEA. Chútes Kinkon (MNHN 2006.0553, MNHN 2006.0554); Kindia, Institut Pasteur (MNHN 2006.0555, MNHN 2006.0556). ‘Agama insularis”: GUINEA. Iles des Los, Ile Roumé (MNHN 2008.0023, TR 2352 — 53; IRSNB 1392 a-c, 1728, 1733, 1729 a-e; MNHN 2008.0023, ZFMK 88247). WAGNER, P., WILMS, T., BAUER, A. & W. BOHME (2009a): Stud- ies on African Agama V. On the origin of Lacerta agama Lin- naeus, 1758 (Squamata: Agamidae). Bonner Zoologische Beiträge 56 (4): 215-223. WAGNER, P., BAREJ, M. F. & A. SCHMITZ (2009b): Studies on African Agama VIL. A new species of the Agama agama-group (Linnaeus, 1758) (Sauria: Agamidae) from Cameroon & Gabon, with comments on 4gama mehelyi Tornier, 1902. Bon- ner Zoologische Beiträge 56 (4): 285-297. WALSH, P. S., METZGER, D.A. & R. Hısucht (1991): Chelex 100 as a medium for simple extraction of DNA for PCR-based ty- ping from forensic material. BioTechniques 10: 506-513. WETTSTEIN, O (1932): Eine neue Eidechse aus Senegambien. Zool. Anz., Leipzig 9% 11/12): 303-305. WERMUTH, H. (1967): Liste der rezenten Amphibien und Rep- tilien: Agamidae. Das Tierreich 86, 1—xiv, 1-127. Agama weidholzi: GAMBIA. Bwiam (ZFMK 75001). MALI. 8km E Kassaro (ZFMK 20060); 5km E Kita (ZFMK 20061); 20km W Kita (ZFMK 20062); 9km N Fatao (ZFMK 20063 — 65); bet- ween Negala and Kassaro (ZFMK 20059). SENEGAL. 5km SO Darsalam (ZFMK 20066 — 068); 20km S Medina Gounas (ZFMK 20069); 13km SE Kolda (ZFMK 20066 — 078); 12km W Kounkané (ZFMK 20070 — 071); between Tiara and Manti- ankanı (ZFMK 20072). Bonner zoologische Beiträge Band 56 Heft 4 Seiten 255-258 Bonn, November 2009 Remarks on the type specimen of Gonocephalus mjobergi Smith, 1925 (Sauria: Agamidae) Wolfgang DENZER! & Ulrich MANTHEY? ! Physical & Theoretical Chemistry Laboratory, University of Oxford South Parks Rd, Oxford OX1 30Z, United Kingdom; E-Mail: denzer@physchem.ox.ac.uk. 2 Kindelbergweg 15, D-Berlin, Germany. E-Mail: Manthey. SSEAH @t-online.de. Abstract. The type specimen of Gonocephalus mjobergi Smith, 1925 has been re-examined. A remarkable character, longitudi- nal folds in the gular region has been discovered which is unique among agamids of the genus Gonocephalus Kaup. G. mjober- gi has been compared to members of the genera Pryctolaemus and Mantheyus and it is concluded that these genera form a mono- phyletic group and G. mjobergi is referred to as “Gen. A.” mjobergi until further material becomes available. Keywords. Agamidae, Gonocephalus mjobergi, type specimen. INTRODUCTION Historically, the genus Gonocephalus Kaup, 1825 includ- ed at some stage species from the genera Acanthosaura Gray, 1831, Coryphophylax Blyth, 1860 and Hypsilurus Peters, 1867 (see for example BOULENGER 1885, SMITH 1935 and WERMUTH 1967). MoobY (1980) removed all species belonging to the Melanesian-Australian radiation and Andaman-Nicobar species from its synonymy with Gonocephalus Kaup and revived the genera Coryphophy- lax (Blyth) and Hypsilurus (Peters) as well as Arua (Do- ria). The Hypsilurus and Arua species complex has been reviewed in detail more recently by MANTHEY & DENZ- ER (2006). Most Gonocephalus species were reviewed by MANTHEY & DENZER in the early 19901es and arranged in species groups (MANTHEY & DENZER 1991) on grounds of mor- phological similarities. There are currently five groups recognised: a) bornensis group including beyschlagi, liogaster, bor- nensis, belli, (denzeri) and the Philippine species sem- peri, interruptus and sophiae assigned to this group here. b) chamaeleontinus group comprising chamaeleontinus, doriae (2 subspecies, G. d. doriae and G. d. abbotti) and kuhlii. c) megalepis group comprising megalepis, klossi, lacuno- SUS. d) robinsonii group comprising robinsonii and mjobergi (tentatively assigned to this group by MANTHEY & DEN- ZER- 1991) e) grandis group comprisiung only grandis. Overall there are currently 17 species recognised with G. denzeri pending further material and verification. MAN- THEY & GROSSMANN (1997) synonymised G. denzeri with G. bornensis because G. bornensis reveals a high variabil- ity in morphometric characters and pholidodis which ın- cludes characters of G. denzeri. The examination of ad- ditional 55 specimen and images of about 30 living spec- imen (MANTHEY unpubl.) including the type specimens of both species (RMNH 3043, 3044 and ZFMK 50527, 50528) gives rise to the assumption that G. bornensis and G. denzeri are conspecific. We note, however, that if just the type specimens are considered there are significant dif- ferences in dorsolateral pholidosis. While in G. bornen- sis dorsolateral scalation is homogeneous G. denzeri clear- ly shows clusters of enlarged scales dorsolaterally. Future phylogenetic studies using biomolecular and genetic meth- ods will hopefully resolve this issue. Morphologically the genus Gonocephalus is highly di- verse and solely defined by a few characters, i. e. basal scales adjacent to the nuchal and dorsal crests, a transverse gular fold, a sharp canthus rostralis and an “angled” supra- 256 Wolfgang DENZER & Ulrich MANTHEY: Remarks on the type specimen of Gonocephalus mjobergi Fig. 1. Dorsolateral view of the type specimen (BMNH 1946.8.13.87) showing the oblique rows of enlarged dorsolateral scales. Fig. 2. Ventral view of the type specimen (BMNH 1946.8.13.87); note the clearly discernible longitudinal folds on the gular re- gion. Bonner zoologische Beiträge 56 257 ciliary ridge. Some of these characters are not well devel- oped in several of the species groups above. The most re- semblance to these generic features is exhibited by mem- bers ofthe G. chamaeleontinus species group. Several species such as G. grandis and G. robinsonii show hardly any resemblance to the remaining species of the genus. An isolated species was described by SMITH (1925) as Gonocephalus mjobergi based on a single female spec- imen collected by Mjóberg on Mt. Murud at an altitude of 7000 feet (2134 m). Until today this has remained the only known adult specimen of the species. Owing to its superficial resemblance and shared ecological niche G. robinsonii and G. mjobergi were tentatively grouped by MANTHEY & DENZER (1991). In the course of our inves- tigations into high altitude forms of Gonocephalus in par- ticular G. robinsonii (Boulenger, 1908) we re-examined the type specimen of G. mjobergi (Smith, 1925). RESULTS Gonocephalus mjobergi was described on the basis of a single female collected on Mount Murud, Sarawak. Sev- eral features of type specimen are in poor condition. We re-examined the holotype (BMNH 1946.8.13.87, former- ly BMNH 1924.8.28.8) and discovered characteristics not mentioned in the original description. While the gular sac is incomplete (SMITH I. c.: “the tip missing”) it is difficult to determine its actual size. From the original drawings it could already be concluded that it might reach further on- to the chest than in any other species of Gonocephalus ex- cept perhaps G. robinsonii. When we examined the holo- type we were surprised that despite the incompleteness it was clear that the onset of a complete the gular sac would have reached the clavicular region. A feature clearly dis- tinguishing G. mjobergi from all other Gonocephalus is the possession of enlarged dorsolateral scales forming oblique rows. The first row is located in the shoulder re- gion and two distinct rows across the back as can be seen in Fig. 1. Dorsolaterally enlarged scales are present in some species of Gonocephalus from Sumatra, in partic- ular G. megalepis (Bleeker, 1860), G. klossi Boulenger, 1920 and G. lacunosus Manthey Denzer, 1991 but nev- er in such a geometrical arrangement as can be found in the type specimen of G. mjobergi. Another feature in G. mjobergi is the possession of an enlarged platelike scale below the tympanum which is separated from the tympa- num by two rows of small scales. Additionally, we dis- covered that G. mjobergi possesses two parallel longitu- dinal gular folds a feature not mentioned at all by Smith (L.c.). No other Gonocephalus species possesses this out- standing character which can clearly be seen in figure 2. Both folds start on the distal part of the gular region ap- proximately bordering the serrated edge in the middle line Fig. 3. Ventral view of the juvenile Gonocephalus sp. collec- ted on Mt. Murud (ZRC 2.5953). of the pouch. The outer one runs nearly parallel to the cen- tre line of the gular pouch and continues onto the anteri- or part of the chest. It partially conceals the Gonocephalus typical transverse fold. The inner folds are shorter and curve inside towards the centre line. This feature is very similar to the longitudinal gular folds known from species of the genera Ptyctolaemus Peters, 1864 and Mantheyus Ananjeva & Stuart, 2001. ANANJEVA & STUART (2001) de- pict the gular region of the latter two genera. The arrange- ment and colouration in G. mjobergi 1s rather compara- ble to that in Pryctolaemus. It consists of two rows (gray- ish white in alcohol) with dark colouration of the skin in between. This indicates that this character independently developed in Ptyctolaemus, Mantheyus and Gonocephalus mjobergi and seems to be an autapomorphy. All three species also lack a transverse gular fold present in Gono- cephalus. However, G. mjobergi is by no means congeneric with ei- ther Ptyctolaemus or Mantheyus. Both genera Pryctolae- mus and Mantheyus have scaled tympani as opposed to G. mjobergi; additionally, Mantheyus has femoral pores. Within this cluster only G. mjobergi shows oblique rows 258 Wolfgang DENZER & Ulrich MANTHEY: Remarks on the type specimen of Gonocephalus mjobergi of enlarged dorsolateral scales. Because of the type spec- imen of G. mjobergi being a female we hypothesize that the gular sac will be even larger in male specimens which would distinguish it further from the genera discussed here with the exception of G. robinsonii. As a consequence we propose to combine Ptyctolaemus, Mantheyus and G. mjobergi ın a genus group with longi- tudinal gular folds being an autapomorph character for the cluster constituting a synapomorphy for the three genera, hence pointing towards monophyly of this group. Owing to the lack of specimens for G. mjobergi we pre- fer to take a conservative view and not define a new mono- typic genus until more material becomes available. Here we suggest to refer this species as “Genus A.” mjobergl. I. Das (University of Malaya) collected a specimen of a juvenile Gonocephalus on Mt. Murud and referred to 1t as G. mjobergi. This specimen has been depicted in Das (2006: 84) and was deposited in the Singapore Zoologi- cal Reference Collection (ZRC 2.5953). The colouration in life is a vivid green with darker reticulate pattern. This is in accordance with SMITH’s (l. c.) description of the colouration of the female which has been described as „grass-green in life“. However, several other species of Gonocephalus show this kind of colouration. In particu- lar juvenile specimens of G. liogaster (Günther, 1872) and G. bornensis (Schlegel, 1848) show a large resemblance to the specimen depicted by Das (l.c.) as can be seen for example in fig. 136 of MANTHEY & GROSSMANN (1997) and fig. 254 of MALKMUS et al. (2002). Additionally, the juvenile specimen from Mt. Murud does not possess enlarged dorsolateral scales or an enlarged scale below the tympanum as described for the type. Such a feature 1s, however, not depending on age and should be developed early on in life. A ventral view including de- tails of the gular region is shown in figure 3. As can be seen the gular region of the juvenile specimen does not show any feature resembling parallel longitudinal folds and therefore propose that ZRC 2,5953 should be referred to as Gonocephalus sp. until more material for compari- son becomes available. Acknowledgements. We would like to thank Dr. Colin Mc- Carthy (Natural History Museum London) who allowed us to examine the type specimen of G. mjobergi and Kelvin Lim (Zo- ological Reference Collection, Singapore) who provided pho- tographs of the juvenile specimen deposited in his collection. REFERENCES ANANJEVA, N. B. & STUART, B. L. (2001): The agamid lizard Ptyctolaemus phuwuanensis Manthey & Nabhitabhata, 1991 from Thailand and Laos represents a new genus. Russian Jour- nal of Herpetology 8 (3): 165-170. BOULENGER, G. A. (1885) Catalogue of the Lizards in the British Museum (Nat. Hist.) I. Geckonidae, Eublepharidae, Uroplati- dae, Pygopodidae, Agamidae. London, 450 pp. Das, L (2006) A photographic guide to snakes and other rep- tiles of Borneo. New Holland Publishers, London, 144 pp. MALKMUS, R., MANTHEY, U., VOGEL, G., HOFFMANN, P. & J. Ko- SUCH (2002): Amphibians and Reptiles of Mount Kinabalu. A.R.G. Gantner, Ruggel, 424 pp. MANTHEY, U. & W. DENZER (1991): Die Echten Winkelkopfaga- men der Gattung Gonocephalus Kaup (Sauria: Agamidae) II. Allgemeine Angaben zur Biologie und Terraristik. Sauria 13 (2):19-22. MANTHEY U. & W. DENZER (2006): A revision of the Melane- sian-Australian angle head lizards of the genus Hypsilurus (Sauria: Agamidae: Amphibolurinae), with description of four new species and one new subspecies. Hamadryad 30 (1&2): 140. MANTHEY, U. & W. GROSSMANN (1997): Amphibien & Repti- lien Súdostasiens. Natur und Tierverlag, Münster, 512 pp. Moopy, S. M. (1980): Phylogenetic and historical biogeograph- ical relationship of the genera in the family Agamidae (Rep- tilia: Lacertilia). Ph.D. Thesis, University of Michigan, 373 p. SMITH, M. A. (1925): On a collection of reptiles and amphib- ians from Mt. Murud, Borneo. Sarawak Museum Journal 8: 5-14. SMITH, M. A. (1935): The Fauna of British Indıa, Ceylon and Burma, including the whole of the Indo-Chinese subregion. Reptilia and Amphibia. Vol. II. Sauria. Taylor & Francis, Lon- don, pp. 1-x111+1-440. WERMUTH, H. (1967): Liste der rezenten Amphibien und Rep- tilien: Agamidae. Das Tierreich vol. 86, Walter de Gruyter, Berlin, 86: pp. i-x1v+1-127. Bonner zoologische Beiträge | Band 56 | Heft 4 Seiten 259-272 Bonn, November 2009 Activity profiles, habitat selection and seasonality of body weight in a population of Arabian Spiny-tailed Lizards (Uromastyx aegyptia microlepis Blanford, 1875; Sauria: Agamidae) in Saudi Arabia Thomas M. WiLms!-*, Philipp WAGNER?, Mohammed SHOBRAK3 & Wolfgang BÖHME? ! Zoologischer Garten Frankfurt, Bernhard-Grzimek-Allee 1, D-60316 Frankfurt am Main, Germany; E-Mail: thomas.wilms@stadt-frankfurt.de. 2 Zoologisches Forschungsmuseum A. Koenig, Adenauerallee 160, D-53113 Bonn, Germany. 3 Biology Department, Science College, Taif University, P.O. Box 888, Taif, Saudi Arabia. 4 Corresponding author. Abstract. A field study was carried out on the Arabian Spiny-tailed Lizard (Uromastyx aegyptia microlepis) in Saudi Arabia (Mahazat as-Sayd Protected Area) focusing on seasonal differences in activity, body condition and on parame- ters influencing selection of burrow sites in this large desert-dwelling lizard. Uromastyx a. microlepis is highly season- al in respect to activity/visibility outside of its burrows which provide shelter against unfavorable climatic conditions. At Mahazat as-Sayd these lizards exhibit a bimodal activity in spring and summer while activity in autumn is unimodal with a peak at early afternoon, but the overall activity is generally low at this time of the year. More than 73 % of total ob- served yearly activity takes place in spring and early summer. Seasonal changes in availability of food result in differ- ences of the animals’ body condition, with a significant decrease of body-mass between spring / autumn and summer / autumn respectively. Selection of burrow sites is largely influenced by soil type and vegetation coverage. Key Words. Reptilia; Sauria; Agamidae; Uromastyx; Uromastyx aegyptia microlepis, seasonality; habitat selection; ac- tivity; body condition; Mahazat as-Sayd Protected Area; Saudi Arabia. INTRODUCTION The spiny-tailed lizards of the genus Uromastyx are in- habitants of the deserts and semi-deserts of North Africa, Arabia and the Middle East. Currently 15 species are con- sidered to be valid, of which 6 are known to occur on the Arabian Peninsula (WILMS et al. 2009). There is a con- siderable scientific interest regarding the taxonomy and phylogeny of these animals but ecological knowledge re- mains scarce and fragmentary with most studies focusing on Uromastyx nigriventris and Uromastyx aegyptia (GRENOT & LOIRAT 1973; GRENOT 1976; BOUSKILA 1986; VERNET et al. 1988; CUNNINGHAM 2000, 2001; AL-Haz- MI 2002; AL-JOHANY 2003; AL-HazM1 et al. 2005). This is especially interesting, considering the ecological impor- tance, prevalence and local abundance of these animals. Uromastyx aegyptia (Forsskál, 1775) ıs by far the largest member of the genus, reaching a maximum body length of even more than 700 mm and a weight up to 2.500 g (Fig. 1). Taxonomically, three subspecies, Uromastyx a. aegyptia (Forsskál, 1775), Uromastyx a. microlepis Blanford, 1875 and Uromastyx a. leptieni Wilms & Böhme, 2001 are distinguised (WILMS et al. 2009). The present study focuses on the Arabian Spiny-tailed Lizard (U. a. microlepis) which lives in deserts and semideserts of Arabia (Saudi Arabia, Yemen, Oman, United Arab Emi- rates, Qatar, Kuwait), in Jordan, Syria, Iraq and coastal Iran. Habitats show a marked seasonality regarding cli- matic parameters (temperature, air humidity, precipitation) as well as regarding availability of food. In contrast to the wide range of daily and annual fluctuations of air and soil temperatures, the temperatures within Uromastyx burrows are remarkably constant (WILMS & BÖHME 2007 & unpub- lished data). Therefore these burrows are considered to play an important role as refuges in respect to thermo- and hydroregulation and as shelter against predators. Burrows of Uromastyx aegyptia can be up to 1025 cm long and reach about 180 cm deep in the ground (BOUSKILA 1983, 1984), but are, at least at Mahazat as-Sayd, normally not longer than about 300-530 cm and 80-120 cm deep (own data). 260 Thomas M. WiLms et al.: Arabian Spiny-tailed Lizards in Saudi Arabia The primary production of Uromastyx habitats is quite low and exhibits strong seasonal and annual differences. There- fore it is somewhat surprising that Uromastyx aegyptia, as all other species of the genus, is primarily herbivorous (MANDAVILLE 1965, KEVORK & AL-UTHMAN 1972, AL- OGILY & HUSSAIN 1983, BOUSKILA 1984 & 1987, ROBIN- SON 1995). Nevertheless remains of beetles (Tenebrion- idae, Carabidae), ants, grasshoppers and even scorpions are also found in the stomach contents as well as in fecal pellets of all age classes (KEVORK & AL-UTHMAN 1972, WiLms 2007, pers. obs.). However, the proportion of animal matter in the food 1s very low and was estimated to be only about 1-2 % of total food intake. The present paper analyzes the seasonality of activity and body condition in this species which is induced by the aforementioned environmental variability. A second focus was put on the selection of habitat structures by Uromastvx a. microlepis. This study was carried out on free ranging populations of Uromastyx aegyptia microlepis at Mahaz- at as-Sayd Protected Area, Saudi Arabia. MATERIAL AND METHODS Study Site All fieldwork was carried out within the boundaries of Ma- hazat as-Sayd Protected Area, Saudi Arabia, which has been protected since 1989. Mahazat as-Sayd (22° 15’ N, 41° 50” E) is located in central-western Saudi Arabia, ap- proximately 170 kilometers north-east of Taif. The reserve is Saudi Arabia’s only completely fenced wildlife reserve and is a reintroduction site for Houbara Bustards (Chlamy- dotis undulata macqueenii), Arabian Oryx (Oryx leuco- ryx) and Sand Gazelles (Gazella subgutturosa). Mahazat as-Sayd covers an area of 2244 km? and lies within one of the hottest regions of the world (MEIGS 1953). It is a hot and semi-arid to arid desert steppe habitat, typical of the central plateau of the Arabian Peninsula. The eleva- tion is between 900-1050 m above sea level (LENAIN et al. 2004). The terrain of this area consisted of flat gravel plains, known as regs, occasionally interdigitated by dry sandy wadis (Fig. 2). Over 95 % of the area is covered by sand and gravel. Air temperature in Mahazat, as record- ed in a standard weather shelter, often exceed 43 °C dur- ing the summer, and occasionally reaches 50 °C (SEDDON 1996, SHOBRAK 1996, WILLIAMS et al. 1999). Rainfall av- erages ca. 100 mm per year (WILLIAMS et al. 1999) and typically occurs between March and May each year, but with occasional important rain events at other times. There is no permanent source of water above ground level in Ma- hazat as-Sayd, but ephemeral pools exist for short peri- ods after heavy rain (LENAIN et al. 2004). The main study site was situated just south-east of the main gate to Mahazat as-Sayd (Al-Muwayh gate), cover- ing an area of approx. 4 km? and was primarily used to assess habitat selection, while a second area nearby was chosen to establish a 60 km long transect for the evalua- tion of activity profiles. The flat gravel plains in this part of the Arabian Desert are intersected by wadis and dominated by sparse vege- Bonner zoologische Beiträge 56 261 Fig. 2. dis. Photo: T. WILMS. tation of perennial grasses, including Stipagrostis spec., Panicum turgidum and Lasiurus scindicus, and small trees, mainly Acacia spec. (MANDEVILLE 1990). This study took place in spring, summer and autumn 2006 as well as spring and summer 2007. Methods applied to assess activity profiles To assess the differences in seasonal activity of the Spiny- tailed Lizards a transect was defined within an Uromastyx colony. The length of this transect was 60 km and it was driven with a 4 WD vehicle eight times per season (equals 480 km each) with an average speed of 40 km/h. These transects were driven only on sunny and calm days. As a result of this protocol the covered time per season was 12 hours (7 am-7 pm). The transect was driven in the fol- lowing seasons: Summer 2006 (28th July-O4!h August), au- tumn 2006 (26th October-17!h November), spring 2007 (30 March—06 April) and early summer 2007 (27th June-O15t July). The following data were registered by two persons (each concentrating on the area on either side of the transect): Aerial view on the study site at Mahazat as-Sayd Protected Area showing reg formation with interdigitating sandy wa- Presence of specimens (time), size class [juveniles (total length < 200 mm), subadults (total length 200-400 mm), adults (total length > 400 mm)], distance of specimen from respective burrow, animal behaviour (e.g. basking, shad- ing, feeding), coloration (dark grey, bright yellow) and habitat (e.g. soil type, vegetation). To estimate the ther- mal environment of the observed specimens physical mod- els were used to measure operative temperatures (T.) in different microhabitats in summer 2006 and spring 2007 (HERTZ et al. 1993, SHINE & KEARNEY 2001). Hollow cop- per cylinders (230 mm long, 78 mm diameter, 0.6 mm wall thickness) were used as models. Miniature data loggers (i-buttons, Dallas Semiconductors, Model 1921) were placed in the model lumen to record T.. Data loggers were calibrated in a water bath against a mercury-in-glass ther- mometer that had a calibration traceable to the National Institutes of Standards and Technology. To mimic the re- flectance of a cold (grey colored) Uromastyx the model was covered with grey duct tape. Temperatures measured with the tape covered copper model followed the body temperatures of a dead Uromastyx better than temperatures measured with the uncovered model. Duct tape covered models were calibrated against dead as well as live Uro- mastyx under the original climate in the field (data will be published elsewhere). 262 Thomas M. WILMs et al.: Arabian Spiny-tailed Lizards in Saudi Arabia Fig. 3. Methods applied to assess habitat selection The main study site was in spring 2006 systematically scanned for Uromastyx burrows by driving parallel tran- sects (distances between transects ca. 50 meter). Each ac- tive burrow was marked with a flag (Fig. 3) and the fol- lowing parameters were registered: Orientation of burrow entrance (N, NE, E, SE, S, SW, W, NW), soil type (fine sand, coarse sand, clay sand, sand/gravel mixture, fine gravel, medium gravel, coarse gravel, rock/sand mixture, rock), slope, vegetation coverage (0 %, <5 %, < 15 %, < 25 %), distance to next Acacia. A total of 206 active Uro- mastyx burrows were registered. Burrows were considered as active (inhabited or visited) either if an Uromastyx was observed directly or if fresh tracks were found near the opened and clean burrow entrances. Otherwise, the bur- row was classified as non-active (abandoned). To assess habitat selection 80 burrows out of the 206 reg- istered ones were selected randomly and tested against 80 additional random localities within the boundaries of the study site, for which the same parameters as for the ac- tive burrows were collected. x2 analyses were performed to assess differences between burrow locations and ran- domly placed locations. Methods applied to assess seasonality of body weight At the main study site Uromastyx were trapped using snares put into the entrance of active burrows. Trapping Uromastyx burrow marked with a flag at the study site in Mahazat as-Sayd. Photo: T. WiLms. was carried out in spring 2006 (08th May-2nd June), sum- mer 2006 (15! July-O15t August), autumn 2006 (20% Oc- tober—11tt November.) and spring 2007 (26'h March-1 5th April). This trapping method has the advantage that a cap- tured animal is still able to retreat into the burrow and therefore the risk of overheating and loosing the animal is minimized. Captured Uromastyx were marked with a passive integrated transponder (PIT), weighted, measured (total length, tail length, head length and width) and sev- eral scale counts were taken. Some specimens were im- planted with temperature data loggers (i-buttons) and equipped with radio transmitters for a separate study on the thermobiology of Uromastyx a. microlepis (WILMS et al. in prep.). The specimens were subsequently released at their original burrow. To assess seasonality of body condition, a body-mass-in- dex (bmi) was calculated for each animal using the fol- lowing quotient: bmi = weight (g) / total length (cm). On- ly data of adult specimens (total length > 400 mm) were included in the analysis (n=62). Statistical analyses of ecological data The Excel 2000 and SPSS (15.0) statistical packages were used to run the analyses. x2 analyses have been selected to evaluate activity data as well as data on habitat selec- tion. To uncover differences in body-mass-indices Mann- Whitney U-test and Kruskal-Wallis statistical test were ap- plied. Bonner zoologische Beiträge 56 263 10 en | ir | Manabi ı a 3 3 3 g 2 3 3 3 3 3 3 3 3 3 2 8 5 8 3 8 2 3 2 3 = 51 5 $ $ > = = = 2 = 2 2 5 = 2 ? 2 2 = = 2 = 2 Mess sá ¢ $ 3 3 8 8 3 4 3 3 € 4 € 4 2 3 4 3 E AA Se = - o & a o = = o o o © eS E > > peep 2 15 — — | 10] : 1 I | | | .— MM i A | eee AA = SIM cs JS "aye ‘si 2 ¢ 8 @ 8 2 E 2 8 = 8 2 E E A rote Saks A'S. is ae Og IE OUR > $2 $8 8 3232 38 3 24282 3 8 8 8 3 8 «322 3 8 8 ole de EG a S o = = o a a ” = E > > ES = = Sa c. Dt 2 bl or 2. i [ 1 A E Dis =i | | wm Seca ote SUMS 081 US 18 8. 3.82 2. 8 38.8 3H 8 2% 8 3-2 ias 2 = 5 8 ¢ 2 @ $ ze es eee ee ge ia Eee Ss 8.808 8 £ £ 32 3 8 8 $8 CEM SPSS AI O EE IE d. ESA 15 10; 5 = SEN. m sans! aa 3 808 08 02-18) $08 4.8.2 83288283808 a E A ER ET A E E ip er Feuer Rm ee Se ea aa 2 28 28 3 8 Fig. 4. Visibility profiles of Uromastyx aegyptia microlepis at Mahazat as-Sayd at different seasons; a. Spring; b. Early Sum- mer; c. Summer; d. Autumn. RESULTS Activity structures A total of 245 Uromastyx were sighted while driving the transects. The absolute numbers for the respective seasons are: 97 (spring), 82 (early summer), 52 (summer), 14 (au- tumn). This is equivalent to 39.59 % (spring), 33.46 % (early summer), 21.22 % (summer) and 5.71 % (autumn) of total observed presence above ground. There is a very restricted activity during winter (deduced from own data on body temperatures, data not shown; BOUSKILA 1986, Ar-Hazmi et al. 2005). These observations show that Uro- mastyx a. microlepis is highly seasonal at Mahazat as-Sayd (y2 = 65.74, P<0.01, df = 3), with a maximum visibili- ty from spring to early summer. a. 100 Fig. 5. Distribution of different behavioral activities as abso- lute numbers (black bars) and as percentage of total observati- ons (grey bars) at different seasons; a. Spring; b. Early Sum- mer; ec. Summer; d. Autumn. Behavior: 1. Basking near burrow entrance; 2. Animal far away from burrow entrance; 3. Shading under Acacia tree; 4. Feeding; 5. Basking on a bush; 6. Basking on a stone. Profiles of daily visibility were produced for every sea- son by adding up the specimens observed within respec- tive intervals of 30 minutes (Fig. 4). The diagrams show, that U. a. microlepis exhibits a bimodal rhythm in spring and early summer with two relatively high visibility peaks in spring (at morning from 10:30-11:00 hrs respectively afternoon from 17:00-17:30 hrs) but only one pronounced peak in early summer (at morning from 8:30-9:00 hrs), followed by a greatly reduced but nevertheless clearly vis- ible peak at early afternoon (13:30-14:00 hrs). The pro- files for spring and early summer exhibit a period with a significantly reduced visibility during the hottest time of 264 Thomas M. WILMs et al.: Arabian Spiny-tailed Lizards in Saudi Arabia a. 120 100 + > mn un 80. 60 | 40 + A 4 ay f MATTE A A Be a ae BEER 3.328 82 2 82 828 32 8 2 8282 32 8 8 2 82 2 32 8 8 3 8 | 8 € 9 E SER Ge SE 8 8 8 8 §& ¢- 8 6 3 $ 3 $s $38 ¢ 6 ¢ 6 $$ ¢ &¢ 3 8 8 sR 82 8 22 8 MEI EEE EEE 4 2 8 72 $ 8 #2 28 1 18 4 Ge Ge fe Es b. 120) 100 | so. so 40. 20, In a 01m MA ea alla et — 2¢e2se2e8e822 8882282828888 238818 2333 23 2:3 68 @ 2 2358 82 2 2 22 s $222 265 95 23 FE. PSF FR SFG FF BE SF Sn Se A a i, Sa Wee Wee Be Bee ne @#e¢es tes &#@ # te = &# # &€ Ss FE Ss c. 120 100 80 60 40 20 | | al... a! - ee 22a 2882888888288 838383 8 [II EE a OR 88 8 8 8 3 ¢ 38 38 3 8 38 3 3 2 3 3 3 38 3 8 s g $ 88 8 8 @ 3 8 ¿ 3 3 ¿$ 2633383834 $ ¿2 £e¢ S$ © @ @ tt 2 8s Fe 2 e 120 | 100 80 | 60 40 20 ‘vs 282228 e822 2828888288888 S$ ¢¢ ¢ sees Skee ee ef 8 BS SEE SBS 8 $ $ 8 $ 8 2 3 3 2 3 8 3 8 3 2 2 23336653 EI ER BB @ e Fb | sg Fig. 6. Profiles showing the number of visible specimens (black bars) as well as average distances (in meters) of all ob- served specimens within each respective interval of 30 minutes (grey bars); a. Spring; b. Early Summer; ¢. Summer; d. Autumn. the day (12:30-14:00 hrs in spring; 12:00-13:30 hrs in ear- ly summer). The profile for the summer season shows principally also a bimodal visibility rhythm with the high- est visibility between 8:30 hrs and 9:00 hrs followed by a rapid decrease of visibility. After 12:00 hrs specimens were seen only occasionally (two specimens between 13:00-13:30 hrs, three specimens between 15:00-15:30 hrs, one specimen at 17:26 hrs and one specimen at 17:34 hrs); but nevertheless some specimens were still visible. The comparison of the profiles from spring, early sum- mer and summer show an obvious trend of reduction of afternoon visibility and a shift of the first visibility peak from late morning to early morning, which is possibly re- lated to the increase of maximum day temperature. The a. | 120 ; | I 100 2 Subadults & Adults Adults 60 | 0. nl 0. HH. ole een so DEP Peri ee 828388883 88283288 3888888288 8 S828 28955 FS 2 eur see WERE PEELE GLa Pid ee b. on 1100: | m Adults Subadults & Adults = | 40 + | „| n f] | 2 { ACA al a dl la Ht 2228282282282 88288 288288 8 MEE EEE EEE TEE EEE | c mo Pa | — |100 4 1 1100 7 mn | 2] | = | 40+ 7 20 1 COUNT PE o ES _ (“ERESS SES SR GRE REESE EER EES | 8222232382323 2232392 82 8 3 L_ _ _ | d. 120 100 - - A A 80 All size classes E PA = AA 40 = e EEE EEE SER 2 = o SS SS SS a SPS EEE Mr WE TE WET IT FE FR ERTETE S49 9 9 6 2.2 A Bu ee PSR eee eee EER ET EEE EEE Es Fig. 7. Profiles showing the number of visible specimens (black bars) as well as the average distances (in meters) from the burrows (grey bars). Vertical bars indicate size class of ac- tive specimens. a. Spring; b. Early Summer; ¢. Summer; d. Au- tumn. visibility profile for autumn is unimodal and shows an overall very restricted level of visibility with a maximum occurring between 12:00 hrs and 15:00 hrs, then the warmest time of the day. These visibility profiles were produced regardless of the type of observed activity, thus counting a visible speci- men as being active. Figure 5 shows the distribution of the different activities like basking, shading, feeding and specimens being far outside of their respective burrow for all four observation periods as absolute numbers (black bars) as well as percentages thereof (grey bars). Calcula- tion of x2 statistics showed that the observed seasonal dif- ferences are significant for the categories “basking near Bonner zoologische Beiträge 56 265 burrow entrance” (x2 = 57.451, P< 0.01, df=5) and “an- imal far outside of its burrow” (y2 = 24.000, P< 0.01, df = 5), while differences in the other four categories were not significant. As a consequence, diagrams showing the number of visible animals as well as the average distance of specimens from their burrows during each 30 minute interval were created (Fig. 6). Temperatures in spring are relatively mild, resulting in a prolonged time the lizards have to spend basking near the burrow entrance to reach their preferred body tempera- tures. After having reached the preferred body tempera- ture some animals start leaving the direct vicinity of their own burrow. From 12:30-14:00 hrs presence of specimens is greatly reduced, but those animals still present exhibit a high degree of activity indicated by the long distance from their respective burrow (Fig. 6a). Evaluating the ac- tivity data in respect to animal size revealed, that from 12:07 hrs until 14:41 hrs only large adults were seen far outside of their burrows (Fig. 7a). Obviously temperatures during midday are even in spring high enough to force the juveniles and subadults under ground, seeking shelter in their burrows as thermal refuges and therefore accounts for the relatively low number of active specimens. In ear- ly summer average as well as absolute temperatures are higher than during spring, thus the basking period near the burrow entrance on early morning is shorter; thus result- ing in an earlier start of “far from the burrow activity” (Fig. 6b). Obviously temperatures at morning are already too high to allow activity of juveniles and subadults away from their burrows because of the risk of overheating. From 8:28 hrs until 14:00 hrs only adults have been seen active. After 14:12 hrs also subadults started “far from the bur- row activity” (Fig. 7b). In summer visibility of Uromastyx was generally greatly reduced. There was no observed ac- tivity of juveniles and subadults and only five adult spec- imens have been seen active during the period between 10:12-11:40 hrs and at 13:08 hrs and 13:25 hrs (Fig. 6c & 7c). In autumn visibility and activity is greatly reduced due to the low temperatures. Activity was observed for all size classes only during early afternoon (Fig. 6d & 7d). Habitat selection Of the five registered parameters at burrow locations three proved to show significant deviations between observed and expected frequencies. These are: orientation of bur- row entrances (P < 0.01), soil type (P < 0.01), and vege- tation coverage (P < 0.01). The slope of the area around burrow entrances as well as the distances of burrow en- trances to next Acacia were not significantly different compared to circumstances at random locations (data not shown). Orientation of burrow entrances. ¥2 analysis revealed a significant deviation of the observed distribution of bur- row entrance directions from the expected distribution of burrow entrance directions (x2 = 30.27; P< 0.01; df =7). The highest deviation from the expected value exists for burrow entrances directed to south-west and south-east (Fig. 8). Selected soil type at burrow location: Nine different soil types were found at the 80 randomly selected locations within the study area (fine sand, coarse sand, clay sand, sand/gravel mixture, fine gravel, medium gravel, coarse gravel, rock/sand mixture, rock). x2 analysis of observed frequencies of selected soil types at burrow locations and of soil types at randomly placed locations within the study === 45 40 35 30 25 20 15 10 5 0 T T T T 1 N NE E SE S SW W NW E | Ml Observed frequency O Expected frequency Fig. 8. | Observed and expected frequencies of Uromastyx burrow entrance directions at Mahazat as-Sayd. 266 Thomas M. WiLms et al.: Arabian Spiny-tailed Lizards in Saudi Arabia barra Fig. 9. Selected soil types at Uromastyx burrow locations a. coarse sand; b. fine gravel; e. medium gravel. area revealed significant differences (x2 = 40.30; P<0.01; df = 8). All but one of the randomly selected Uromastyx burrows were located at places with medium gravel, fine gravel and coarse sand (Fig. 9). The only exceptional bur- row was found between black rocks. Occasionally the original soil type in the vicinity of Uromastyx burrows was covered by a thin layer of drifting sand, which did not in- terfere with the physical properties of the under laying soil. In such cases only the main substrate was considered for the statistical analysis. Sandy soil was therefore defined as loose deep sand. Fig. 10. Vegetation coverage estimated within a radius of five meters around entrances; a. 0%; b. < 5%; e. < 15%; d. < 25%. Bonner zoologische Beiträge 56 267 Selected vegetation coverage at burrow location: Vege- tation coverage was estimated within a radius of five me- ters around the burrow entrances respectively around the randomly selected locations (0%, < 5%, < 15%, < 25 %; Fig. 10). x2 analysis revealed a significant difference be- tween vegetation coverage frequency at burrow locations and at randomly placed locations (x2 = 49.40; P< 0.01; df = 3). The data show that Uromastyx avoid area with 0% coverage but show a preference for areas with < 5% vegetation coverage. For places with < 15 % and < 25 % vegetation coverage no differences between observed and expected frequencies were evident. Seasonality of body weight Morphometric data of 62 adult specimens (total length > 400 mm) caught in spring 2006, summer 2006, autumn 2006 and spring 2007 were used to calculate body-mass- indices (bmi). Bmi in spring 2006 was 1.1961-3.2414 (2.4348 +/ 0.6033), in summer 2006 it was 1.1057-3.1269 (2.2172 +/- 0.6146), in autumn 2006 it was 0.8454-2.2981 (1.4972+/-0.4257) and in spring 2007 it was 1.1872-2.8921 (1.9828 +/- 0.4901). Kruskal-Wallis sta- tistics were performed to test data for significances (Fig. 11), which revealed that bmi differs significantly in respect to season (P < 0.001; Fig. 11 a) but not regarding to the a.) Season: 1 = spring 2006; 2 = summer 2006; 3 = autumn 2006; 4 = spring 2007 season | Mean Rank Chi-Square df Asymp. Sig. a Kruskal Wallis Test b Grouping Variable: season Fig. 11. sex of the respective specimens (P < 0,189. Fig. 11 b). To check if between seasons bmi differences are statistical- ly significant, data for all four seasons were pair wise test- ed using Mann-Whitney U-test. These tests show, that dif- ferences between spring 2006 and autumn 2006 were high- ly significant (P < 0.001) as well as for summer 2006 and autumn 2006 (P < 0.004). All other pairs showed no sig- nificances. To further evaluate the relationship of bmi between spring 2006 and autumn 2006 respectively summer 2006 and au- tumn 2006 Mann-Whitney U-tests were performed on da- ta for males and females separately. As result, differences for males were significant between spring 2006 and au- tumn 2006 (P < 0.001) but not for females while between summer 2006 and autumn 2006 differences were signif- icant for females (P < 0.016) but not for males. After FOWLER et al. (1998) Mann-Whitney U-test may be used with very low sampling units, but in that case there must be no overlap of observations between the two samples to reject Hp. This precondition of the test was met by the dataset for females with values for summer being 1.81-2.74 and for autumn 1.08-1.43, while it was not met by the dataset for males with values for spring being 1.99-3.24 and for autumn 0.85-2.30. Therefore we reject H, only for the females’ dataset. b.) Sex: 1 = male; 2 = female sex N Mean Rank Chi-Square df Asymp. Sig. a Kruskal Wallis Test b Grouping Variable: sex Kruskal-Wallis test statistics testing body-mass-index data for significance regarding season (Fig. lla) and sex (Fig. 11b). 268 DISCUSSION Activity structures As has been shown in this study Uromastyx aegyptia mi- crolepis 1s highly seasonal, with this seasonality not be- ing restricted to behavioral means of avoidance of tem- porarily unsuitable microenvironments but also to changes in the physical constitution of the animals (changes of bmi). The evaluation of thermoregulation in this species is not the primary objective of the present paper and will be discussed elsewhere (WILMS et al. in prep.). Neverthe- less, the observed seasonality accounts to some degree to interactions between the thermal environment and the lizard and is therefore here briefly discussed. Differences in temporal patterns of activity, the use of space, and body temperature relationships are not inde- pendent. Many lizards behaviorally regulate body temper- atures by shuttling between sun and shade or hot and cold microenvironments to alter heat flux, by modifying pos- ture to alter surface areas exposed to heat sources or sinks, and by regulating activity times (HUEY 1974, PIANKA 1986). These are well accepted paradigms of reptile ecol- ogy and especially true for extreme habitats like those in- habited by the studied Uromastyx at Mahazat as-Sayd. The visibility profiles of Uromastyx at Mahazat as-Sayd show a bimodal structure in spring, early summer and to a lesser degree in summer combined with a general de- crease of visibility from spring to summer. Visibility struc- ture changes from a bimodal to an unimodal rhythm from summer to autumn reflecting changes in the availability of suitable thermal microhabitats. The visibility profile for autumn shows an overall very restricted level of visibili- ty with a maximum occurring between 12.00 hrs and 15.00 hrs, then the hottest time of the day. These results are in good accordance with the observations of AL-HAzZMI et al. (2005) made in central Saudi Arabia (Al-Gassim region), with two activity peaks in spring (April) and summer (July) (9.00-11.00 hrs and 15.00-17.00 hrs in spring, 7.00—10.00 hrs and 16.00-18.00 hrs in summer) and only a single activity peak in autumn (mid October) (11.00-13.00 hrs). In this study observa- Thomas M. WiLms et al.: Arabian Spiny-tailed Lizards in Saudi Arabia tions were also conducted in early spring (February) show- ing that the animals have only one daily activity period in this season (11.00-15.00 hrs). After BOUSKILA (1986) the period with the highest activity is from March to Sep- tember in a population of Uromastyx a. aegyptia in Israel, while from October to February only 70 % of the observed Uromastyx burrows showed signs of activity. The lowest levels of activity were recorded from December to Mid- February, were all observations of active specimens were made on clear and sunny days only. In another study conducted by AL-JOHANY (2003) near Riyadh (Thumama; King Khalid Wildlife Research Cen- ter) an unimodal activity structure was observed in spring (May) starting between 9.00 and 10.00 hrs and ending around one hour before sunset. Summer activity (August) was found to be bimodal with one activity period starting 8.00-9.00 hrs until midday and another period from 15.00-16.00 hrs until sunset (18.30 hrs). We suggest, that the different spring activity profiles established in the pres- ent study compared with the result of AL-JOHANY (2003) might be explained by the different methodology used to assess activity. While in the present study a transect method was applied AL-JOHANY (2003) observed the an- imals from a hiding place within the study area, which probably lead to biased results by overestimating activi- ty during midday by observing single actively roaming specimens. As has been shown in the present study, few specimens are visible above ground during midday but some of these show an extraordinary high activity by walking far from their respective burrow. All observed specimens being active during the hottest period of the day were adult. Adult Uromastyx a. microlepis have a body mass of up to 2.500 g and have therefore a higher ther- mal inertia than smaller conspecifics. This property is re- ducing the risk of overheating for large specimens and al- lows them a prolonged activity phase under unsuitable thermal conditions compared to smaller specimens. Operative temperatures (T.) for observation periods in spring 2007 and summer 2006 are available from three dif- ferent microhabitats (sandy plain, gravel plain and 40 cm deep in the entrance of an Uromastyx burrow). There is only a slight difference regarding T, (average of all days with observations) between spring and summer with al- Table 1. Operative temperatures of three different microhabitats in spring and summer in Mahazat as-Sayd (only temperatures between 07.00 hrs and 19.00 hrs were used for calculation). Te (min, spring) To (min, summer) To (max, spring) Sandy plain 27.8 °C 27.6 *C 54.4 °C Gravel plain 28.8 °C 28.4 °C 33.3 °C Burrow entrance 29.8 °C 32.41.7C 39.4 °C T average T average T e (max, summer) e (spring) e (summer) 303 E 45.3 °C 47.20€ 54.8 °C 44.6 °C 46.4 °C 36.1 “E 36.7 °C 37. 25E Bonner zoologische Beiträge 56 269 errrerrrerrerrrerrererrerrrererrer| Fig. 12. Profiles of T, in gravel plain in spring (solid line) and summer (dashed line). Adults (Ad.) Fig. 13. Estimation of Uromastyx Ty, based on average T.; a. Spring; b. Summer; — — — T, burrow entrance (approx. 40 cm deep); - - - -T, gravel plain; T. sandy plain most identical minimum values but slightly higher max- ima at summer (Tab. 1). The main observed differences lie in the temperatures of the Uromastyx burrow entrance and in the profiles of T. in spring and summer (for the pro- file of T, in the gravel plain see Fig. 12). During spring, temperatures in the entrance of the burrow differ between 7.00 hrs and 19.00 hrs about ten degrees centigrade while in summer this variation accounts only to three degrees. The profiles of mean Ty sravel plain IN Spring and summer are quite similar until 10.40 hrs. After that time T, rises faster in summer and reaches a higher maximum than in spring. Afternoon T, in summer remains significantly higher until sunset (see Fig. 12). Based on the combined visibility/activity profiles and the mean T. for the respective seasons, body temperatures (T,,) for the first specimens seen outside of their burrows are estimated to be around 30-31 *C in spring and 33 *C in summer (because all specimens observed on the transects have been found in a gravel or in a mixed gravel/sandy microhabitat we suppose that T,, is within the range of T. sandy plain and To gravel plain): 1» estimates for specimens ob- served active far from their burrows are 44-52 *C for subadults and 48-54 °C for adults in spring and 47-55 °C for adults in summer (only adult specimens have been ob- served during summer). These temperature ranges were de- duced using the actual measurements of T, for each ob- served specimen (Fig. 13 gives an approximated visuali- zation of these data!). These estimates are based on op- erative temperatures which reflect the interactions between biophysical and to some degree morphological factors in- fluencing T, of an ectotherm but lacking physiological (and behavioral) control (HERTZ etal. 1993) and therefore overestimate T,, at least in the upper temperature range. The experimentally established critical maximum temper- ature (CTyax) 18 51°C for this taxon (AL-JOHANY 2003) and temperatures of 55 °C are lethal within 4-5 hours (EL- GOHARY & ASHOUR 1975), therefore it is suggested that Uromastyx aegyptia microlepis is able to control T,, by physiological and behavioral control mechanisms at least at high temperatures (data on T;, control based on semi continuous T, measurements will be published elsewhere). It has to be pointed out, that preferred body temperatures change in respect to season, food composition and amount, hormonal status and other physiological traits and that therefore critical temperatures might also be subject to sea- sonal changes (PHILLIPS & HARLOW 1981, Woop 1989, SEEBACHER & FRANKLIN 2005, TRACY et al. 2005). Time windows during which operative temperatures in spring and summer would allow activity on a gravel plain far from the burrows were estimated for adult and subadult Uromastyx (Fig. 14) using the above listed T), estimates (spring data of subadults were used to estimate time win- dow during summer). The resulting diagrams show, that estimated time windows for subadults are considerably longer in spring and in summer compared to actually ob- served activity. In spring estimated time window is ca. four hours (ca. two hours at morning and two hours at after- noon), while observed activity is restricted to the morn- ing totaling to ca. two hours. In summer estimated time window is ca. 3.5 hrs during two periods, one late morn- ing and the other late afternoon, while observed activity was zero. For the adults, the estimated time window in spring is approximately 4.5 hrs and the period of observed activity is with ca. 4 hrs almost identical, while the esti- mated activity window for summer is about 6.5 hrs with the observed activity is only about two hours. The differ- ences between the estimated and observed activity phas- es may be explained by the costs associated with ther- moregulation, by other needs adversely influencing above ground activity like the necessity to maintain an ad- equate water balance and the seasonally different avail- 270 Thomas M. WiLms et al.: Arabian Spiny-tailed Lizards in Saudi Arabia a 160 . IM time window summer | ||] time window spring Spee Been 150 + u ER, as | y \ \ | | ra \ | ra \ | | id \ 40 + L | | ZL estimated estimated | / NJ ln | estimated estimated observed ese222222222222222228232 | RAR OwORaBRSSoE ETHAN AMO b 60 - - 2 1 time window summer | [] time window spring 50 u re! z = fo = xO \ y \ ” A NACO A estimated bs / MN, lso observed observed estimated 120 o observed, "esessssseesseesesesasesseessssessss| ARO IO OO EFE id Fig. 14. Estimated time windows during which operative tem- peratures in spring (solid line) and summer (dashed line) would allow activity on a gravel plaın far from the burrows for adult and subadult Uromastyx; a. for Subadults; b. for Adults. ability of food sources. We suggest that the high activity in adults during spring 1s associated with reproduction; in this case the benefits of successful reproduction may out- weigh the costs of thermoregulation. Habitat selection There is only one published study on habitat selection of Uromastyx aegyptia (BOUSKILA 1986). This study was car- ried out on the nominotypic subspecies (Uromastyx a. ae- gvptia) in southern Israel (near Hazeva Field Study Cen- ter, Arava) and focused on the following habitat parame- ters: habitat topography, soil type, distance to Acacia trees and orientation of burrow entrance. Selected soil type at burrow location: After BOUSKILA (1986) Uromastyx aegyptia significantly prefers reg soil and avoids dry water courses, loose saline soil and sandy reg. This result is in accordance with the present study were medium gravel, fine gravel and coarse sand on the surface was significantly preferred. All of these three men- tioned soil types can be subsumed as reg soil. Reg soil is characterized by a layer of closely packed stones over a layer of fine sediment several centimeters to meters thick (BREED et al. 1997). We suggest that this preference for reg soils is mainly because of the physical properties of this layer of fine sediments, providing a stable under- ground for the lizards to dig their burrows. Sandy soil [loose deep sand in old water courses (wadis) or sandy reg] might be disadvantageous to dig burrows in, because of its low stability and in the case of sandy reg because of the drifty sand being blown into the burrow entrance. Orientation of burrow entrances: BOUSKILA (1986) did not find any evidence for a preference of specific orientation of the burrow entrances, which was principally confirmed by AL-OciLY & Hussam (1983) for Uromastyx burrows in the Riyadh Region. However these authors report a slightly higher number of burrow entrances oriented to the northeast. In the present study x? analysis revealed a sig- nificant deviation of the observed distribution of burrow entrance directions from the expected distribution of bur- row entrance directions, with the highest deviation from the expected value exists for burrow entrances directed to south-west and south-east (Fig. 8). One reason for this ob- servation could be the prevailing wind direction, which is NE at 0-30° N. Unfortunately this suggestion can not be tested with actual data of wind directions in Mahazat as-Sayd because of a malfunctioning weather station dur- ing this study. Selected vegetation coverage at burrow location: Vege- tation coverage at burrow locations revealed to be signif- icantly different compared to vegetation coverage frequen- cy at randomly placed locations. Visibility/activity profiles of the present study are likely to explain this observed preference, because of the generally very high percentage of specimens seen in direct vicinity of their burrow en- trance. Having a food source directly available near the retreat entrance may lower ecological stress for the lizards due to the supposedly relative low costs associated with foraging. Based on these observations it seems obvious, that for Uromastyx a. microlepis at least some vegetation in the vicinity is essential for the selection of a burrow lo- cation. One main difference between the results of the present study and those of BOUSKILA (1986) is the failure to demonstrate a significant relationship between burrow location and distribution of Acacia trees. In his study BOUSKILA found a significant correlation in the distance between Uromastyx burrow respectively random locations and the nearest Acacia tree. His medians of the distances were 21 m (burrow — Acacia) respectively 28 m (random location — Acacia). In the present study both medians are considerably lower (14.5 m) and even the median of all 206 registered burrows is comparably low (14.0 m). These findings raise again the question of the importance of Aca- cia for Uromastyx in Mahazat as-Sayd. We take our data as a high availability of Acacia for the lizards in Mahaz- at as-Sayd which we consider as a strong factor minimiz- ing the need of an active selection of burrow locations in this particular respect. Nevertheless we regard Acacia trees as very important for the survival of Uromastyx provid- ing food and opportunities for thermoregulation (Fig. 15). Bonner zoologische Beitráge 56 271 Fig. 15. Adult Uromastyx thermoregulating under an Acacia at Mahazat as-Sayd. Photo: T. WiLms. Alternatively a second hypothesis explaining the “ob- served selection of locations” may be that Uromastyx is in fact a seed disperser and actively vegetates the area near the burrow entrance. The occurrence of seed dispersal through reptiles is well demonstrated (GODÍNEZ-ÁLVAREZ 2004, VALIDO & OLESEN 2007). Spiny-tailed lizards defe- cate regularly near their own burrow which can be proven by the abundances of fecal pellets, and these pellets con- tain regularly seeds of different plant species. In Mahaz- at as-Sayd they consist in summer to a high degree of undi- gested seeds of Acacia. AL-HAZMI (2002) evaluated the amount of seeds in the stomach of 40 Uromastyx a. mi- crolepis per season (spring, summer, autumn, winter) to- taling 6.7 % (spring), 40 % (summer) and 48.2 % (autumn) of total organic matter, thus representing a significant por- tion of the total food intake. In winter no food at all was found in the stomachs. Therefore a study testing the ef- fects of an intestinal passage on the germination of desert plant seeds and a possible impact of Uromastyx on the veg- etation coverage would be highly beneficial to understand the ecology of the species and this desert community. Seasonality of body weight Seasonal changes in body weight of an Uromastyx species (U. nigriventris; nomenclature after WILMS et al. 2009) had already been evaluated by GRENOT (1976) and VER- NET et al. (1988). In this species changes in body condi- tions are quite pronounced and associated with seasonal differences in food availability, which is also related to rainfall distribution. Extracellular and intracellular fluid volumes vary widely during the year (VERNET et al. 1988). This is also the case in U. a. microlepis with significant differences in body mass indices between seasons and with marked differences in extracellular water in the body cav- ity (observation made during implantation of data loggers). This difference in seasonal body weight is most likely as in Uromastyx nigriventris, also attributed to the season- ality of food availability. Acknowledgments. We thank H.H. Prince Bandar ibn Saud (NCWCD, Riyadh) for the support and continuous interest in this study in Saudi Arabia. Ahmed Boug (Director NWRC, Taif) as well as the staff of the NWRC and all Rangers at Mahazat as- Sayd helped in many ways to make this study possible and we thank all of them for their help, support and friendship. Uwe Wünstel, Dr. Nicola Lutzmann, Steven Arth and Marco Wage- mann worked as field assistants in this project; we thank them for their engagement and for valuable input. Dr. Ingrid Galal, Jeddah, and Gerda Kuhfittich, Jeddah provided support and gave two of us (TW and PW) a home base in Saudi Arabia. REFERENCES AL-HAZMI, M. A. (2002): Feeding Behavior and Food Selection of Dhab spiny-tailed Lizard Uromastvx microlepis from wild vegetation. Journal of the Egyptian German Society of Zool- ogy 37A: 185-203. AL-HAZMI, M. A., ASSAGGAFF, A. I. & A. A. 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Bonner zoologische Beiträge | Band 56 | Heft 4 | Seiten 273-278 Bonn, November 2009 Phylogeny of the genus Agama based on mitochondrial DNA sequence data Adam D. LEACHE!, Rebecca A. CHONG!, Theodore J. PAPENFUSS!, Philipp WAGNER3, Wolfgang BÓHMES, Andreas SCHMITZ#, Mark-Oliver RÓDELS, Matthew LEBRETON®, Ivan INEICH®, Laurent CHIRIO®, Aaron BAUER’, Edem A. ENIANGS, & Sherif BAHA EL DIN? ! Museum of Vertebrate Zoology, University of California, Berkeley, CA, USA 2 E-Mail: aleache@ucdavis.edu 3 Zoologisches Forschungsmuseum Alexander Koenig, Bonn, Germany 4 Museum histoire naturelle, Department of Herpetology and Ichthyology, Geneva 6, Switzerland > Museum für Naturkunde at the Humboldt University Berlin, Germany 6 Muséum national d’Histoire naturelle, Department Systématique et Evolution (Reptiles), Paris, France 7 Department of Biology, Villanova University, Villanova, PA, USA $ Department of Forestry and Wildlife, University of Uyo, Akwa Ibom State, Nigeria 2 3 Abdala El Katib St, Dokki, Cairo, Egypt Abstract. We present a preliminary phylogeny for 19 species of African 4gama lizards based on a maximum likelihood phylogenetic analysis of 1,181 bp of mitochondrial DNA sequence data. Monophyletic radiations of species in East, South and West Africa are supported, as well as a clade containing two species (4. doriae and A. sankaranica) distributed across the Sahel region. West African populations of A. agama are paraphyletic with respect to A. finchi from westernmost Kenya, providing further evidence for a biogeographic corridor between West and East Africa. Populations of A. agama form four phylogeographic groups, which suggests that 4. agama may be composed of multiple independent evolutionary lin- eages. Key words. Africa, phylogeography, Systematics, biogeography, Agama agama. INTRODUCTION African lizards in the genus Agama are among the most diverse and widespread terrestrial squamates in Africa, making them an ideal group for testing biogeographic hy- potheses and conducting comparative ecological and evo- lutionary studies. However, the robust phylogenetic framework for Agama that is needed to meet these goals is currently lacking. Constructing a phylogeny for Aga- ma requires detailed investigations of phylogenetic rela- tionships across multiple spatial and temporal scales. In general, the higher-level relationships within the Agami- dae are poorly understood (MAcEY et al. 2006), and dense sampling within Agama and other African and Asian taxa is necessary for testing the monophyly of and identifying the sister taxon to Agama. At the population level, sever- al widespread and polytypic species, such as Agama aga- ma in West Africa and A. lionotus in East Africa, have un- clear species limits (BOHME et al. 2005) and may be com- posed of multiple independent evolutionary lineages. These broadly distributed taxa pose some difficult species delimitation problems (WAGNER 2007; WAGNER et al. 2008a, WAGNER et al. 2008b) and require detailed phylo- geographic investigations. The goals of our collaborative research on Agama lizards are to combine our sampling efforts from separate biodi- versity fieldwork programs in different countries across Africa to 1) infer a comprehensive molecular phylogeny for the genus Agama, 2) resolve species limits in wide- spread, non-exclusive taxa, 3) conduct phylogeographic investigations of the 4. agama and A. lionotus complex- es, and 4) describe new species, where warranted. In this paper, we present a preliminary phylogeny for the genus Agama based on an analysis of mitochondrial DNA se- quence data (mtDNA). In addition, we investigate the phy- logeography of A. agama across West Africa. 274 Adam D. LEACHE et al.: Phylogeny of the genus Agama MATERIAL & METHODS A total of 19 species of Agama were included in the phy- logenetic analyses (Table 1). We included 15 specimens of A. agama from 10 countries across West Africa to in- vestigate the phylogeographic relationships among pop- ulations. We selected four species from the Agaminae to represent outgroup taxa, including Acanthocercus atricol- lis, Laudakia stellio, Trapelus mutabilis, and Xenagama taylori. We rooted our phylogeny with Laudakia stellio, although we note that the higher-level relationships with- in the Agaminae are poorly understood (MAcEY et al. 2000; 2006). DNA was extracted from tissues following the PureGene Animal Tissue DNA Isolation Protocol (Gentra Systems, Inc). Two portions of the mitochondrial genome were PCR amplified and sequenced, including a small fragment of the 16S rRNA gene (16S) and a portion of the ND4 pro- tein coding gene (ND4) and the adjacent histidine, serine, and leucine tRNA genes. Oligonucleotide primers used for PCR and sequencing are provided in Table 2. The 16S rRNA gene was PCR amplified for 30 cycles (95°C 30s, 58°C 30s, 72°C 50s) and the ND4 was amplified for 35 cycles (95°C 30s, 55°C 40s, 72°C 1 min). PCR products were purified using ExoSAP-IT (USB Corp.). Cycle se- quencing products were ethanol precipitated, and then se- quenced using an ABI 3730 automated sequencer. 1004 CM MCZ18456! 100}"GA ZFMK73185 gol [GH MVZ249617 NE MVZ238891 1001 GH LSUMZH20336 2% Er MLAMNH 109799 MR ZFMK76838 West Africa 98L. GH LSUMZH20085 Agama finchi Radiation TD 29061 GN ULM200 CM MCZ 184562 NG MVZ253099 EG CAS207958 100 | CM ZFMK8376 1 Agama sankaranica CM X3853 Agama planiceps Agama doriae ] Sahel Radiation Agama kaimosae Agama mwanzae East Africa Agama lionotus Radiation Agama rueppelli Agama caudospinosa 100 Agama aculeata 96 “ ” 93 Agama “agama 85 100 100 100 100 91 100 54 78 99 100 100 ul» 100 89 Laudakia stellio 0.1 substitutions per site Agama armata Agama hispida | South Africa Agama atra Radiation Agama anchietae Agama weidholzi Agama boulengeri West/ North Agama impalearis Africa Agama spinosa 100 Xenagama taylori Acanthocercus atricollis Trapelus mutabilis Fig. 1. Phylogenetic relationships within the genus Agama based on a maximum likelihood analysis of mtDNA sequence data using the GTR+T model of nucleotide substitution. Maximum likelihood bootstrap values > 50% are shown. Bonner zoologische Beitráge 56 27 un Table 1. Voucher numbers and locality data for specimens used in the study. All sequences are deposited in GenBank (Accessi- on Nos. GU128430 — GU128502). Voucher abbreviations are as follows: CAS, California Academy of Sciences; MVZ, Museum of Vertebrate Zoology; ZFMK, Zoologisches Forschungsinstitut und Museum; MCZ, Museum of Comparative Zoology; LSUMZ, Louisiana State University Museum of Natural Science; ULM, University of Louisiana at Monroe Museum of Natural History; AMNH, American Museum of Natural History; AMB, Aaron M. Bauer personal collection; specimen numbers beginning or en- ding with “X”, “T°, or “TR”, personal collector numbers for authors from the Museum national d Histoire naturelle. Species Agama aculeata Agama agama Agama agama Agama agama Agama agama Agama agama Agama agama Agama agama Agama agama Agama agama Agama agama Agama agama Agama agama Agama agama Agama agama Agama agama Agama anchietae Agama armata Agama atra Agama boulengeri Agama caudospinosa Agama doriae Agama finchi Agama hispida Agama impalearis Agama kaimosae Agama lionotus Agama mwanzae Agama planiceps Agama rueppelli Agama sankaranica Agama spinosa Agama weidholzi Acanthocercus atricollis Laudakia stellio Trapelus mutabilis Xenagama taylori Locality Botswana, Kgalagadi District Cameroon Cameroon, Mende, monts Takamanda Cameroon, Yaounde Cameroon, Yaounde Equatorial Guinea, Bioko Island, Malabo Gabon, Barrage de Tchimbele Ghana, Greater Accra Region, Tesano Ghana, Northern Region, Buipe Ghana, Volta Region Guinea, Diaragbela, Niger River Mali Mauritania, Selibabi Niger, Niamey Nigeria, Cross River National Park Tchad, Bol Namibia, Khorixas District Tanzania South Africa, Northern Cape Province Mauritania, Nouakchott Dist. Kenya, Naru Moru Cameroon, Daré Ville Kenya, Malaba South Africa, Northern Cape Province Mauritania, Nouakchott District Kenya, Nandı Kenya, Nakuru Kenya, Mara Namibia, Outjo Dist., Kamanjab Somalia, Awdal Region Ghana, Volta Region Djibouti, Dikhil Senegal Uganda, Rukungiri District Turkey, Antalya Province Egypt Somalia, Galbed Region Voucher MVZ 198076 ZFMK 83761 X 3853 MCZ 184561 MCZ 184562 CAS 207958 ZFMK 73185 LSUMZ H20336 LSUMZ H20085 MVZ 249617 ULM 200 AMNH 109799 ZFMK 76838 MVZ 238891 MVZ 253099 2906 I AMB 7582 ZFMK 84990 CAS 193435 MVZ 235764 ZFMK 83662 4218 X ZFMK 83653 AMB 4800 MVZ 235766 ZFMK 83660 ZFMK 83646 ZFMK 82076 AMB 7638 MVZ 241340 MVZ 249656 MVZ 236458 TR 481 CAS 201726 MVZ 230213 ZFMK 64395 MVZ 241356 Contiguous DNA sequences were aligned and edited us- ing Sequencher v4.8, and multiple sequence alignments were generated using Muscle v3.6 (EDGAR 2004). The open reading frame for the ND4 gene was identified us- ing MacClade v4.08 (MADDISON & MADDISON 2005). The 16S rRNA alignment included indel-rich loop regions that could not be aligned unambiguously and were excluded from the phylogenetic analysis. Phylogenetic relationships were inferred using maximum likelihood using RAxML v7.0.4 (STAMATAKIS 2008), and implemented the GTR+T model of nucleotide substitution. Support values for in- ferred relationships were estimated from 100 non-paramet- ric bootstrap replicates. RESULTS The maximum likelihood analysis of the combined mtD- NA sequence data (1,181 characters) supports the mono- phyly of the genus Agama (bootstrap = 100%; Fig. 1). The outgroup taxa Xenagama taylori and Acanthocercus atri- collis form a strongly supported clade (bootstrap = 100%) that is sister to Trapelus mutabilis. Within Agama, a clade containing A. impalearis and A. spinosa is sister to all re- maining Agama, although this relationship is not accom- panied by strong support (bootstrap <50%; Fig. 1). Some regional species assemblages are monophyletic, including a clade of Southern African species (A. aculeata, A. an- 276 Cameroon * Cameroon A Equitorial Guinea Nigeria 10? 0° 10° Adam D. LEACHE et al.: Phylogeny of the genus Agama A. planiceps 20° 30° Fig. 2. Phylogeographic structure of Agama agama across sub-Saharan Africa. Samples of Agama agama are labeled by coun- try in the unrooted phylogeny. The maximum likelihood phylogenetic analysis places Agama planiceps sister to Agama agama. chietae, A. atra, A. armata and A. hispida) and a clade of East African species (4. kaimosae, A. mwanzae, A. liono- tus, A. caudospinosa and A. rueppelli), each of which are strongly supported (bootstrap = 99% and 100%, respec- tively; Fig. 1). Strong support is also provided for a clade of species distributed across the Sahel region of sub-Sa- haran Africa, including A. doriae and A. sankaranica (bootstrap = 100%). Agama planiceps is sister to a clade containing all 15 specimens of A. agama and A. finchi, al- though this relationship is weak (bootstrap < 50%; Fig. 1). The interrelationships among these major clades are not supported by bootstrap values > 50%, except for a strongly supported clade (bootstrap = 100%) containing the Sahel species (A. doriae and A. sankaranica) and A. planiceps, A. agama, and A. finchi (Fig. 1). Populations of Agama agama in West Africa are para- phyletic with respect to 4. finchi from westernmost Kenya (Fig. 1). Populations of A. agama form four phylogeo- graphic groups (Fig. 2). The geographic proximity be- tween samples is a poor predictor of phylogenetic rela- tionships, and three of the four clades overlap spatially (Fig. 2). One of the most well sampled phylogeographic groups within 4. agama extends across West Africa from Mauritania to Gabon (Fig. 2). Two other clades with broad distributions include a group extending from Guinea to Cameroon and another extending from Tchad to Kenya (A. finchi) (Fig. 2). Finally, a clade with a more restricted dis- tribution includes populations from southeastern Nigeria, western Cameroon, and Bioko Island (Equatorial Guinea) (Fig. 2). DISCUSSION This study represents the first detailed molecular phylo- genetic investigation of the genus Agama. Monophyletic radiations of Agama lizards occur in East Africa, South Bonner zoologische Beitráge 56 IT Table 2. Primers used to amplify and sequence mitochondrial DNA. Gene Primers Reference 16S 16Sf: CGCCTGTTTAACAAAAACAT This study 16sR: CCGGTCTGAACTCAGATCACGT ND4 ND4:CACCTATGACTACCAAAAGCTCATGTAGAAGC AREVALO et al. (1994) LEU: ACCACGTTTAGGTTCATTTTCATTAC Africa, the Sahel region, and even West Africa (Fig. 1), but reconstructing the biogeographic history of the group is impeded by several factors. First, the inter-relationships among the major lineages of Agama inferred by the mtD- NA data are not accompanied by strong support, which would make any biogeographic scenarios based on the cur- rent phylogeny speculative. Second, our taxonomic cov- erage is not comprehensive and lacks approximately 10 species that are distributed throughout Africa. Finally, the sister taxon of Agama remains unclear. Despite these chal- lenges, there are strong biogeographic signals in the cur- rent phylogeny (Fig. 1). For instance, the phylogenetic placement of 4. finchi from western Kenya within the West African A. agama complex provides evidence for a biogeographic corridor between West and East Africa, a result that provides further evidence for a close biogeo- graphic affinity between these regions (WAGNER et al. 2008c). In addition, the close relationship between A. plan- iceps from Namibia and the West African A. agama com- plex provides support for a biogeographic connection be- tween West and South Africa (Figs 1 and 2). This relation- ship is not surprising given the close similarity in mor- phology and colour pattern shared between these species (BÓHME et al. 2005). The intraspecific phylogeny for Agama agama highlights the taxonomic problems that this widespread taxon intro- duces to Agama lizard systematics. This species 1s para- phyletic with respect to 4. finchi (Figs 1, 2) and appears to be composed of multiple distinct clades. The argument could be made that 4. finchi should be synonymized with A. agama to retain a monophyletic 4. agama, but A. finchi is clearly distinct based on morphology and coloration. The major phylogeographic groups found within 4. aga- ma based on mtDNA may represent distinct species, and represent a good starting point for testing species bound- aries with multiple nuclear markers. Determining if gene flow is absent among these mtDNA groups is an impor- tant next step in resolving the systematics of the 4. aga- ma complex. The paraphyly of 4. agama also underscores the need for additional detailed comparative morpholog- ical investigations that do not rely solely on adult male coloration characteristics (see GRANDISON 1968). The phylogeographic relationships within the Agama aga- ma complex exhibit a surprising amount of spatial over- lap between clades. Phylogeographic studies of other Aga- ma lizards, including A. impalearis (BROWN et al. 2002) and 4. atra (MATTHEE € FLEMMING 2002) recovered more typical phylogeographic patterns whereby populations formed geographically exclusive clades. The spatial overlap of mtDNA clades in the A. agama complex could be evidence of the presence of multiple distinct species, which co-occur throughout West Africa and are as of yet morphologically cryptic. Conversely, this spatial pattern could reflect the recent expansion of distinct lineages that were formerly restricted to exclusive geographic areas. Distinguishing among these different hypotheses of pop- ulation history await the addition of more specimens from throughout West Africa and the collection of nuclear DNA data. This work is currently underway. The collaborative nature of this research project is pro- viding an opportunity to conduct detailed investigations of the phylogeny and phylogeography of Agama lizards than would have otherwise been impossible. Our current work benefits from having expanded taxonomic sampling and multiple individuals of each species, more detailed outgroup sampling, dense population sampling in 4. aga- ma and A. lionotus species, and the addition of nuclear DNA sequence data. Acknowledgements. We thank reviewers for useful comments on previous versions of this manuscript, and to curators at the MVZ, CAS, MCZ, FMNH, AMNH, AMCC, and LSUMNS for tissue loans. REFERENCES ARÉVALO, E., Davis, S. K. & J. W. SITES, JR. (1994): Mitochon- drial DNA sequence divergence and phylogenetic relationships among eight chromosome races of the Sceloporus grammicus complex (Phrynosomatidae) in central Mexico. Systematic Bi- ology 43: 387-418. BÖHME, W., WAGNER, P., MALONZA, P., LOTTERS, S. & J. KÖH- LER (2005): A new species of the Agama agama group (Squa- mata: Agamidae) from western Kenya, East Africa, with com- ments on Agama lionotus Boulenger, 1896. Russian Journal of Herpetology 12: 143-150. 278 Adam D. LEACHE et al.: Phylogeny of the genus Agama Brown, R. P., SUÁREZ, N. M. & J. PESTANO (2002) The Atlas mountains as a biogeographic divide in North-West Africa: ev- idence from mtDNA evolution in the Agamid lizard 4gama impalearis. Molecular Phylogenetics and Evolution 24: 324-332. EDGAR, R. C. (2004): MUSCLE: multiple sequence alignment with high accuracy and high throughput. Nucleic Acids Re- search 32: 1792-1797. GRANDISON, A. G. C. (1968): Nigerian lizards of the genus Aga- ma (Sauria: Agamidae). Bulletin of the British Museum (Nat- ural History) Zoology 17: 65-90. Macey, J. R., SCHULTE IL J. A., LARSON, A., ANANJEVA, N. B., WANG, Y., PETHIYAGODA, R., RASTEGAR-POUYANI, N. & T. J. PAPENFUSS (2000): Evaluating trans-Tethys migration: an ex- ample using acrodont lizard phylogenetics. Systematic Biol- ogy 49: 233-256. Macey, J. R., SCHULTE Il, J. A., FONG, J. A., Das, I. & T. J. Pa- PENFUSS (2006): The complete mitochondrial genome of an agamid lizard from the Afro-Asian subfamily Agaminae and the phylogenetic position of Bufoniceps and Xenagama. Mol- ecular Phylogenetics and Evolution 39: 881-886. MADDISON, W. P. & D. R. MADDISON (2005): MacClade 4: Analy- sis of Phylogeny and Character Evolution. Sinauer Associates, Sunderland, MA. MATTHEE, C. A. & A. F. FLEMMING (2002): Population fragmen- tation in the southern rock agama, Agama atra: more evidence for vicariance in Southern Africa. Molecular Ecology 11: 465-471. STAMATAKIS, A, HOOVER, P. & J. ROUGEMONT (2008): A rapid bootstrap algorithm for the RAxML web-servers. Systemat- ic Biology 57: 758-771. WAGNER, P. (2007): Studies in African Agama I. On the taxo- nomic status of Agama lionotus usambarae Barbour & Loveridge, 1928. Herpetozoa 20: 69-73. WAGNER, P., KRAUSE, P. & W. BÖHME (2008a): Studies on Afri- can Agama III. Resurrection of Agama agama turuensis Lo- veridge, 1932 (Squamata: Agamidae) from synonymy and ele- vation to species rank. Salamandra 44: 35—42. WAGNER, P., BURMANN, A. & W. BOHME (2008b): Studies on African Agama 1. Resurrection of Agama agama kaimosae LOVERIDGE, 1935 (Squamata: Agamidae) from synonymy and its elevation to species rank. Russian Journal of Herpetology 15: 1-7. WAGNER, P., KOHLER, J., SCHMITZ, A. & W. BOHME (2008c): The biogeographical assignment of a west Kenyan rain forest rem- nant: further evidence from analysis of its reptile fauna. Jour- nal of Biogeography 35: 1349-1361. Band 56 Bonner zoologische Beiträge Heft 4 Seiten 279-284 Bonn, November 2009 A new species of Bronchocela (Squamata: Agamidae) from Nicobar Island Jakob HALLERMANN Biozentrum Grindel & Zoologisches Museum, Martin-Luther-King-Platz 3, D-20146 Hamburg, Germany; E-Mail: hallermann@uni-hamburg.de. Abstract. A new species of the genus Bronchocela from central part of Nicobar Islands is described. It is similar to 2. cristatella but differs by having a red gular patch in males, fewer scale rows around midbody and only up to 3 dorsal body scales rows pointing upwards. It was found on seven islands of the central part of Nicobar but not in the northern or southern part of the archipelago. A colonization scenario of the Nicobar Islands by the genus Bronchocela ıs discussed. An arrival of an ancestor from Sumatra through the Great Channel is more probable than from north through the Ten De- gree Channel by island hopping. Key words. Squamata, Agamidae, Draconinae, Bronchocela, new species, India, Nicobar archipelago. INTRODUCTION The genus Bronchocela comprises nine species distributed in South East Asia, the Indo- Australian Archipelago, the Philippines and in western New Guinea (WERMUTH 1967, DE Rooty 1915, DIONG & Lim 1998, HALLERMANN 2005). Recently a taxonomic review of the genus was published including biogeographical data was given for each species (HALLERMANN 2005). The Andaman and Nicobar Archipelagos are politically part of India. The herpetofauna of the Andaman Islands 1s considered to be of Chinese-Indian affinities, resulting from the connection of these islands to the mainland dur- ing late Pleistocene glaciations with sea level lowering of about 100 m. On the other hand, Nicobar Islands are con- sidered to be of volcanic origin, with Indo-Malayan affini- ties (DAs 1999). Its fauna established mainly through waif dispersal across the Great Channel from Sumatra. For the size Of land area the islands of central Nicobar show the highest proportion of endemic snake species (Das 1999). Das (1999) summarized the history of herpetological ex- ploration from its beginning in the middle of 19th centu- ry to the late 19901es. The inventory of the Nicobar her- petofauna was relatively poor until the 1960ies. An expe- dition of the Zoological Survey of India in the 1960ies, however, increased our knowledge of the herpetofauna of the Nicobar Islands. Tiwary & Biswas (1973) described Bronchocela danieli from Great Nicobar. Indraneil Das, during several field trips between 1994 and 1998 discov- ered new species and country records and clarified prob- lematic species records e.g. Pseudocalotes archiducissae Fitzinger, 1861, a nomen nudum, as Bronchocela cristatel- la. Other species such as Calotes calotes reported by BLYTH (1863) as C. ophiomachus and by SMITH (1935) as well as Calotes jubatus could not be verified to exist on Nicobar, since no voucher specimens exist (Das 1999, 2000). Das (1999) presented the currently most complete list of amphibians and reptiles from the Nicobar Islands. Bronchocela cristatella ıs known to occur on several is- lands in the north and south of Nicobar (BISWAS & SANYAL 1980). B. danieli ıs restricted to Great Nicobar (see Fig. 1). S. P. Vijayakumar studied the herpetofauna of Nico- bar Islands in his PhD Thesis. On several islands of the central Nicobar a lizard of the genus Bronchocela occurs which differs morphologically from the common B. cristatella and all other species of this genus. It was found on Katchal Island, Trincat Island, Camorta Island, Nan- cowry Island, Bompoka Island, Tarassa (Teressa) Island and Chowra Island. It is more similar morphologically to B. cristatella than to B. danieli, but differs from it in several characters. A comparison with all other species of the genus shows that it represents a new species which I describe below. 280 Jakob HALLERMANN: A new species of Bronchocela from Nicobar Island E \ Car Nicobar 2 Chowra, } Tillanchong Tarassa Go Camorta Y Trinkat Nanco' Katchall ane y Little Nicobar 1 att & $ lora Nicobar j L he o 0 10 20 Kilometers Les | Fig. 1. Map of Nicobar Islands. Black coloured islands illustra- te occurrence of B. rubrigularis sp. nov. MATERIAL AND METHODS The specimens examined for this study are listed in the appendix. Museum acronyms follow LEVITON et al. (1985). The following data were recorded: Snout-vent length (SVL), tail length (TL from cloaca to tip of tail, if tail was complete), head length (HL, between tip of snout and hind border of tympanum), head width (HW, at an- gle of jaw), number of supralabials (SL, on each side) and infralabials (IL, on each side), number of scale rows around midbody (M), hind limb length (HLL, from groin to tip of fourth toe, without claw), foot length (FL, with- out claw), as well as the ratios TL/SVL, HLL/SVL, HW/HL. In several specimens diameter of the tympanum and orbit, and the ratio of the two as well as the number of scales between the nasal scale and anterior border of orbit along the canthus rostralis, were also checked. Bronchocela rubrigularis sp. nov. Holotype. ZMH R09271 (Fieldnumber Tri 25) (see Fig. 2). An adult male collected by S.P. Vijayakumar on Trinkat Island, central Nicobar Islands, Nicobar in 2004. Paratype. ZMH R09272 (Fieldnumber Kat 43). A subadult male, (Katchall Island), central Nicobar Islands, same collector as holotype. Six paratypes will be inven- toried at ZSI: Fieldnumber BOM 21: Bomboka Island: Fieldnumber NAN 15: Nancowry Island; Fieldnumber CAM 76: Camorta Island; Fieldnumber TER 39: Teressa (Tarassa) Island): Fieldnumber CHO 05: Chowra Island: Fieldnumber BOM 27: Bomboka Island. Diagnosis. A relative robust species with a SVL of 81-106 mm, and a long tail (280-377 mm (336-378 % of SVL). Tympanum large, more than half diameter of orbit, often dusky coloured, Ratio tympanum /orbit 0.74-0.89. Head covered above with small keeled scales, up to two slight- ly enlarged scales between orbit and tympanum. 7—10 supralabials and infralabials on each side. Canthus rostralis sharp, two small erect, compressed scales behind supra- ciliary edge. Nuchal crest formed by 7—10 lanceolate erect scales, bordered laterally by smaller erect scales, longest little longer than diameter of orbit. Dorsal crest smaller than nuchal crest, continues nuchal crest by a small gap. Mental wider than high, three postmentals, gular sac small in males, gular region covered with little enlarged keeled scales, smaller than ventrals. Body scales mucronate, keeled, homogenous in 52-58 midbody scales. 1-3 uppermost scale rows next to dorsal crest pointing upward, 4-5 rows directed parallel, others scale rows directed downwards. Ventrals about two times larger than dorsals, strongly keeled. It can be distinguished from other congeners of the Nicobar Islands: from B. cristatella by a larger nuchal crest, only 1-4 upper scale rows pointing upwards (versus 5—10), and a red gular patch (white in alcohol) in males; from B. danieli by a larg- er tympanum (versus only half diameter of orbit) , longer fifth toe than fourth finger (versus fifth toe smaller than fourth finger) and only two times larger ventrals (versus five times larger) Colouration. In life: Known for males only: body green to brown above, head light green to light brown, upper side of head sometimes mottled with red, tympanum black, or- bit with a blue ring around eye, in some males body same colour as head (light brown), ventral side light to yellow- ish green with a V-shaped red patch on gular (Figs 3-6). In alcohol: Males are dark brown to nearly black above, lighter brown on venter and on lower side of legs. A no- tably white, triangular patch (apex directed rostrally) on the gular region Description of holotype (ZMH R09271, Fig. 2). Snout vent length 106.4 mm, tail length 377 mm, 355 % of SVL (original tail broken and separate), hind limb length from base of femur to tip of 4" toe (without claw) 88 mm, foot length to tip of fourth toe: 34 mm, length of toes (with- out claws) 1-5 in mm: 5.7 / 9.4/ 18.2 / 21.7 / 14.9, length of fingers 1-5 in mm: 3.8 / 7.3 / 13.2 / 13.0 / 7.7, head Bonner zoologische Beitráge 56 281 qe ñ 4 IN > a 2, Der) Fig. 2. Preserved holotype of Bronchocela rubrigularis sp. nov. (ZMH 9271). A= complete holotype in dorsal; B= complete ho- lotype in ventral; C= head in ventral view; D= head in lateral view. Photo: J. Hallermann. 282 Jakob HALLERMANN: A new species of Bronchocela from Nicobar Island Fig. 3. Bronchocela rubrigularis male in life from Camorta Is- land. Photo: S.P. Vijayakumar. width in temporal region 16.0 mm; head length from tip of snout to hind margin of tympanum: 30.4 mm. 9 upper labials and 8 lower labials on each side. Diameter of or- bit 4.84 mm, diameter of tympanum 4.34 (ratio: tympa- num/orbit 0.89). Head concave, covered above with small uniform keeled scales, two slightly enlarged scales inter- mixed on an upper line between orbit and tympanum 5 scales between nasal and anterior border of orbit along canthus rostralis. Mental wider than high, followed by three postmentals. Gular region covered by keeled scales, gular sac small, covered with little enlarged keeled scales, smaller than ventrals. A skin fold stretches from angle of mouth to insertion of foreleg. Nuchal crest formed by 10 lanceolate erect scales, bordered laterally by smaller erect scales, longest little longer than diameter of orbit. Mid- dorsal scale row forming a small dorsal crest by triangu- lar erect scales, which ıs lower than nuchal crest, sepa- Fig. 4. Bronchocela rubrigularis same specimen as in Fig. 5, showing red gular patch. Photo: S.P. Vijayakumar. rate from from ıt by a small gap. 54 scale rows around midbody, dorsal and lateral body scales small, mucronate, keeled; 2 uppermost scale rows directed upwards, next 5 below parallel to middorsal scale row, other dorsoventral scale rows directed downwards. Ventrals about two times larger than dorsals, strongly keeled. Limbs relatively strong, covered by keeled scales. Coloration. Coloration in life not known for holo- and paratype, but supposedly green body colouration. In al- cohol (Fig. 2): upper side of body and head dark brown to almost black. A white gular patch, V-shaped, apex di- Table 1. Measurements and scale counts of Bronchocela rubrigularis sp. nov. specimen no SVL PI sex. M ear orbit SL/IL (% of SVL) HW HW/HL HLL FL HLL/SVL % __ remarks ZMH R09271 106.4 377 (354) m 54 434 484 99/88 30.4 6.0 0:52 88 34 82,7 Tail broken holotype ZMH R09272 80.3 304 (378) m 58 351 439 99/89 3 3.0 0.55 68.6 28 85.4 paratype sad BOM 21 81.9 -lost m 55 3,34 99/98 1.9 75.6 92.3 Nuchal crest 9 NAN 15 83.4 283 (339) 1 52 5.1 87/97 2.8 71.8 86.0 Eggs no gular patch CAM 76 85.4 -lost m 55 3.88 9 8/10 10 12.8 77.9 91.2 No patch ? TER 39 85.3 287 (336) m 56 2.8 9 10/9 10 12.7 78.2 91.6 Red patch CHO 05 86.5 205 (reg.) f 52 3.46 9 9/9 10 3.0 76.2 88.0 Pale gular patch BOM 27 85.4 290 (339) 1 58 3.82 10 10/10 10 2.7 80.0 93.6 Eggs no gular patch Snout-vent length (SVL), tail length (TL from cloaca to tip of tail, if tail was complete), head length (HL, between tip of snout and hind border of tympa- num), head width (HW, at angle of jaw), number of supralabials (SL, on each side) and infralabials (IL, on each side), number of scales around midbody (M), hind limb length (HLL, from groin to tip of fourth toe, without claw), foot length (FL, without claw), as well as the ratios TL/SVL, HLL/SVL, HW/HL. In several specimens diameter of the tympanum and orbit. Bonner zoologische Beiträge 56 283 Fig. 5. Bronchocela rubrigularis male in life, from unknown locality in central Nicobar in green morph. Photo: S.P. Vijaya- kumar. rected rostally. Tympanum dark, lower side of body be- tween fore legs, underside of upper and lower legs and re- gion around cloacae light brown, lower side of hands and foots whitish. Venter and lower side of tail brown. Variations (see table 1). Females differ from the holotype by a lower nuchal crest, and by the lack of a red gular patch. One female has a pale gular patch and one subadult male has no gular patch. Tail length varies (280-377 mm [336-378 % of SVL]). Body colouration varies (see di- agnosis). Body colouration (in alcohol) of venter and ven- tral side of legs and tail of paratype (ZMH R09272), subadult male, is very similar to the holotype. Scalation (see table 1) is similar. The nuchal crest is lower, no en- larges scales are present between orbit and tympanum. Fig. 6. Bronchocela rubrigularis male in life, from Trincat Is- land in brown morph. Photo: S.P. Vijayakumar. Distribution. The new species is only known from the seven island of Central Nicobar (Fig. 1), and its distribu- tion seems to be restricted to Central Nicobar. Etymology. The species is named after its red gular (lat- in rubber = red, gula (feminin) = gular) DISCUSSION Bronchocela rubrigularis sp. nov. was observed and col- lected on several islands of Central Nicobar (Katchal Is- land, Trincat Island, Camorta Island, Nancowry Island, Bompoka Island, Tarassa Island and Chowra Island) while it is missing on the northern group (Carl island ) and south- ern group of the Nicobar (Little and Great Nicobar). Pat- tern of distribution suggests that the new species is restrict- ed to the central region of Nicobar. While B. cristatella occurs in the northern part, B. danieli was only recorded on Great Nicobar. From a biogeographical viewpoint it is important to know about geological history of Andaman and Nicobar Archipelago. Andaman and Nicobar are both parts of the Sunda Land and not of the Indian Subconti- nent (DAs 1999). During glacial sea-level lowering in the Pleistocene (about 120.000 years ago) a land connection between Birma (Myanmar) and Andaman Archipelago ex- isted. On the other hand, the Nicobar Islands were isolat- ed during all times by the Great Channel from Sumatra in the southeast and by the Ten Degree Channel in the north. It is more plausible that a faunal colonization has occurred by dispersal through the Great Channel from Sundaland than by island hopping from north via An- daman since Bronchocela is missing on Andamar Island and Myanmar (DAs 1999). I can not yet estimate the time of arrival of a Bronchocela species on Nicobar nor can I resolve the genetic relationships as long as molecular da- ta are still lacking Acknowledgement. For loan of specimens we thank, Ivan In- eich (MNHN), I am very grateful to Wolfgang Bóhme (ZFMK) for reading and providing helpful comments on an earlier ver- sion of the manuscript, and Indraeil Das (Kota Samarahan, Malaysia) for providing a map of Nicobar Islands. S. P. Vijayaku- mar provided me two specimens for description, and some in- complete measurement data of six specimens of the same species but no catalog numbers. Also, data on colouration of these spec- imens and habitat notes are still lacking. REFERENCES Biswas, S. & D. P. SANYAL (1980): A repoart on the Reptilia fau- na of Andaman and Nicobar Islands in the collectionof Zoo- logical Survey of India. Rec. Zool. Surv. India, Calcutta 77: 255-292. BLYTH, E. (1863): Report of the curator, zoological department. Jour. Asiatic Soc. 32 (1): 73-90. 284 Jakob HALLERMANN: A new species of Bronchocela from Nicobar Island Das, I. (1999): Biogeography of the amphibians and reptiles of the Andaman and Nicobar Islands, India. In: Tropical Island herpetofauna. Origin, current diversity and conservation. (ed. H. OTA) Developments in animal and veterinary science 29: 43-77, Elsevier, Amsterdam-Lausanne-New York-Shannon- Singapore-Tokyo. DE Roos, N. (1915): The reptiles of the Indo-Australian Arch- ipelago. I. Lacertilia, Chelonia, Emydosauria. E.J. Brill Ltd., Leiden. 348 pp. Dione, C. H. & S.S. L. Lim (1998): Taxononic review and mor- phometric description of Bronchocela cristatella (Kuhl, 1820) (Squamata: Agamidae) with notes on other species in the genus. Raff. Bull. Zool. 46 (2): 345-359. HALLERMANN, J. (2005): A taxonomic review ofthe genus Bron- chocela (Squamata: Agamidae), with description of a new species from Vietnam. Russ. J. Herpetol. 12 (3): 167-182. Appendix: Material examined Leviton, A. E., GIBBS, R. H. Jr., HEAL, E. & C. E. DAWSON (1985): Standards in herpetology and ichtyology: Part 1. Stan- dart symbolic codes for Institutional resource collectios in herptology and ichtyology. Copeia 1985 (3): 802-832. SMITH, M. A. (1935): The fauna of British India, including Cey- lon and Burma. Reptilia and Amphibia. Vol. II Sauria. Taylor and Francis, London. 440 pp. Tiwari, K. K. & S. Biswas (1973): Two new reptiles from the Great Nicobar Island. J. Zool. Soc. India 25: 57-63. WERMUTH, H. (1967): Liste der rezenten Amphibien und Rep- tilien: Agamidae. Das Tierreich 86: 1-127. Bronchocela rubrigularis. India: Nicobar Island: Trincat Island (ZMH RO09271 (field no Tri 25) holotype); Katchin Island (ZMH R09272 (field no Kat 43) paratype), Bompoka Island (No catalogue no. (field No. BOM 21)); Nancowry Island (No catalogue no. (field NAN 15)); Camorta Island (No catalogue no. (field no. CAM 76)), Teressa (Tarassa) Island (No catalogue no. (field TER 39)); Chowra Island (No catalogue no. (field no. CHO 05)), Bompoka Island (No catalogue no. (field no. BOM 27. Bronchocela cristatella. Indonesia: Sumatra: Serdang 3°30°N 98°50°E (ZMH RO608; ROS602-3); NE Sumatra: Kwalu (ZMH RO05623), Sumat- era Barat: Indrapura 2°04°S,100°56’E (ZMH RO5600);Tandjong: Padang District 0°38°S 100°52’E (ZMH RO05624); Environs of Pispis 3°10’N,99°01’E, mountain region (ZMH R04928-9): Pulo Weh Island: Sabang (ZMH R05625); Indrgiri: Riau: Sungei Lalak 0°27’°S 102°59°E (ZMH R06075-6); Nias Island (ZMH R04893-5). Bonner zoologische Beiträge | Band 56 | Heft 4 Seiten 285-297 Bonn, November 2009 Studies on African Agama VII. A new species of the Agama agama-group (Linnaeus, 1758) (Sauria: Agamidae) from Cameroon & Gabon, with comments on Agama mehelyi Tornier, 1902 Philipp WAGNER*!, Michael F. BAREJ! & Andreas SCHMITZ? ! Zoologisches Forschungsmuseum A. Koenig, Adenauerallee 160, D-53113 Bonn, Germany. 2 Museum d’histoire naturelle, C.P. 6434, CH-1211 Geneva 6, Switzerland. * Corresponding author: philipp.wagner.zfmk(@uni-bonn.de. Abstract. In the course of recent taxonomic studies in the African Agamidae the West African species were examined and new species have been identified. In this publication a new species of the genus Agama Daudin, 1802 is described from Cameroon and Gabon. Males of the new species differ from all other known Agama in the unique combination of the reticulate colouration of the throat and the blue tip of tail. It is compared with the other Cameroonian members of the genus and several other Agama species from Africa. Additionally, the status of Agama mehelyi Tornier, 1902, only known from its holotype, is discussed. Key words. Reptilia: Sauria, Agamidae, Agama sp. n., Agama mehelyi, Africa, Cameroon, Gabon, morphology, taxon- omy. INTRODUCTION After research activities on the genus Agama Daudin, 1802 in the last few years many species were described, syn- onymised or revalidated (PADIAL 2005, WAGNER 2007, WAGNER et al 2008a, WAGNER et al. 2008b). Currently, the genus contains 35 recognised species, but preliminary re- sults (especially from the West African clade of the genus) show several cryptic taxa within the Agama agama (Lin- naeus, 1758) complex. Therefore, many populations are under investigation and especially vouchers from Cameroon were of special interest to the authors, because the type locality of Agama agama has been restricted to this country. However, until recently it was not possible to character- ize the ‘true’ Lacerta agama Linnaeus, 1758, because the syntypes illustrated in SEBA (1734) are not available and the proposed type material in the Museum Adolphi Fri- derici collection of the Naturhistoriska Riksmuseet (= The Swedish Museum of Natural History) is not identical with the illustrated specimens. WAGNER et al. (2009b) clarified this situation by designating a neotype (ZFMK 15222) from Cameroon. The authors followed MERTENS (1938) who restricted the erroneous type locality ‘America’ of Agama agama to Cameroon but failed to fix this decision, according to article 76 of the Internatio-nal Code of Zo- ological Nomenclature (ICZN 1999) by designating a lec- to- or neotype. Now, after the proper de-finition of a neo- type (WAGNER et al. 2009b), a taxo-nomic revision of this species group is possible and the already recognised new species can be described. CHIRIO & LEBRETON (2007) listed seven species of Aga- ma for Cameroon: Agama a. agama (Linnaeus, 1758); Agama doriae benueensis Monard, 1951; Agama gracil- imembris Chabanaud, 1918; Agama mehelyi Tornier, 1902; Agama paragama Grandison, 1968; Agama sankaranica Chabanaud, 1918 and Agama sylvanus MacDonald, 1981. In the adjoining countries of Cameroon, no other than the above mentioned Agama species occur (WAGN- ER, unpubl. data) but CHIRIO & LEBRETON (2007) also identified four probable new Agama species from Cameroon and one of them (Agama sp. 1) turned out to be identical with the new species recognized by us before, and we describe it herein. 286 Philipp WAGNER et al.: Studies on African Agama VII. New species of the Agama agama-group MATERIAL & METHODS The type material of the new species is deposited in the Zoologisches Forschungsmuseum A. Koenig, Bonn, Zo- ologisches Museum für Naturkunde, Berlin and the Museum d’histoire naturelle, Geneva. It was compared with vouchers from Cameroon and other African countries housed in those collection (see appendix) and with data from literature (GRANDISON 1968, CHIRIO & LEBRETON 2007). Measurements were taken with a digital calliper to the nearest of 0.1 mm. Measurements and scale counts were done following GRANDISON (1968), MoobY (1980) and Moopy & BÖHME (1984). Molecular data were collected to examine sequence vari- ation between the new species and its morphologically closest relatives. A portion of the mitochondrial 16S rRNA gene of the holotype of the new species (ZFMK 87698; GenBank accession number: GU133316), a series of oth- er specimens of the new species (no voucher; GenBank accession number: GU133315; ZFMK 73241; GenBank accession number: GU133317; ZFMK 75376; GenBank accession number: GU133318; ZFMK 83762; GenBank accession number: GU133319; ZFMK 83766: GenBank accession number: GU133320), as well as the proposed neotype of Lacerta agama (ZFMK 15222; GenBank ac- cession number: GU133323) from northern Cameroon, Agama sankaranica (ZFMK 84992; GenBank accession number: GU133327) from an unknown locality, Agama paragama (ZFMK 15244; GenBank accession number: GU133321) from Cameroon, Waza, Logone et Chari, Aga- ma agama africana (ZFMK 73845, ZFMK 73846; Gen- Bank accession numbers: GU133311, GU133312) both from Senegal, Dakar-Bel Air, Agama agama ssp. (MH- NG 2689.53; GenBank accession number: GU133310) from Benin, Agama boueti (ZFMK 80057) from Mauri- tania, 30km NW of Rosso, and Agama finchi (ZFMK 83652; GenBank accession number: GU133314) from Kenya, Malaba (type localıty) were sequenced. Sequences for Agama planiceps (GenBank accession number: AF355476) and Agama castroviejoi (GenBank accessıon number: AY522929) were added to the dataset from al- ready published sequences (MATTHEE & FLEMMING 2002, PADIAL 2005). Acanthocercus atricollis (ZFMK 41748; GenBank accession number: GU133322 Botswana, Gaborone) was chosen as outgroup. DNA was extracted using QuiAmp tissue extraction kits (Quiagen) or a modified Chelex-Protocol (WALSH et al. 1991, SCHMITZ 2003). The primers 16sar-L (light chain; 5’— CGC CTG TTT ATC AAA AAC AT — 3”) and 16sbr- H (heavy chain; 5” - CCG GTC TGA ACT CAG ATC ACG T — 3’) of PALUMBI et al. (1991) were used to am- plify a portion of the mitochondrial 16S ribosomal RNA gene. PCR cycling procedure was as described in SCHMITZ et al. (2005). PCR products were purified using Quiaquick purification kits (Quiagen). Sequences were ob- tained using an automatic sequencer (ABI 377). Sequences were aligned using ClustalX (THOMPSON et al. 1997; de- fault parameters) and manually checked using the origi- nal chromatograph data in the program BioEdit (HALL 1999); this resulted in a total of 505 bp for the chosen sec- tion of the 16S gene. Phylogenetic trees were calculated in the program Paup* 4.0b10 (SwoFFORD 2002) using the neighbor-joining algorithm (NJ) and bootstrapping with 20000 pseudo-replicates to estimate node support as well as with a maximum parsimony approach (heuristic search with TBR branch swapping, stepwise random addition with 100 replicates, bootstrapping with 2000 pseudo-repli- cates). Paup* 4.0b10 was also used to compute the uncor- rected pairwise distances for all sequences. The following measurements and scale counts were part- ly used to compare the different species (see table 1): Snout-vent length (SVL): measured from mental scale to cloaca; tail length (TL): measured from cloaca to tip of tail; head length (HL): measured from jugale to rostral scale; head height (HH): measured at the jugal-postorbital region; head width (HW): measured across the jugal-pos- torbital region just anterior to the external auditory mea- tus; Midbody scales (MS): scale rows around midbody; vertebral scales (VS): number of midline vertebral scales, counted from midpoint of pectoral region to midpoint of pelvic region; dorsal scales (DS): number of midline lon- gitudinal dorsal scales, counted from midpoint of pectoral region to cloaca; cloacal pores (CP). RESULTS & DISCUSSION Species of the genus Agama, unlike chameleons, the sis- ter taxon of the Agamidae (e.g. MACEY et al. 2000), are very conservative in body form, without developing or- naments like e.g. horns or ear laps. Only in some species body ornamentations like enlarged nuchal or tail crest are present. If one takes into consideration the results from the chameleons (ZIEGLER & BOHME 1997; BOHME & ZIEGLER 2008), where morphologically similar species show high variations in hemipenis structures, a similar sit- uation in the Agamidae could be expected. In contrast, pre- liminary results (BOHME 1988; WAGNER et al., unpubl. da- ta) show a high level of homoplasy and conservative mor- phology in the hemipenis structures in the genus Agama. Consequently, beside from body form and hemipenis structure, there must be another mechanism of species recognition. A pre-mating mechanism was identified by several authors (LOVERIDGE 1933, THYS VAN DEN AUDE- NAERDE 1963, MCLACHLAN 1981, BOHME et al. 2005, Bonner zoologische Beitráge 56 287 Acanthocercus atricollis ZFMK 41748 82 53 94 57 100 <> 100 100 100 0.1 Agama sankaranica ZFMK 84992 Agama planiceps AF355476 Agama finchi ZFMK 83652 Agama agama ssp. MHNG 2689.53 84 = Agama agama africana ZFMK 73845 100 94 Agama agama africana ZFMK 73846 Agama agama agama ZFMK 15222 Neotype 84 80 Agama paragama ZFMK 15244 Agama lebretoni no voucher 100 100 Agama lebretoni ZFMK 87698 Holotype Agama lebretoni ZFMK 73241 Agama lebretoni ZFMK 75376 Agama lebretoni ZFMK 83762 Agama lebretoni ZFMK 83766 Agama boueti ZFMK 80057 Agama castroviejoi AY522929 Fig. 1. Phylogenetic tree based on 505 bp of the mitochondrial 16S rRNA gene. Values above the nodes represent neighbour-joi- ning bootstrap (20000 pseudo-replicates) values in percent while the values blow the nodes are the corresponding maximum par- simony supports. Significantly supported values are in bold. Values below 50% are not shown. The branch length from the out- group (Acanthocercus) to the ingroup (4gama) has been shortened for visual purposes. 288 Philipp WAGNER et al.: Studies on African Agama VII. New species WAGNER 2007, WAGNER et al. 2008a, 2008b) as the colouration of head, throat, forelimbs and tail of nup- tial males. In conclusion, the colouration of dominant males and, sometimes, of pregnant females can be used for the identification of the different species in the genus Agama, but not for analysing phylogenetic relation- ships. One of the main characters is the colouration pat- tern of the throat (JACOBSON 1992; MCLACHLAN 1981; WAGNER 2007; WAGNER et al. 2008a, 2008b). Dominant males present their throat, head and forelimbs in a be- havioural context to other adult male competitors and, especially, females. Consequently, these characters are useful to distinguish different species and to compare the herein described new species with closely related taxa. Therefore, we compare the new species not only in pholidosis and morphometrics with other Agama species, chiefly with that ones occurring in Cameroon, but we turn our attention especially to the colouration of dominant males. Genetics: For the genetic dataset both phylogenetic cal- culation methods completely agree in the topology of the recovered phylogenetic tree (fig. 1), and only dif- fer in the degree of support recovered for the individ- ual nodes. The phylogenetic tree is rather well resolved with three major groups apparent. The first and basal fully sup- ported clade includes Agama boueti and the only recent- ly described A. castroviejoi (NJ: 100/MP: 100). The po- sition of this clade corresponds well with the position- ing of other extreme West- or North African taxa in the more complete phylogeny of LEACHE et al. (2009), therefore it is likely that these two taxa are part of a more comprehensive north-western clade within the genus Agama. The second clade is only well support- ed by the neighbour-joining analysis and is represent- ed in this tree by a single species (Agama sankarani- ca), and represents the “Sahel Radiation”-clade identi- fied by LEACHE et al. (2009). The third and largest clade includes a basal unresolved polytomy because the phylogenetic position of A. plan- iceps and A. finchi could not be resolved with the used gene fragment. But here the three well supported sub- clades are of importance: the first subclade includes the proposed neotype of Agama a. agama and the closely related A. paragama (NJ: 84/MP: 80), the second sub- clade includes the two Senegalese vouchers of Agama a. africana as well as a specimen from Benin (NJ: 84/MP: 62), while the third clade fully supported clade (NJ: 100/MP: 100) contains all the included vouchers of “Agama sp. n.”. OLESSEAV Sdaotunjd puDsp trZSsI MINAZ Dlupsp.ind vunsp 99LER MINAZ u ds mojaiga] DUDS 0 0 Cc cel 0 IZvO'0 +ysO cLr00 ITrOO 09L0'0 L860°0 0F60'0 9TFO'0 L280'0 ESILO 86010 € 6rr00 6St0'0 6rr00 6br0'0 e800 8rr0'0 6rr00 oFrO'0 6FF0'0 C err00 pol 18£0'0 00 18£0'0 967 €Tr0'0 vO0 c 8£0'0 Ibt0'0 C 00000 10€E0°0 7800 28200 8E00 00000 c 78200 18200 v8LO'O LOOL'O 1E60°0 18£0'0 00000 00000 00000 00000 ri Qr, Fa] OLEX MWAZ U “ds 110]9.1(/9] DUD. c Isco'O = 18e0'0 ZL6I'0 18£0'0 v8LO'O LOOTO 1600 18£0'0 00000 10E0°0 0000'0 00000 tl OLESL MINAZ u “ds 1072.19] DUDSP 18200 18t00 ceolO 18£0'0 vy8LO'0O LOOLO 1600 1800 r8L00 LOOLO 1600 18 00000 TOt0'0 00000 00000 00000 al IPZEL MWAZ u ds iojaiga] DIOS Y (adAJO[OH]) 869.8 MWAZ u “ds 110/2.19] DIOS Y £61'0 I8E00 7 18£0'0 18£0'0 1820'0 00 c t 00000 10t0'0 of the Agama agama-group ce61 0 18200 [8£0°0 P8L0'0 LOOL'O 1E60°0 18£0'0 00000 10£0'0 (19YINOA ou) "u ‘ds 111072.19] DUDS 18£0°0 18€0°0 TEL oO 18E0°0 P8L00 LOOLO 1E60°0 L800 10£0'0 8 ANAZ 1y9/ puvsp 2668 MINAZ PO1UDADYUDS vumdy cS9 10E0°0 1E61'0 cTLO'0 L1£0'0 €8L0°0 00L0°0 LPOl0 [L60°0 OTZOO 07ZO0'0 6v61'0 10LO'0 Ir70'0 L801'0 28010 ISTIO £6110 10200 6T6TZS AV 10/0140.1I80) DUDS COLT O 99110 L80L'0 LIOTO 00000 Lrr0'0 0070'0 LSO08 MINAZ Vanog puvsp OY8€L MINAZ pupaLıfp DUDSD DUDS 00070 cloro Toro 11010 un Te81'0 SP8EL MINAZ Yuva DUDED DUDE Y (adAXJ09N) 22751 MINAZ PDD DUDID DUDS Y ces 0 Lyt0'0 00700 8S9t0'0 ELOTO ES’689T ONHW ‘dss DUDID DUDS Y St/L It MINAZ $111091.409D SNOIAAIOYIUDIP 20610 UOXEL 91 SI rl tl zı II OL LI ‘Apnjs sıyy ur posn sasuanbas YN AS9] [erıpuoyvonuw oy} Jo dq SOS 10] saQue]sIp-d PSPSMODUN “E EL Bonner zoologische Beiträge 56 289 Fig. 2. Images of Agama lebretoni sp. n. alive: A. Pregnant female of Agama lebretoni sp. n. from Mt. Nlonako, Cameroon (ZFMK 75376) | B. Non-pregnant female of Agama lebretoni sp. n. from Nyasoso, Mt. Kupe, Cameroon | C. Close-up of the head of the living holotype of Agama lebretoni sp. n. from northeast of Mamfe, Mukwecha, Amebisu, Cameroon. | D. Holotype of Agama le- bretoni sp. n. in life Pe 290 Philipp WAGNER et al.: Studies on African Agama VII. New species of the Agama agama-group Fig. 3. Preserved holotype of Agama lebretoni sp. n. (ZFMK 87698) from nordeast of Mamfe, Mukwecha, Amebisu in Came- roon. Bonner zoologische Beiträge 56 291 Table 1 summarizes the uncorrected p-distances for the used gene fragment. The neotype of Agama a. agama shows only a comparatively low genetic difference (2.96%) to its sister taxon A. paragama, while it differs from its western ‘subspecies’ Agama a. africana by 4.47% and from Agama sp. n. by 3.81%. The latter two species are separated by an equal genetic distance (3.81%). These distances are of similar magnitude than to other well es- tablished species within this large clade (A. a. agama [neo- type]-A. planiceps 4.72%; A. a. agama [neotype]-A. finchi 3.17%; A. finchi-A. planiceps 3.59%). Regarding the ge- netic distances of vouchers which we morphologically confirmed to be members of the same taxon we find them to be genetically identical to each other; this is true both for the two Agama a. africana specimens as well as for all six vouchers of in the third subclade (Agama sp. n.). The identified genetic structure (low to non-existent in- tra-clade distances and moderately higher but more or less equidistant inter-clade distances) is typical for rather young radiations. Our genetical analyses therefore sup- ports the idea of previous studies (e.g. WAGNER 2007; WAGNER et al. 2008a, 2008b) that many of the previous- ly recognised species in the genus Agama in reality com- prise of several taxa, which are often very difficult to sep- arate on the basis of external morphology alone, and the same now holds true for Agama agama sensu lato. The inclusion of morphologically well established species (A. planiceps, A. finchi, A. paragama), which show near- ly equal genetic distances to the neotype of A. a. agama prove that the uncovered distance of Agama sp. n. to all other closely related congeners confirms the specific dis- tinctness of Agama sp. | and together with morphologi- cal evidence (see below) warrants the formal recognition of this species. Agama lebretoni sp. n. Holotype (figs. 2c-d, 3, 4.1) ZFMK 87698 (field number MMO075): Cameroon, north- east of Mamfe, Mukwecha, Amebisu, N 05°53.866° E 009°33.495’, leg. J. Wurstner & M. Barej, September, 2007. Paratypes ZFMK 87694 — 697, 87699 (paratopotypes): Cameroon, Mamfe region, Mukwecha, Amebisu, leg. J. Wurstner & M. Barej, September, 2007. ZFMK 87700: Cameroon, Mamfe, leg. J. Wurstner & M. Barej, September, 2007. ZFMK 61243: Cameroon, Korup, Mundemba, leg. C. Wild, February 1989. ZFMK 51686: Cameroon, Magba, leg. F. Schütte, Febru- ary 1990. ZFMK 75376: Cameroon, Mt. Nlonako, Nguengue, leg. H.-W. Herrmann & A. Schmitz, January 2000. ZMB 55709, 37061: Cameroon, Makum. MHNG 2713.29 (field number AMC-360): Mofako Balue, Rumpi Hills, Cameroon, leg. locals people, July 2009. MHNG 2713.30 (field number AMC-248): Big Massaka, Rumpi Hills, Cameroon, leg. M. Bare) & A. Schmitz, Ju- ly 2009. MHNG 2713.31 (field number AMC-009): Foyer du Marin, Douala, Cameroon, 01.07.2009, leg. M. Barej & A. Schmitz. MHNG 2713.32 (field number AMC-178): Big Massaka, Rumpi Hills, Cameroon, 09.07.2009, leg. by locals. Diagnosis. A fairly large species of Agama (total length of adult male above 25 cm), which is characterized by a reticulated pattern on the throat, a bright vertebral stripe and a deep blue tail tip in adult males. Males of the new species are distinguishable from all other Agama-species by the combination of throat and tail colouration. The throat colouration (fig. 4.1) is a reticular pattern of red lines, which is so far only known from Agama paragama (fig. 4.2), Agama sylvanus (fig. 4.3) and an undescribed species from Uganda (fig. 4.5). The tail colouration is unique within the genus because of the blue tip of a tri- coloured tail (see fig. 2d). Females are similar to those of other species of the Agama agama species complex, and can only be safely determined by their genetics. From southern and eastern African species A. /ebretoni sp. n. differs as follows: From A. planiceps, A. turuensis, A. mwanzae, A. kaimosae and A. atra the new species differs in their unique throat colouration, the blue colouration of the tip of the tail and in having a vertebral stripe. From A. knobeli the new species differs additionally in not possessing a tail crest. From A. mucosoensis the new species differs in having a red head instead of a yellow. A. lebretoni sp. n. is very dif- ferent in tail colouration (tri-coloured instead of banded narrow white and blue) from A. lionotus and A. kirkii. Agama species occurring in Cameroon and Gabon differ from A. lebretoni sp. n. as follows: Agama agama differs in having a black colouration of the tail tip instead a blue one in the new species. Both species also differ in characters of throat colouration: Agama aga- ma has a more or less striated to uniform red throat (fig. 4.4), which is dissimilar to the reticulated throat of the new species. A. agama is also lacking the typical pale verte- bral stripe which is present in nuptial coloured males of Philipp WAGNER et al.: Studies on African Agama VII. New species of the Agama agama-group 159) Ke) ine) Fig. 4. Throat patterns of some West African adult males of Agama species: 1= Agama lebretoni sp. n. (holotype, ZFMK 87698); 2= Agama paragama (ZFMK 15244); 3= Agama sylvanus (ZFMK 40252); 4= Agama agama (neotype, ZFMK 15222); 5= Agama sp. n. (ZFMK 88809); 6= Agama a. africana (ZFMK 20125); 7= Agama finchi (NMK L2716); 8= Agama planiceps (ZFMK 55062); 9= Agama lionotus lionotus (ZFMK 83624); 10= Agama lionotus el- gonis (ZFMK 82065); 11= Agama lionotus dodomae (ZFMK 83706); 12= Agama kaimosae (NMK L2715); 13= Agama doriae benueensis (ZFMK 29615); 14= Agama doriae doriae (ZFMK 55545); 15= Agama sankaranica (ZFMK 40468); 16= Agama gra- cilimembris (ZFMK 33719). Bonner zoologische Beitráge 56 293 the new species. In contrast, Agama agama has a slight- ly lower count of scale rows around midbody. According to GRANDISON (1968) A. agama from Nigeria has 59 to 77 rows, whereas A. lebretoni sp. n. has 73 to 80 rows [23 specimens]. From Agama doriae benueensis the new species differs in having a different throat pattern in dominant males: 4. doriae with a prominent black dot on the base of the throat (fig. 4.13, 4.14), instead of a reticulated throat in A. le- bretoni sp. n. Additionally, A. doriae benueensis is one of the few 4gama species which have the nostril below the canthus rostralis instead of on the canthus like in the most Agama’s. Agama gracilimembris is much smaller in size (100 to 120 mm in A. gracilimembris in difference to more than 250 mm in A. /ebretoni sp. n.) and does not have spinose scales around the ear opening as in the most other Agama species. Again, also this species differs in the colouration of the throat of adult males: striated in 4. gracilimembris (fig. 4.16) and reticulated in the new species. In morphology, A. gracilimembris differs in possessing strongly keeled head scales and it is also one of the few Agama species which have the nostril below the canthus rostralis. How- ever, both species are similar in the count of scale rows around midbody (70 to 85 in A. gracilimembris and 73 to 80 in the new species). From Agama paragama the new species differs in pos- sessing a lower nuchal crest, in having a red instead of a yellow-whitish head and in having a higher number of scale rows around midbody (Wagner, unpubl. data). Ad- ditionally, adult males of A. paragama show a black in- stead of blue tail tip. GRANDISON (1968) described the colouration of the throat of A. paragama (similar to the herein described new species) as ‘a dark network on a cream ground which takes the form of isolated, round, cream spots’ (fig. 4.2) and as dissimilar to Agama agama which has a ‘longitudinal arrangement of darker lines or blotches’. However, GRANDISON (1968) also mentioned, that the typical throat colouration of A. paragama is pres- ent in both sexes, which is not comparable with 4. lebre- toni sp. n.. A similar situation exists in Agama turuensis and Agama lionotus elgonis which have a very similar colouration of the throat to each other (see fig. 4) but are recognized as distinct taxa by WAGNER et al. (2008a). Agama sankaranica is completely different in body colouration, but also in other aspects of morphology: this species is much smaller in size and it is one of the few Agama lizards which have the nostril below the canthus rostralis. Additionally, A. sankaranica has a fewer count of scale rows around midbody (64-78 [69.6] in sankaran- ica versus 73 to 80 [76.2] in lebretoni sp. n.) and a very large occipital scale, which is as large as one and half di- ameter of the tympanum. Agama lebretoni sp. n. differs clearly from Agama syl- vanus in having a vertebral stripe instead of a complete- ly blue body in A. sylvanus, but both species show a sim- ilar pattern of throat colouration in adult males (figs 4.1, 4.3). Agama lebretoni sp. n. differs from the holotype of 4ga- ma mehelyi (for comments see below) in having a homog- enous body scalation. From the three other proposed new species mentioned by CHIRIO & LEBRETON (2007) the new species differs as fol- lows: from Agama sp. 2 in the larger size; from Agama sp. 3 in having the nostril on the canthus rostralis; from Agama sp. 4 in having a vertebral stripe and completely blue forelimbs instead of red and blue coloured ones in Agama sp. 4. Comparison of 4. lebretoni sp. n. with probably valid synonyms of Agama agama Agama colonorum var. congica Peters, 1877 was described from Chinchoxo, Cabinda, Angola. One syntype (ZMB 9196) resembles in colouration more Agama agama. The typical characteristics of A. lebretoni sp. n. (reticulated throat, white speckled body and blue tip of tail) are lack- ing in this specimen. The other syntype (ZMB 67193) is lacking colouration but shows a relatively large body sca- lation in difference to A. lebretoni sp. n.. It is most prob- able that Agama colonorum congica 1s a valid species but further investigations are needed. Agama picticauda Peters, 1877 was described by a series of six syntypes from Ada Foah in Ghana (Adafer in Mau- ritania fide LOVERIDGE 1957, which is obviously in error), from Acera in Guinea (= Accra in Ghana?) and from an unknown locality in Cameroon. DENZER et al. (1997) des- ignated the specimen ZMB 403 from Ada Foah as lecto- type. This subadult specimen resembles in colouration and morphology Agama agama. Nevertheless, the paralecto- type series mistakenly included an adult male of A. /ebre- toni sp. n. (ZMB 8299) from Cameroon. Therefore, be- cause of the chosen lectotype, this taxon must be recog- nized as synonym of Agama agama. Differences between A. agama and A. lebretoni sp. n. have already been dis- cussed above. Description of the holotype (figs 1 c—d, 2, 4.1). Habitus stout, snout-vent length (SVL) 140.0 mm, tail length (TL) 199.6 mm, head length (HL) 36.0 mm, head with (HW) 26.0 mm, head height (HH) 15.8 mm. 294 Philipp WAGNER et al.: Studies on African Agama VII. New species of the 4gama agama-group O 1 02 Cameroon @13 Equatorial Guinea Fig. 5 Distribution of Agama lebretoni sp. n. CAMEROON. 1= Makum (ZMB 55709, 37061); 2= Magba (ZFMK 51686, 54906 - 907); 3= Metchum, Wum (ZFMK 15194 — 15200); 4= Amebisu [=Amebesu] [type locality]; (ZFMK 87694 — 699); 5= Mamfe (ZFMK 87694 — 699, 87700); 6= Nguengue, Mt Nlonako (ZFMK 69017); 7-9= Rumpi Hills: Mofako Balue, Big Massaka (MHNG 2713.29 — 30, 2713.32); 10= Mundemba (ZFMK 61243); 11= Douala (MHNG 2713.31); 12= Limbe (Victoria) (ZFMK 18891 — 894). EQUATORIAL GUINEA. 13= Bioko Island (Fernando Poo), San Carlos (ZFMK 9353 — 359). GABON 14= Ngouassa (IRSNB 15686 — 687); 15= Fougamou (ZFMK 73239 — 245). Bonner zoologische Beiträge 56 295 Large triangular nasal scale slightly above the canthus ros- tralis and pierced with the nostril in the posterior part, di- rected and supplied obliquely upwards. Between the nasal scales, two narrow longitudinal, keeled scales are visible, posterior followed by one smooth transverse scale. Nine supralabial scales, eight sublabial scales on both sides. Head scales between the eyes smooth, directed sideward from a midline between the eyes; head scales between pos- terior end of the eyes and neck keeled, directed forwards; head scales of the temporal region keeled, directed side- wards; free anterior margins of head scales with numer- ous sensory pits; supraocular scales smooth. Parietal shield more or less pentagonal, pineal organ visible, pierced in the middle of the shield; parietal shield surrounded by scales more or less equal in size. Ear hole large, about the same size as the eye, margin being composed by spiny scales, surrounded by five tufts (three anterior, two pos- terior) of more or less spiny, mucronate scales; tympanum superficial. Nuchal crest low, consisting of 17 lanceolate scales. Gular scales flat, smooth, juxtaposed and becom- ing smaller towards the gular fold. Dorsal body scales strongly keeled and mucronate, equal in size, in 67 scales from midpoint of pectoral region to midpoint of pelvic re- gion. Ventral body scales smooth, slightly imbricate at their posterior margins, in 86 scales from midpoint of pec- toral region to midpoint of pelvic region. Together in 79 scales rows around midbody. Tail scales strongly keeled and mucronate. One row of nine precloacal scales. Scales on the upper side of the forelimb strongly keeled, smooth on the underside, on the upper arm scales thrice as large as the dorsal body scales, becoming smaller towards the underside and the manus. 4th finger longest, digital length decreasing 3-2-5-1, subdigital lamellae keeled and mu- cronate. Scales on the upper side of the hindlimb strong- ly keeled becoming smooth on the underside, on the up- per tights equal in size as the dorsal body scales becom- ing larger towards the lower tights. 4'h toe longest, digi- tal length decreasing 3-2-5-1. Colouration (in alcohol after 5 months of preservation). Head and neck red. Limbs and body blue, body darker as the limbs. Posterior part of the neck, body and upper parts of the hindlimb speckled with white scales. Between the limbs with a whitish vertebral stripe. Tail at the base blue, followed by white, red and blue. Belly and underside of the limbs blue, tail anterior whitish, posterior bluish. Throat, gular fold and parts of the extreme lower head red with a white speckling, resembles at the throat a reticu- late pattern. Colouration in life. Males: head and nape deep red; body and limbs electric to dark blue, body and sometimes up- per parts of the limbs speckled white to yellowish; whitish vertebral stripe between the limbs; tail four-coloured: blue (extending from the body), whitish in a transition zone to red and dark blue at the tip; belly and underside of limbs blue; throat with a reticulate pattern of red and white, sometimes more or less red with a white speckling. Fe- males: Non-pregnant females and juveniles are vividly coloured. Ground colour is grey-brown, head and forepart of the body blotched with yellow spots, stripes and bars. Body usually with a fine and pale vertebral stripe; yellow and black framed blotches on the lateral parts; along the vertebral zone hexagonal black stripe markings visible; body is also blotched with dark dots and short stripes; sometimes an orange coloured broad stripe on the lateral parts obvious (see fig. 2b). Pregnant colouration of females is clearly different: body becomes uniform brown in ground colour; yellow to orange blotches on head more or less blurred. Body colouration ıs dominated by broad orange waved or serrated bands (see fig. 2a). Variation. For some measurements and scale counts in A. lebretoni sp. n. see also table 1. Colouration in males and females 1s variable within the typical pattern of the species and depends on many different factors (see also colouration). Non-dominant males resemble the non-preg- nant female respectively juvenile colouration. But also dis- turbed dominant males can loose their typical colouration rapidly. Dominant males vary in having a distinct or non- distinct vertebral stripe and in the intensity of spots on the body. Females vary in pregnant and non-pregnant coloura- tion. A turnover colouration is unknown. Etymology. The new species 1s dedicated to MATTHEW LE- BRETON, Yaoundé (Cameroon) in recognition of his con- tributions to the herpetology of Cameroon. Distribution. The new species is so far documented by us from Cameroon, Gabon and Bioko Island (= Fernan- do Poo, Equatorial Guinea) (for details see fig. 5). Accord- ing to CHIRIO & LEBRETON (2007) A. lebretoni sp. n. (=Agama spec. 1) is very widespread in western Cameroon. So far only two populations are known from Gabon, but also here a more widespread distribution can be assumed. In Cameroon, the new species also occurs along the western border to Nigeria and therefore an oc- currence of A. /ebretoni sp. n. can be expected for this country. Relationships. Agama lebretoni sp. n. should be closely related to Agama agama. As mentioned in this latter in- troduction this species also occurs in Cameroon with its type locality in the northern part of the country. Both species are very similar in colouration and can be distin- guished by the pale vertebral stripe, the white body speck- ling and the blue tip of the tail in 4. lebretoni sp. n. 296 Philipp WAGNER et al.: Studies on African Agama VII. New species of the Agama agama-group Comments on Agama mehelyi Tornier, 1902 TORNIER (1902) described this species on the basis of a single female specimen (ZMB 18149) from Bipindihof which was collected by the botanist Georg August Zenker (1855-1922). Since that time no other voucher of the species was collected or observed. While Moopy (1980), DENZER et al. (1997) and CHIRIO & LEBRETON (2007) list- ed A. mehelyi as valid species none of these authors ex- plained their decision about the validity. WERMUTH (1967) on the other hand stated clearly that he considered the validity of A. mehelyi as doubtful. Therefore, the type was examined and compared with other 4gama-species with a heterogeneous body scalation. From (1) A. hispi- da Kaup, 1827 and A. etoshae McLachlan, 1981 the species differs in having a large ear hole. From (2) A. ar- mata Peters, 1855 it differs in having no reticulated throat pattern and no strongly keeled dorsal scales. From (3) A. aculeata it differs in lacking a network pattern on the cen- tral part of the throat. The specimen resembles in scala- tion and colouration and especially in throat colouration pattern A. anchietae Bocage, 1896. Both species show dark wavy strips on the throat. A distinct colouration as in e.g. A. aculeata is lacking and both species are char- acterized by a pale vertebral stripe, four to five cross bands on the body (lacking in the preserved voucher of A. mehe- /yi) and a banded tail. Because of this striking similari- ties Agama mehelyi Tornier, 1902 is considered by us as a synonym of Agama anchietae Bocage, 1986. The locality of the specimen is doubtful. *Bipindihof” is a well known locality because of the collections made by Zenker on his own plantation. But Bipindihof is a rain- forest area where e.g. Lepidothyris fernandi (Burton, 1836) (Sauria: Scincidae), a typical rainforest lizard was found (WAGNER et al. 2009a). In Africa, no true rainfor- est agamid species occur and only species of the genera Acanthocercus and Agama sylvanus MacDonald, 1981 oc- cur in dense dry forest. Especially the members of the 4. hispida-group, like e.g. A. anchietae, are typical ground dwellers of arid regions. However, between 1889 and 1896 Zenker operated the Post Office of Yaounde and he later moved to Bipindihof to build up a plantation. From 1906 he also operated the Post Office of the area. As a passion- ate collector Zenker collected ethnographic curiosities, photographs, herbaria, skulls, stuffed birds, mammals, am- phibians and lizards. For this collections he probably al- so used his contacts to the Post Office to get material from other regions of Africa. Therefore, the given locality for A. mehelyi is most probably erroneous. Acknowledgements. We are thankful to Wolfgang Böhme and Aaron Bauer for comments on a further draft of the manuscript, to Mark-Oliver Ródel for the loan of important material from the collection of the Museum ftir Naturkunde in Berlin and to Adam Leaché for sequencing some of the used specimens. We are grateful to Julia Wurstner for photographs of the new species. REFERENCES BÖHME, W. (1988): Zur Genitalmorphologie der Sauria: Funk- tionelle und stammesgeschichtliche Aspekte. Bonner zoolo- gische Monographien 27: 1-176. BÖHME, W., WAGNER, P., MALONZA, P., KOHLER, J. & S. LOT- TERS (2005): A new species of the Agama agama group (Squa- mata: Agamidae) from western Kenya, East Africa, with com- ments on Agama lionotus Boulenger, 1896. Russian Journal of Herpetologie 12 (2): 143-150. 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Material examined Agama agama: SENEGAL. Dakar: Bel-Air: ZFMK 73845-849; CAMEROON. Diamaré, Mokolo: ZFMK 15222 [neotype]; unknown locality: ZFMK 83759 — 766. Agama colonorum var. congica: ANGOLA. Cabinda, Chinchoxo: ZMB 9196, ZMB 67193. Agama boueti: NIGER. Agadez: ZFMK 33706 — 714, 36595 — 598; Assaouas: ZFMK 20050 — 051; Dabnou: ZFMK 20051; El Meki: ZFMK 20048 — 049, 20046 — 047; Agadez: ZFMK 20044 — 045. SENEGAL. Dakar: 17169 — 174; Mboro-sur-mer: ZFMK 17176 — 183. Agama d. doriae: ETHIOPIA. Eritrea: ZFMK 20847; Gorgora Land, Lake Tana: ZFMK 19454 — 455; Kaffa, Omo river: ZFMK 15861 — 864. Sudan. Equatoria, Rihi river: ZFMK 29614 — 618; Dinder Pare: ZFMK 38397 — 399; Agama doriae benueensis: CAMEROON. Adamaoua, Martap: ZFMK 15192 — 193; Diamara: ZFMK 15176; Logne et Chari, Waza: ZFMK 15177 — 191; Margui-Wandala, Gouria: ZFMK 20085 — 088, Margui-Wandala, Koza: ZFMK 15175; Margui-Wandala, Mora: ZFMK 8855; Mokolo: ZFMK 33746; Mora: ZFMK 33738 — 745. CENTRAL AFRICAN REPUBLIC. Koumbala: ZFMK 33726 — 727; Ndélé: ZFMK 33747, 33728 — 733. NIGERIA. Beni Sheik: ZFMK 33734 — 737. Agama finchi: Kenya. Malaba: NMK L/2716/1, 4, 6-7, 9-10, ZFMK 83652-656. Aga- ma gracilimembris: CAMEROON. Benoué, Boki: ZFMK 15257 — 259; Benoué: ZFMK 33717. CENTRAL AFRICAN REPUBLIC. Koumbala: ZFMK 33718 — 721; Kotissako: ZFMK 33722 — 725. Agama kaimosae: KENYA: Ngoromosi: NMK L/2715/1, 3-4, ZFMK 83658 — 660; TANZANIA: Tanganyika District: Mwamalasa: MHNG 877.65; MHNG 2684.001 — 006. Agama kirkii: MALAwt. Mt. Mlanje: ZFMK 30696 — 697; Zomba: ZFMK 54533 — 534. ZIMBABWE. Zimba, Umfuruzi: ZFMK 51254. Agama lebretoni sp. n.: CAMEROON. Douala, Foyer du Marin: MHNG 2713.31; Fako (Mt. Cameroon), Limbe (Victoria): ZFMK 18891 — 894; Korup, Mundemba: ZFMK 61243; Magba: ZFMK 51686, 54906 — 907; Makum: ZMB 55709, 37061; Mamfe: ZFMK 87694 — 699, 87700; Mamfe, Mukwecha, Amebisu: ZFMK 87694 — 699; Metchum, Wum: ZFMK 15194 — 15200; Mt. Nlonako, Nguengue: ZFMK 69017, Rumpi Hills, Mofako Balue: MHNG 2713.29; Rumpi Hills, Big Massaka: MHNG 2713.30, 2713.32. EQUA- TORIAL GUINEA. Bioko Island, San Carlos: ZFMK 9353 — 359. GABON. Fougamou: ZFMK 73239 — 245; Ngouassa: IRSNB 15686 — 687. Agama mehelyi: CAMEROON: Bipindihof (in error): ZMB 18149 (holotype). Agama paragama: CAMEROON. Logone et Chari, Waza: ZFMK 15242 — 256. NIGER. Agadez: ZFMK 36599; Tessaoua: ZFMK 33749 — 750. Agama cf. paragama: CAMEROON. Benoué, Boki: ZFMK 15227 — 241. Agama pic- ticauda: GHANA: Ada Foah: ZMB 403. CAMEROON: ZMB 8299. Agama planiceps: NAMIBIA. Brandberg: ZFMK 33040 — 041; Frauenstein: ZFMK 18395 — 398; Kaoko, Werda: ZFMK 21961 — 962, Okahandja: ZFMK 2696, 2694 — 695; Omandumba: ZFMK 18399, 33007 — 009; Windhuk: ZFMK 46453 — 456. Agama sankaranica: BURKINA FASO. Ouagadougou: ZFMK 39032. MALI. Kassaro: ZFMK 20053; Monsombougou: ZFMK 20056; Sandaré: ZFMK 20057; Nioro du Sahel: ZFMK 20055. Toco. Lome: ZFMK 40468 — 484, 40525. Agama sylvanus: CAMEROON. Benoué: ZFMK 33751 — 754. CENTRAL AFRICAN REPUBLIC. Koumbala: ZFMK 40251 — 260; Mélé: ZFMK 33766; Ndélé: ZFMK 33755 — 7654; Sibut: ZFMK 33765. Agama turuensis: TANZANIA. Mount Hanang: ZFMK 74930 — 943, 82292 — 94, 82324 — 328, 82357 — 360, 82278 — 279. APPENDIX De “Agamis Abstracts of presentations hold on DEAGAMIS the 18ST INTERNATIONAL SYMPOSIUM ON AGAMID LIZARDS ORAL PRESENTATIONS History of Classifications and Phylogenies Scott M. Moopy Department of Biological Science, Irvine Hall, Ohio University, Athens, OH, 45701, USA; Email: moody(wohio.edu. DARWIN in his 1859 “Origin of species ...” eloquently demonstrated that taxonomic classifications could (and should) reflect the past evolutionary history of higher taxa and relationships of the extant species. However the major her- petological works, for example, BOULENGER’s Catalogue of Lizards, published over the next 100 years did not attempt reconstructions of phylogeny. Indeed BOULENGER reversed the earlier named (now recognized again) genera and high- er taxa proposed especially by W. PETERS and L. FITZINGER. Although, CAMP (Classification of Lizards, 1923) and HENNIG (Revision of Draco, 1936) published pioneering works (they were decades ahead of others) the taxonomic herpetologists ignored them until the “rebirth” of phylogenetic systematics in the 1960’s and 1970’s. Moopy (1980) published a phylogenetic analysis based on 122 morphological characters and reviewed the historical biogeography of the agamid genera and proposed several subfamilies. Within the past two decades beginning with JOGER (1991, a molecular phylogeny of agamid lizards) several workers have employed molecular analyses. The hypothesized mo- lecular phylogenies by MACEY, LARSON, ANANJEVA, PAPENFUSS, OTA, HONDA, SCHULTE and MELVILLE will be com- pared with my original and revised morphologically based hypotheses. The controversy over the familial status of the various higher taxa within the “acrodont” section of the suborder Iguania that ensued following FROST & ETHERIDGE (1989) also will be reviewed. The arid corridor from Middle East to Africa — Insights from the Agamidae Philipp WAGNER Zoologisches Forschungsmuseum A. Koenig, Adenauerallee 160, D-53113 Bonn, Germany; Email: philipp.wagner.zfmk(@uni-bonn.de. The similarities between fauna and flora of the arid southwest and northeast corners of Africa have been pointed out by several authors. E.g. POYNTON underlined this distribution pattern of an arid corridor with instances from Bufonid amphibians, whereas WAGNER found instances among reptiles, especially agamid lizards. These arid areas are still connected by a strip of arid country through Kenya, Tanzania and northeastern Zambia to southern Africa. This arid corridor allowed a faunal and floral exchange during dry phases of the Quaternary and influenced the colonization history of the continent. The relations and differences between Africa and Middle East are analysed using the agamid taxa Trapelus, Pseudotrapelus, Acanthocercus, Agama, Xenagama and Uromastyx. 300 DEAGAMIS — Abstracts Taxonomic, morphological and ecological diversity of Asian agamids (Agamidae:Acrodonta: Sauria:Reptilia) Natalia B. ANANJEVA Zoological Instutute, Russian Academy of Sciences, Sanct-Peterburg 194034, Russia: Email: azemiops@Zin.ru. One of important aspect of integrative study of lizards of Agamidae family is an analysis of morphological diversity within this group of squamates according to recognizing compositional, structural and functional biodiversity (Noss 1990). Agamidae is morphologically and ecologically diverse family belonging to Iguania that is a sister group to all the remaining squamates (SUKHANOV 1961; Moopy 1980; Estes 1983, 1985). Agamid lizards are characterized by acro-pleurodont dentition, lack of the intravertebral autotomy fracture plan existing in most other lizards and high structural diversity of integumental derivatives: scale sense organs, epidermal holocrine glands (femoral pores, cal- lose scalation), and so on. We analyze the representation of different evolutionary lines of agamid lizards in Palearc- tic and Oriental Asia, as well as morphological variety of dentition and integumental structures in these lines. The study of structure and development of dentition revealed a special type of anlage of the egg-teeth in Iguania in com- parison with another sugamates (SERGEYEV 1940; ANANJEVA & ORLOV 1986). The single egg-teeth anlage is synapo- morphy for Iguania whereas the paired tooth germs are shared by Gekkota, Scincomorpha, Anguinomorpha and all Ophidia. The next developmental transformations lead to paired state of egg-teeth in Gekkota including dibamids (Un- DERWOOD & LEE 2000) and unpaired (as a result of reduction of one of the two egg tooth germs, or as a result of fu- sion) state of egg-teeth in another squamates (ANANJEVA & ORLOV 1986). Attempts to nest integumental derivats into the phylogenetic tree of acrodonta lizards (MACEY et al. 2000; ANANJEVA 2004) and to consider association of morphological and ecological diversity of Asian agamids are discussed. Phylogenetic relationships and evolution of the agamid lizard subfamily Draconinae James SCHULTE II Department of Biology, 177 Clarkson Science Center, MRC 5805, 8 Clarkson Avenue, Clarkson University, Potsdam, New York, 13699-5805, USA; Email: jschulte@clarkson.edu. The agamid lizard clade (subfamily) Draconinae is widespread throughout the Indian subcontinent, Southeast Asia, Indonesia, the Philippines, and associated islands of the Indian and Pacific Oceans. Phylogenetic analyses are pre- sented for this clade at several hierarchical levels. Nuclear DNA recover very robust support for the agamid clade Agaminae as the sister taxon to draconines. Within Draconines, relationships are investigated using mitochondrial and nuclear DNA. Mantheyus phuwuanensis is strongly supported as the sister taxon to all other draconine species with Ptyctolaemus and Draco forming a clade that is sister to all remaining species. A detailed phylogeny of Calotes based on mtDNA also will be presented. Finally, the diversification rate of draconine agamids will be compared with Chamaeleons, Agaminae, and Amphibolurinae to identify possible evolutionary important differences between these clades. Status survey of the Indian Spiny-tailed lizard Uromastyx hardwickii in the arid regions of Rajasthan, North-Western India Madhuri RAMESH Group for Nature Preservation & Education, New No. 30, Block II, Gandhi Mandapam Road, Kotturpuram, Chennai 600 085 India; Email: madhurir@hotmail.com. The Indian Spiny-tailed Lizard Uromastyx hardwickii is endemic to the arid region between India and Pakistan. Known as the ‘Sanda’ in India, it is protected by law (Schedule II, Indian Wildlife Protection Act, 1972). Existing informa- EEE Bonner zoologische Beitráge 56 301 tion on this species is largely anecdotal and so sparse that the most detailed observations are almost a century old — this species inhabits self-excavated burrows and occurs in clusters; it is believed to be threatened by local trade in its meat and oil, and habitat destruction caused by anthropogenic activities. In order to effectively conserve this unique species basic information such as distribution, location of clusters and habitat characteristics is essential. Therefore, the objectives of this ongoing survey are to map the distribution of the Sanda in the arid regions of Rajasthan (north- western India), locate large clusters and identify habitats that can support such clusters. In addition, information on threats to the species including extent of exploitation is also being collected. The survey commenced in March and will continue till September 2007: so far, a combination of vehicular transects as well as interviews with locals has been used to assess presence/absence; abundance and habitat parameters have been measured using belt transects. Fur- ther, semi-structured interviews with hunting tribes has provided valuable information on exploitation. The survey will also be followed by a more detailed study on the ecology of this species since it is the only herbivorous lizard in the Indian subcontinent. The genera Pseudocalotes and Bronchocela in south East Asia: Taxonomy and distribution Jakob HALLERMANN Zoologisches Museum Hamburg, Martin-Luther-King-Platz 3, 20246 Hamburg, Germany; Email: hallermann@uni-hamburg.de. Uncertainty exists about occurrence and differentiation of the species of the genus Pseudocalotes in South-East-Asia. The situation in Vietnam and Laos is problematic due to problems in identification, occurrence of different colour morphs or undescribed species. New voucher specimens of that genus from different localities in Vietnam and Laos were examined and compared with type material. A first record of a female P poilani demonstrate sexual dimorphism and differences between P. poilani and P. floweri. Both species have a restricted distribution to southern Laos (poilani) and to southern Cambodia/ Thailand (floweri). In Vietnam P. microlepis brevipes are more widespread distributed than known. And a third unknonw species occurs in Kon Tum Province /Vietnam. A new species of Bronchocela from Nicobar Islands is presented with its variation and distribution. A distribution map for Pseucocalotes and Bronchocela is presented. A preliminary phylogeny of the genus Acanthosaura Gray 1931, inferred from mitochondrial and nuclear genes Perry Lee Woop JR. Villanova University, Department of Biology, 800 Lancaster Ave, Villanova, PA, 19085, USA; Email: perry. wood@villanova.edu. An analysis of mitochondrial and nuclear DNA from the Southeast Asian lizard genus Acanthosaura, was use to ass- es the phylogenetic relationships of the group. The specimens examined encompass the entire range of the genus, (Myan- mar, Thailand, Cambodia, Laos, China, and Malaysia (including its off shore islands, Pulau Tioman, Pulau Aur, Pu- lau Perhintian, and Pulau Langkaw1)) from all seven recognized species. A phylogenteic analysis using parsimony, likelihood, and bayesian analyses was used to test the hypotheses (1) Acanthosaura cf. crucigera from the Cardamom Mountains forms a distinct group separate from Acanthosaura crucigera from its type locality in Thailand (2) Acan- thosaura sp. from Pulau Aur forms the sister group to other A. armata, (3) to see if the Acanthosaura from Pulau Langkawi is more closely related to the Cambodia’s A. cf. crucigera then from the type locality in Thailand, follow- ing other biogeographic patterns. This will be the first time all recognized species will be included in a phylogenic context. 302 DEAGAMIS — Abstracts Who”s who — The Specific status of Leiolepis belliana and Leiolepis reevesi Jesse L. GRISMER Villanova University, Department of Biology, 800 Lancaster Ave, Villanova, PA, 19085, USA; Email: jesse.grismer@villanova.edu. The taxonomy of the lizard genus Leiolepis has long been problematic. Much of this is seeded in the high degree of morphological variation in the two most wide spread species, Leiolepis belliana and L. reevesi. To date all studies in- volving these species have use specimens from the distant ends of their distribution adding to the evidence that these species are distinct from one another. Through recent collection efforts at the contact zone of these species in south- ern Indochina, has produced morphological data suggesting that these two species are conspecfic and morphological- ly grade into one another. A phylogenetic analysis of mitochondrial and nuclear DNA was preformed to test the hy- pothesis of these two species representing one species. This would indicate that all other species of Leiolepis have arisen from one wide-ranging species. Australian Agamid Lizards: an overview of species diversity, biogeography and evolutionary relationships Jane MELVILLE Museum Victoria, GPO Box 666, Melbourne, VIC 3001, Australia; Email: jmelv@museum.vic.gov.au. Approximately 70 species of Agamid lizards are currently recognised in Australia and are known as the sub-family Amphibolurinae, which is genetically distinct from other agamid clades. Over the last decade we have undertaken a revision of the systematics of this subfamily, using molecular and morphological techniques. We have completed DNA sequencing of more than 2000 individuals for mtDNA and the nuclear region RAGI. I will be presenting an overview of these results. We have found that species diversity is underestimated in the current taxonomy, particularly in the genera Diporiphora and Tvmpanocryptis. In addition, these two genera show distinct geographic centres of species diversity, with multiple sympatric species — Diporiphora in north-western Australia and Tympanocryptis in north-east- ern South Australia. The genetic structuring across arid and semi-arid Australia is highly complex in the genera Di- poriphora, Tympanocryptis, Amphibolurus and Lophognathus, with greater levels of genetic divergence than expect- ed. Our results will provide important insight into evolutionary patterns, historical biogeography and speciation in arid and semi-arid Australia. Viviparity in the family Agamidae Scott M. MooDYy Department of Biological Science, Irvine Hall, Ohio University, Athens, OH, 45701, USA; Email: moody(wohio.edu. When MoobY (1980) reviewed reproductive mode within the family Agamidae only two clades were known to be vi- viparous. Cophotis ceylanica is viviparous although the other two closely-related genera (Lvriocephalus and Ceratopho- ra) have oviparous species. The genus Phrynocephalus has several viviparous species although others are oviparous. Surprisingly none of the desert-adapted amphibolurines of Australia are viviparous. The family Chamaeleontidae has both oviparous and viviparous clades and since this family is arguably embedded cladistically within the family Agami- dae sensu lato, the paucity of viviparity within the large radiation of agamids (Draconinae Moopy) in southeastern Asia is enigmatic. I was pleasantly surprised when Bjorn Lardner recently sent to me photographs of a female agamıd with a full-term fetus collected in Borneo. Thus a third case of viviparity has been documented Hylagama borneen- sis MERTENS. I will review the evolution of viviparity within the agamids in light of ecological and physiological adap- tations and restraints, and speculate as to why viviparity has not evolved more frequently within the Agamidae. Bonner zoologische Beiträge 56 303 POSTER PRESENTATIONS On Systematics and phylogeography of sun watcher toad agamas Phryno- cephalus helioscopus (PALLAS, 1771) Daniel MELNIKOV!, Natalia B. ANANJEVA!, Mehdi RAJABIZADEH? & Konstantin MILTO! !'Zoological Institute, Russian Academy of Sciences, St. Petersburg, Russia. 2Department of Biology, Faculty of Science, Shahid Beheshti University, Evin, Tehran, Iran. Results of our study on historical background and taxonomic status of two Iranian-Transcaucasian forms — Phyno- cephalus persicus DEFILIPPL, 1863 and Phynocephalus helioscopus horvathi MEHELY, 1894 was showed recently (MEL- NIKOV et al. 2008). Results of our current study on toad agamas of helioscopus-group from whole species range with the same mitochondrial marker (ND2, 700 bp) and its congruence with morphology are presented. Two main lineage — Iranian-Transcaucasian and Turanian — were detected on molecular analysis data. Specimens from southernmost lo- cality Abadeh (Southern Iran, Zagros mountains) occupy basal position in Iranıan-Transcaucasian lineage. Apical po- sitions in this lineage have specimens from Saveh (Central Iran, type territory of Ph. persicus) and specimens from Aralik (Turkey, type territory of Ph. h. horvathi), Armavir (Armenia), lughli (north-western Iran). These two forms also have good morphological differences in distance between nostrils and L/Led ratio (Melnikov et al., 2008). Con- trary to our previous opinion about horváthi as a form of Turanian lineage, now we have strongly supported data that it belongs to Iranian-Transcaucasian lineage. Basal position of agamas from Charyn and Taskarasu (south-eastern Kaza- khstan, left bank of Ili River, near to type territory of Ph. h. cameranoi BEDRIAGA, 1907) is most surprising in Turan- ian lineage. Other lizards from the south-eastern Kazakhstan (Otar, Burylbaital, Ay, Panfilov) are clustered with aga- mas from Fuyun (western China, type territory of Ph. varius Eichwald, 1831) and occupy apical position in this lin- eage. Specimens from Baskunchak, Tautobe (south-eastern Russia, Astrakhan region) and Akespe, Kamyshlibash (west- ern Kazakhstan, Aral Sea) have another apical position in this lineage. Type territory of Ph. helioscopus (PALLAS, 1771) is situated between these two areas. Phylogenetic positions of specimens from Kagan and Tamdy, Eddum (southern Uzbekistan) are not clear because of scanty material. Agamas from supported clades demonstrate some difference in morphology. Ph. h. helioscopus specimens have jet-black tip of tail, that easy to distinguish even on old collection material. Coloration of lower surface of tail end in males Ph. h. helioscopus is ensanguined, in Ph. h. varius 1s vi- nous, in animals from Charyn and Taskarasu 1s orange-red. This red coloration 1s fast to disappear in collection spec- imens. Morphological features suitable for identification of collection material are working out. Distribution patterns of agamids in the North Caspian region Konstantin D. MILTO Department of Herpetology, Zoological Institute, Russian Academy of Sciences, St. Petersburg, Russia; Email: coluber@zin.ru. Five species of agamid lizards (Trapelus sanguinolentus, Phrynocephalus mystaceus, Phrynocephalus helioscopus, Phrynocephalus guttatus, Phrynocephalus interscapularis) occur in the North Caspian Region. Phrynocephalus mys- taceus and Phrynocephalus guttatus are psammophilous species and they demonstrate a wide north-caspian distribu- tion in the Caspian Depression reaching the Ergeni Hills in the west and Terek-Kuma Sands in the south. The north- ernmost agamid species, Phr. guttatus, is distributed in the Sarpa Depression in the north. The sclerophillic Phr. he- lioscopus occupies only eastern part of the Caspian Depression from the Western Kazakhstan in the east to the Vol- ga River in the west. In the European part of the range this species is distributed sporadically and very rare. 7! san- guinolentus demonstrates Caucaso-Middle-Asian range disjunction and inhabits the Terek-Kuma Sands in the west and eastern extremity of the Caspian Depression in the east. Phr. interscapularis is represented by an isolated relic population in the Caspian Karakum Desert. Three types of distributional patterns could be distinguished for agamids in this region: 304 DEAGAMIS — Abstracts l. continiuous north-caspian distribution (Phr. mystaceus, Phr. guttatus), 2. disjunctive north-caspian distribution (7. sanguinolentus), 3. relic north-caspian distribution (Phr. helioscopus, Phr. interscapularis). Phr. interscapularis ıs an endemic species of the Middle Asian and the East Iranian deserts. 7. sanguinolentus, Phr. helioscopus and Phr. mystaceus have Middle-Asian ranges and were described as a subendemic species of the Mid- dle Asia and the Eastern Iran. The wide distributed in the Middle and Central Asia, Phr. guttatus has possibly Cen- tral-Asian origin. Two species in the North Caspian Region are represented by Turanian subspecies (Phr. h. heliosco- pus and Phr. i. interscapularis). A subspecific endemism are known for three species of agamids in the North Caspi- an Region. Trapelus sanguinolentus, Phrynocephalus mystaceus and Phrynocephalus guttatus are represented here by endemic subspecies (7. s. sanguinolentus, Phr. m. mystaceus, Phr. g. kalmykus). In the territory of the Middle Asia these species are represented by Turanian subspecies (7. s. aralensis, Phr. m. galli, Phr. g. guttatus). This work was supported by the grant of Scientific School N° 4212.2006.4. Phylogeography « taxonomy of the Agamid lizards (Sauria: Agamidae) of East Africa: morphological and genetic analysis Alexander BURMANN, Philipp WAGNER, Bernhard Misor, Martin HAASE & Wolfgang BÖHME Zoologisches Forschungsmuseum Alexander Koenig, Adenauerallee 160, D-53113 Bonn, Germany. The East African members of the 4eama lionotus complex are analyzed. The results clearly show differences in mor- phology and colouration between the 4. lionotus subspecies. Especially the throat of adult males is a characteristic to determine the different species and subspecies. Buchbesprechung 305 MANTHEY, Ulrich (2008). Terralog Vol. 7a — Agamid Lizards of Southern Asia, Draconinae 1. Edition Chimaira, Frankfurt am Main / Rodgau. 160 S., Hardcover. ISBN 978-3-89973-357-0. This first part of the volume 7 of TERRALOG portrays 95 agamid species of the subfamily Draconinae from Asia. All species are roughly described with pictograms only (without text) and with a distribution map including the type locality. This book contains the most beautiful pho- to collection of agamid lizards of southern Asia that I have ever seen. Usually adult males, females and juveniles are figured for all species (often from more than one locali- ty), totalling in more than 530 colour figures. The book not only pays deference to the tremendous colour varia- tion of individuals in some of the treated species, but al- so to the great species diversity of this subfamily (species lacking in this book will be published in the second part of the volume). Additionally, for some species also the holotype or the habitats are illustrated. Most of the images are from the author himself but several other well-known agamid reserchers contributed images, e.g. J. McGuire, L. L. Grismer, W. Bóhme, T. Ziegler or N. Orlov. In the introduction the genera and some of the species are characterized (though not very exhaustive) and in some cases detailed information about the current taxonomic sta- tus (e.g. Draco abbreviatus) ıs considered as valid, or the distribution are given. These data include new findings, e.g. a first record of Japalura tricarinata for Tibet. The introduction includes also a part “How to use the book’, some information on husbandry of species and a list of bibliographical references. The text comes bilingual in English and German. Summarizing, the book is not a taxonomic, ecological or naturalists’ guide, but a perfect photographic account for professional and amateur herpetologists who are interest- ed in the family Agamidae in general, or more specifical- ly in the subfamily Draconinae. of Southern Asia Draconinae 1 Philipp WAGNER Zoologisches Forschungsmuseum Alexander Koenig, Bonn CALL FOR PAPERS International Symposium August 16-20, 2010 Zoological Institut, St. Petersburg DEAGAMIS 2nd International Symposium on Agamid Lizards Topics: O Taxonomy, phylogeny & biogeography O Population genetics € reproductive modes O Physiology & ecology O Life history O Behaviour ® Conservation Congress language: English Key note: lectures 45 min or 20 min (Discussion time included) Prof. Dr. Natalia Ananjeva Philipp Wagner Head of Division of Herpetology Zoologisches Forschungsmuseum A. 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Otherwise, suggest another journal. — Is it substantial and novel enough to justify publication? — Would the reader easily grasp the significance of the work from the introduction and discussion? — Do the conclusions follow from the information presented? — Can the paper be made clearer and more concise? — Could the title be shortened or made more informative’ — Does the abstract indicate the methods used and the important re- sults? — Are alternative interpretations sufficiently considered? — Are contents and length of the paper well balanced? — Are there any important references to the subject that have not been considered? — Are all tables and illustrations necessary? — Are the tables and figures sufficient? SMITHSONIAN INSTITUTION LIBRARIES NENA, 3 9088 01533 1598 Special Issue: On Agamid Lizards “DEAGAMIS — 1% International Symposium on Agamid Lizards’ Preface 211 Editorial 213 WAGNER, Philipp; WILMS, Thomas M.; BAUER, Aaron & BÖHME, Wolfgang: PANS) Studies on African Agama V. On the origin of Lacerta agama Linnaeus, 1758 (Squamata: Agamidae) SANDERA, Martin & STAROSTOVA, Zuzana: 225 A record of Asian agama of the genus Calotes Cuvier, 1817 (Squamata: Agamidae) in Kenya ISARUNARATHNA, D. M. S. Suranjan; AMARASINGHE, A. A. Thasun & STÓCKLI, Edi: 229 Taxonomic and biological study on Calotes ceylonensis Müller, 1887 (Reptilia: Agamidae) of Sri Lanka WAGNER, Philipp; INEICH, Ivan; LEACHE, Adam D.; WıLms, Thomas M.; TRAPE, Sébastien; 239 BÖHME, Wolfgang & SCHMITZ, Andreas: Studies on African Agama VI. Taxonomic status of the West African Agama (Sauria: Agamidae) with prominent tail crests: Agama boulengeri Lataste 1886, Agama insularis Chabanaud, 1918 and Agama cristata Mocquard, 1905 DENZER, Wolfgang & MANTHEY, Ulrich: 255 Remarks on the type specimen of Gonocephalus mjobergi Smith, 1925 (Sauria: Agamidae) WiLms, Thomas M.; WAGNER, Philipp; SHOBRAK, Mohammed & BOHME, Wolfgang: 259 Activity profiles, habitat selection and seasonality of body weight in a population of Arabian Spiny-tailed Lizards (Uromastyx aegyptia microlepis Blanford, 1875; Sauria: Agamidae) in Saudi Arabia Go LEACHE, Adam D.; CHONG, Rebecca A.; PAPENFUSS, Theodore J.; WAGNER, Philipp; Dif. BÖHME, Wolfgang; SCHMITZ, Andreas; RODEL, Mark-Oliver; LEBRETON, Matthew; INEICH, Ivan; CHIRIO, Laurent; BAUER, Aaron; ENIANG, Edem A. & BAHA EL Din, Sherif: Phylogeny of the genus Agama based on mitochondrial DNA sequence data HALLERMANN, Jakob: 279 A new species of Bronchocela (Squamata: Agamidae) from Nicobar Island WAGNER, Philipp; BAREJ, Michael E & SCHMITZ, Andreas: 285 Studies on African Agama VII. A new species of the Agama agama-group (Linnaeus, 1758) (Sauria: Agamidae) from Cameroon & Gabon, with comments on Agama mehehii Tornier, 1902 Appendix: Abstracts of “DEAGAMIS — 1% International Symposium on Agamid Lizards’ 299 Buchbesprechungen / Book Reviews MANTHEY, Ulrich (2008) 305 Terralog Vol. 7a— Agamid Lizards of Southern Asia, Draconinae 1 (P. WAGNER) Titelbild/Cover illustration: Agama planiceps Peters, 1862 from Namibia. Photo by Dieter M. 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