Bonn Vomime 57 Issue 2 zoological 010 Bulletin formerly: Bonner zoologische Beitrage Herpetologia Koenigiana Festschrift in honour of Prof. Wolfgang BOhme Guest editor: Philipp Wagner An open access journal of organismal zoology,.published by Zoologisches Forschungsmuseum Alexander Koenig, Bonn Bonn zoological Bulletin (BzB), formerly “Bonner zoologische Beitrage”, is published by the Zoologisches Forschungsmuseum Alexander Koenig (ZFMK), Bonn. Two regular issues of BzB are published per year; supplements on focus topics are produced in irregular succession. Subscription price is 46 € per volume (year), including supplements and shipping costs. For subscription, back issues and insti- tutional exchange, please contact the ZFMK library (ZFMK, Bibliothek, Frau Diane Steinebach, Adenauerallee 160, D-53113 Bonn, Germany, tel. +49 228-9122-216, fax: +49 228-9122-212; d.steinebach.zfmk@uni-bonn.de). The online version of BzB is avail- able free of charge at the ZFMK homepage: http://www.zfmk.de/web/Forschung/Buecher/Beitraege/index.en.html. © 2010 Zoologisches Forschungsmuseum Alexander Koenig, Bonn, Germany. ISSN 2190-7307 Produced by Eva-Maria Levermann, Kaiserstr. 129, D-53113 Bonn, Germany; emlevermann@netcologne.de. Printed and bound by DCM, Werner-von-Siemens-Str. 13, D-53340 Meckenheim, Germany. Bonn zoological Bulletin Editor-in-Chief Fabian Herder, Zoologisches Forschungsmuseum Alexander Koenig (ZFMK), Ichthyology Section, Adenauerallee 160, 53113 Bonn, Germany, tel. +49 228-9122-255, fax: +49 228-9122-212; f.herder.zfmk@uni-bonn.de Guest Editor Philipp Wagner, Zoologisches Forschungsmuseum Alexan- der Koenig (ZFMK), Herpetology Section, Adenauerallee 160, 53113 Bonn, Germany, tel. +49 228-9122-254, fax: +49 228-9122-212; philipp.wagner.zfmk@uni-bonn.de Editorial Board Dirk Ahrens, Insects: Coleoptera, ZFMK, tel. +49 2289122286, fax: +49 228-9122-332: d.ahrens.zfmk@uni-bonn.de Wolfgang Bohme, Amphibians and Reptiles, ZFMK, tel. +49 228-9122-—250, fax: +49 228-9122-212; w.boehme.zfmk@uni-bonn.de Netta Dorchin, Insects: Diptera, ZFMK, tel. +49 228-9122-292, fax: +49 228-9122-212; n.dorchin.zfmk@uni-bonn.de Renate van den Elzen, Birds, ZFMK, tel. +49 228-9122-231, fax: +49 2289122212; r.elzen.zfmk@uni-bonn.de Bernhard Huber, Invertebrates except Insects, ZFMK, tel. +49 2289122294, fax: +49 228-9122-212; b.huber.zfmk@uni-bonn.de Rainer Hutterer, Mammals, ZFMK, tel. +49 228-9122—261, fax: +49 228-9122-212; r.hutterer.zfmk(@uni-bonn.de Gustav Peters, Mammals, ZFMK, tel. +49 2289122262, fax: +49 2289122212: g.peters.zfmk@uni-bonn.de Bradley Sinclair, Canadian National Collection of Insects, Ottawa Plant Laboratory — Entomology, CFIA, K.W. Neat- by Bldg., C.E.F., 960 Carling Ave., Ottawa, ON, Canada K1A 0C6, tel. + 1 613-759-1787, fax: + 1 613-759-1927; bradley.sinclair@inspection.ge.ca Dieter Stiining, Insects except Coleoptera and Diptera, ZFMK, tel. +49 228-9122-—220, fax: +49 228-9122-212; d.stuening.zfmk@uni-bonn.de Advisory Board Theo C. M. Bakker, Rheinische Friedrich-Wilhelms-Univer- sitat, Institut fiir Evolutionsbiologie & Okologie, 53113 Bonn, Germany, tel. +49 228—73-5 130, fax: +49 228-73- 2321; t.bakker@uni-bonn.de Aaron M. Bauer, Villanova University, Department of Biolo- gy, 800 Lancaster Avenue, Villanova, PA 19085-1699, USA, tel. +1 610-519-4857, fax: +1 610-519-7863; aa- ron.bauer@villanova.edu Wieslaw Bogdanowicz, Museum and Institute of Zoology, Polish Academy of Sciences, Wilcza 64, 00-679 Warszawa, Poland, tel. +48 22628-7304, fax: +48 22629-6302; wieslawb@miiz.waw.pl Matthias Glaubrecht, Museum fiir Naturkunde Berlin, Leib- niz-Institut fiir Evolutions- und Biodiversitatsforschung an der Humboldt-Universitat zu Berlin, Invalidenstrasse 43, 10115 Berlin, Germany, tel. +49 30-2093-8504/ 8400, fax: +49 030-—2093-8565; matthias. glaubrecht@mfn-berlin.de Jeremy D. Holloway, The Natural History Museum, Depart- ment of Entomology, Cromwell Road, London, SW7 5BD, U.K.; j-holloway@nhm.ac.uk Boris KryStufek, Slovenian Museum of Natural History, P. O. Box 290, Ljubljana, Slovenia; boris.krystufek@zrs.upr.si Wolfgang Schawaller, Staatliches Museum ftir Naturkunde, Rosenstein 1, 70191 Stuttgart, Germany, tel. +49 711-8936-221, fax: +49 711-8936-100; schawaller.smns@naturkundemuseum-bw.de Ulrich K. Schliewen, Department of Ichthyology, Bavarian State Collection of Zoology, Minchhausenstr. 21, 81247 Munchen, Germany, tel. + 49 89-8107—110; schliewen@zsm.mwn.de Michael Schmitt, Ernst-Moritz-Arndt-Universitat, Allge- meine & Systematische Zoologie, Anklamer Str. 20, 7489 Greifswald, Germany, tel. +49 3834864242, fax: +49 3834 86-4098: michael.schmitt@uni-greifswald.de W. David Sissom, Dept. of Life, Earth and Environmental Sciences, W. Texas A. & M. University, WITAMU Box 60808, Canyon, Texas 79016, USA, tel. +1 806-651-2578, fax: +1 806-651-2928; dsissom@mail.wtamu.edu Miguel Vences, Technical University of Braunschweig, Zoo- logical Institute, Spielmannstr. 8, 38106 Braunschweig, Germany, tel. + 49 531-391-3237, fax: + 49 531-391-8198; m.vences@tu-bs.de Erich Weber, Eberhard-Karls-Universitat, Zoologische Schausammlung, Sigwartstr. 3, 72076 Tubingen, tel. +49 70712972616, fax +49 7071—295170; erich.weber@uni-tuebingen.de Editorial On occasion of his 66th birthday and his retirement from his position as curator of herpetology after nearly 40 years at the Museum Alexander Koenig, I had the great pleas- ure to be the editor of a Festschrift, and the co-ordinator of a colloquium honouring the work of Wolfgang Bohme. The title of this special issue of the Bonn zoological Bul- letin and the colloquium is a combination of Herpetolo- gia, referring to several symposia (e.g. the two SEH meet- ings) he organized during his time at this institute, and Koenigiana, referring to ‘his’ museum. During his time at curator from 1971 onwards, he in- creased the at this time ‘sleeping’ herpetological collec- tion with less than 9500 specimens to one of the leading collections in Germany and Europe with about 100.000 vouchers. His scientific work 1s documented in more than 530 publications and with his enthusiasm he influenced and stimulated 32 PhD (ten more candidates are still work- ing on their theses), 145 diploma and 35 state-examen stu- dents, some of them working today as scientists in renowned German and European institutions. As his student, I was influenced since my first time at the University of Bonn and the Museum Koenig by his cours- es and lectures and he positively forced my decision to work with herpetological systematics and the Afrotropi- cal region. These influences Wolfgang B6hme had on many upcom- ing scientists and his numerous cooperations are in part reflected by the impressive set of articles about various topics published in the present issue of Bonn zoological Bulletin, written by friends, colleagues and former stu- dents. These articles are partly dealing with his main sci- entific fields of interests: Africa, chameleons, or monitor lizards, but also with ‘side’-fields like lacertids, nomen- clature or the history of herpetology. Many articles are first descriptions of new amphibian and reptile species named after Wolfgang Bohme, which are the ‘living manifesta- tions’ of his impressive work. Moreover, the two non-her- petological articles show that he has also some impact in other zoological disciplines. As guest editor, I would like to thank all the reviewers for their helpful comments. Without their fast work it would not have been possible to have this issue ready right in time. Especially, I would like to thank Brian Sinclair who did the English in review in some of the articles. Pe ladear Philipp Wagner (Guest Editor) Bonn, November 2010 Pe 0a PGP TER eit ph | iter), Saas pines ‘ice . Me wn Sa Leth Age oe Wy | ne es ae ee a Co 5 ‘Morita: ine eT TR pasate ll RE ea ree ae td es ha bby = Mas Preface This volume is dedicated to a scientist, who during the past four decades transformed the herpetology collection of Museum Koenig into one of the most prominent European research facilities for reptile systematics and biology. Un- der his management, the comparatively young collection of reptiles and amphibians in Bonn has grown tenfold and became one of the most important ones in Germany. More- over, he dedicated his time and energy not only to science, but also to the development of the institute, which has been his second home. Thanks to his commitment and person- ality, he became the undisputed head of the vertebrate de- partment, which he still leads today. He also was, and still is, the link between the museum and the many experts among laypersons, who contribute substantially to the growth of scientific knowledge and otherwise shy from direct contact with professionals. With the same spirit he cooperated with the Alexander Koenig Gesellschaft, the friends of the museum society. Wolfgang Bohme is a passionate zoologist whose enthu- siasm infected several generations of students, many of which later became successful scientists themselves. He regularly takes students on field courses, where he teach- es them how to discover, observe and capture snakes, lizards and frogs, as well as identify other animals such as grasshoppers and bees. His favourite destination for this purpose has always been Lake Neusiedl that became a popular destination among biology students in Bonn. Wolfgang Bohme frequently invited like-minded zoolo- gists from other countries to work with him in Bonn or to attend German conferences, and it was obvious that he was thrilled for the opportunity to learn of other people’s discoveries out of pure fascination by animals and with- out any jealousy. During the past 40 years, the Museum’s directors were grateful for his public-relations work. He frequently of- fered evening lectures for the general public with an en- joyable combination of adventure, discovery, and BOhme’s characteristic humour. He authored popular articles for the media and for the institute’s series “Tier und Museum” (later “Koenigiana’’), a publication for which he acted as an editor. Similarly, he was always ready to take part in non-scientific events, where he participated in activities such as reading poems by his ancestor Wilhelm Busch, whose funny bone he obviously inherited. With this volume on the occasion of his retirement from official duties, we want to thank him for his outstanding contribution to herpetology and to the development of Mu- seum Koenig. I hope this is just the beginning of a new period of active research, for which we will always have a space available for him in our institute. J. Wolfgang Wagele (Director) Bonn, November 2010 j Sidaniteite ‘ sine we ak i bia ety Nl Se) wee ey wo Wi Te . Ls “ ie ahi te F eetlornty ma) ith ay A ant ie 10 Greetings from the SEH I met Wolfgang Bohme for the first time at the very be- ginning of my carrier, just after my graduation. I remem- ber that he was extremely friendly when talking about lizards, the Mediterranean basin and his deep love for Africa and the tropics, making my first impact with the international professional herpetological world easy and smooth. Throughout our long professional relationship, Wolfgang has always been extraordinarily open to new ideas, sharing research projects with a number of scien- tists worldwide and prompting the creation of a network of enthusiastic believers, such as for the circum-Mediter- ranean Podarcis fans. | remember very well the begin- ning of the “saga” of the Symposia on the Lacertids of the Mediterranean, together with Nicholas E. Arnold, Massi- mo Capula, Valentin Pérez-Mellado, Efstratios Valakos and, which 1s still successfully ongoing. In the nineties, as President of the Societas Europaea Herpetologica, Wolfgang Bohme put a tremendous effort into establish- ing collaborations between the national European herpeto- logical societies. He also opened the doors of the herpeto- logical collection of the Alexander Koenig Museum to a multitude of island zoologists, inducing the full scientif- ic exploitation of such an extraordinary biological treas- ure. His outstanding scientific activity, together with his humanity and intellectual generosity make Wolfgang Bohme one of the reference point in and out my profes- sional life and Ill always be grateful to him and his love- ly family for such a warm and positive friendship. Claudia Corti (President Societas Europaea Herpetologica) Bonn, November 2010 ar Wee ee ij ys Tete eh iy eae ; “a aii oi nin wi on ees Rover See ie} it iti: aay, wih ig oe iowoaanladi dota sal poenllle, - ry) eaten) at bidil OE | ned tis Ae dna li weit hah @ aa Salat Hiriay i ribs rie an Aly AD quiere fey / hens & wean Ty LD ET EC eo Greetings from the DGHT On the occasion of the retirement of Prof. Dr. Wolfgang Bohme, curator of herpetology, head of the vertebrate de- partment, and deputy director of the Zoologisches Forschungsmuseum Alexander Koenig, we were honoured with an invitation to direct a few words to the readers of this special issue of the Bonn Zoological Bulletin. We sincerely appreciate this opportunity, as the name Wolfgang Bohme is not only connected to a distinguished career in herpetology, but also to a person who always made an effort to keep a close connection between pro- fessional scientific research and amphibian and reptile am- ateurs all over the world. Wolfgang Bohme’s achievements in this particular field of interaction are not less than out- standing. There is probably no other herpetologist whose research benefited as much from data supplied by ama- teurs, and there are countless thankful amateurs whose pastime has been deeply enriched by the scientific advice and the stimulating attitude of Wolfgang Bohme. Sharing data, knowledge and specimens with people operating out- side the scientific community was paramount for many spectacular discoveries and the furtherance of knowledge about the biology and ecology of amphibian and reptile species, many of which are considered to be well-known today. Decades ago, it was mainly these amateur enthu- siasts who travelled to remote and previously unexplored exotic areas in search of their “pets” and returned with treasures of data and photographs that gave rise to subse- quent research projects. It is Wolfgang Bohme’s merit and strength to have fostered and encouraged this tradition of interaction for several decades. Bringing professionals and amateurs together and promot- ing their dialog is also one main characteristic of the Deutsche Gesellschaft fiir Herpetologie und Ter- rarienkunde (DGHT), as is apparent by the society’s name. As a consequence, there was, and is, some kind of “nat- ural relationship” between Wolfgang Bohme and the DGHT, and this relationship has been a sustained and fruit- ful one. Shortly after he took his position at the Museum Koenig, he founded the “DGHT Stadtgruppe Bonn” in 1973. Later, Wolfgang B6hme became president of the so- ciety and held this position from 1983 to 1991. He was furthermore a founding member of the DGHT Work Group Literature and History, and a regular member of the Work Groups Chameleons and Lacertids. In 1994, Bohme was made an honorary member of the DGHT, and very recently, the current executive board asked him to join its advisory council. And so it is for many reasons that Bohme’s name and personality are closely interlinked with the DGHT. The DGHT as a society, DGHT members and associates, as well as many students and colleagues owe a lot to Wolf- gang Bohme. Despite facing his ‘official’ retirement as a herpetologist now, we are convinced that few things will change and that his fruitful relationships with the DGHT will continue for many years to come. Dear Wolfgang Bohme, thank you for everything. The DGHT executive board (Peter Buchert, Jorn Kohler, Axel Kwet, Stefan Lotters, Wolfgang Schmidt, Holger Vetter) Bonn, November 2010 it i (Aon w da K cy wile Ws 4 1a a \ Wyn my a ni anil teh ap |) a Toke vubeunite ry byt if eis Me, eligte’ by A aie wud rn i tie sal Curriculum Vitae of Wolfgang Bohme Born November 21, 1944 at Schdnberg near Kiel (Schleswig-Holstein, Germany) as the 3rd child of the mu- sician Ferdinand B6hme (1906-1971) and his wife Hed- wig, born Stange (1913-1992). Ferdinand Bohme was vi- olinist of the first desk at the municipal orchestra in the opera house of Kiel and subject teacher for violine, both privately and at the Pedagogic College of Kiel. From a first marriage (1970-1974) father of one daugh- ter (Judith). Since 1974 married with Roswitha Bohme. From this marriage two sons (Moritz and Peter). Wolfgang Bohme finished highschool (“Kieler Gelehrten- schule”’) in Kiel, April 1965, and subsequently studied zo- ology, botany and paleontology at the “Christian-Al- brechts-Universitat” of Kiel. Doctoral degree with a the- sis on hemipenis morphology in lacertid lizards in June 1971 under supervision of Prof. Dr. Wolf Herre. From August 1971 until December 2010 head of the Her- petology Section of the “Zoologisches Forschungsmuse- um Alexander Koenig” (ZFMK) in Bonn, Germany. Since then, increase of the herpetological collection from less than ca. 9.600 specimens to ca. 100.000. From October 1992 to his retirement Head of the Vertebrate Department and Vice Director of the Forschungsmuseum. Since winter semester 1980/81 participation in teaching at the “Rheinische Friedrich-Wilhelms-Universitat” Bonn; habilitation (thesis on genital morphology in the Sauria) and venia legendi received in May 1988. Since then reg- ular teaching and supervision of more than 129 master and 32 doctoral theses, plus 35 theses for state examen. Award- ed full professorship (“‘apl. Professor’) April 1996. Fields of Research. Systematics, ecology and biogeog- raphy of amphibians and reptiles, with taxonomic focus on lizards (chameleons, monitor lizards) and its Tertiary and Quaternary predecessors (e.g. lizard amber fossils); genital morphology of squamates. Geographical focus on the western Palearctic Region (founder and editor of “Handbuch der Reptilien und Amphibien Europas’”, edi- tion of 5 vols. between 1981 and 1999) and West Africa (six larger field excursions to West/Central Africa, each with 1—3 months duration: Spring 1973 Cameroon, win- ter 1973/74 Cameroon, winter 1975/76 Senegal/Gambia, 1993 Guinea, 1998 Cameroon, 1999 crossing the Sahara in both directions: Morocco, West Sahara, Mauritania, Senegal). In 2002 an extensive educational trip through the eastern half of the USA. Currently 530 sci. publica- tions. Working group. Members of his ZFMK herpetological working group were/are active in the tropics of Central and South America (Costa Rica, Venezuela, Peru, Bolivia, Chile), Africa (Guinea Bissau, Gambia, Benin, Cameroon, Gabon, Kenya, Zambia) and Madagascar, and of SE Asia (Vietnam, Indonesia). Societies. September 1979 host and founding member of the “Societas Europaea Herpetologica”, which publishes “Amphibia-Reptilia’, simultaneously founded at ZFMK, and today the leading journal of its discipline in Europe. Elected for president of the society 1993 in Barcelona, re- elected for another 4 years 1997 in Prague. From 1983 to 1991 Chairman of the “Deutsche Gesellschaft fiir Herpetologie und Terrarienkunde” (DGHT) which during this period grew from ca. 2000 to over 5000 members, thus becoming the largest associa- tion of its discipline in the world. Wolfgang Bohme has the Honorary membership of the DGHT since 1994. From 2000 to 2002 founding chairman of the working group “Literatur und Geschichte der Herpetologie” of the DGHT with its own periodical “Beitrage zur Literatur und Geschichte der Herpetologie und Terrarienkunde’”, after two issues, from 2003 onwards, continued as “Sekretar”. From 2001 to 2005 member of the Commission of the “In- ternational Committee of Zoological Nomenclature” (ICZN). From October 1996 to December 2006 President of the “European Association of Zoological Nomenclature” (EAZN). Elected as ‘Honorary foreign member’ of the American Society of Ichthyologists and Herpetologists in 2008. Awarded with the Alexander Koenig Medal in 2010, by the Alexander-Koenig-Gesellschaft, sponsoring society of ZFMK. Ph. Wagar Philipp Wagner (Guest Editor) Bonn, November 2010 eT Liars War Lah F a wu ; See eal al fay POLED | i ; Pal wed rie a he be) 2a} tan SelveW eet “nai ire \, 4 be rv 7, v ia ave votive : Poe MY — ; ay ste a 1 ene peo sis anti Bonn zoological Bulletin Volume 57 Issue 2 pp. 111-118 Bonn, November 2010 Sharing resources in a tiny Mediterranean island? Comparative diets of Chalcides ocellatus and Podarcis filfolensis in Lampione Miguel A. Carretero!*, Pietro Lo Cascio, Claudia Corti3.! & Salvatore Pasta‘ 'CIBIO, Centro de Investigag¢ao em Biodiversidade e Recursos Genéticos, Campus Agrario de Vairao, P-4485-661 Vairao, Portugal; E-mail: carretero@mail.icav.up.pt 2Associazione “Nesos”, Via Vittorio Emanuele 24, 98055 Lipari (ME), Italy; E-mail: plocascio@nesos.org. 3Museo di Storia Naturale dell’ Universita di Firenze, Sezione di Zoologia “La Specola”, Via Romana 17, I1-50125, Firenze, Italy; E-mail: claudia.corti@unifi.it 4via V.F. 19, 60/A, I-90126, Palermo, Italy; E-mail: salvatore.pasta@alice.it *corresponding author: E-mail: carretero@mail.icav.up.pt tel.+351252660400 fax.+351252661780 Abstract. The insular lizard microcommunity inhabiting the Lampione islet (Pelagian islands, S Italy) is constituted by a skink (Chacides ocellatus) and a lacertid lizard (Podarcis filfolensis). Their diet composition (taxa and sizes) during spring-early summer were analysed based on 131 faecal pellets, which could be individually assigned to a lizard species and sex (only in P. filfolensis). The diet of C. ocellatus was biased towards hard prey of medium to large sizes (Coleoptera, insect larvae). Podarcis filfolensis displayed a more diverse prey spectrum including Formicidae, Coleoptera, insect lar- vae and minor prey not consumed by the skink, but restricted to the small items; differences between sexes were mini- mal. Both species were partially herbivorous. Evidence of cannibalism was found for P. filfolensis and C. ocellatus preyed upon P. filfolensis. Pseudocommunity analysis does not support community structure but instead points to convergence in trophic strategies between both species due to insular conditions. Evolutionary history, rather than resource partition- ing, seems responsible for the moderate trophic overlaps recorded and even may explain why both species coexist under the harsh conditions of this tiny islet. Keywords. Diet; Chacides ocellatus; Podarcis filfolensis; community ecology; islands; Lampione. INTRODUCTION For decades, lizards have constituted fruitful model organ- isms for studies in community ecology, diet being the most studied ecological dimension (see review by Luiselli 2008). Many of the initial and current studies are focused on the most complex assemblages, namely those in trop- ical or desert areas (Arnold 1984; Pianka 1986; Vitt & Caldwell 1994; Vitt & Carvalho 1995; Vitt & Zani 1998; Vitt et al. 2000; Akani et al. 2002, amongst many others), where environmental stability could allow interspecific re- lationships promoting detectable community structure (Winemiller & Pianka 1990). In contrast, studies on lizard assemblages inhabiting temperate regions are less abun- dant (but see Pérez-Mellado 1982; Strijbosch et al. 1989; Pollo & Pérez-Mellado 1991; Carretero & Llorente 1993; Capula & Luiselli 1994; Carretero et al. 2006; Kuranova et al. 2005; Rouag et al. 2007). This is probably because these are composed of less species but also because abi- otic restrictions of seasonal climates overcoming the role of species interactions would make community structure less expectable to appear (Barbault 1991). In fact, a Bonn zoological Bulletin 57 (2): 111-118 recent meta-analysis concluded that lizards of most (80%) communities worldwide do not partition their food re- sources but are randomly organised in the trophic niche axis (Luiselli 2008). Instead, increasing evidence is demonstrating that the influence of evolutionary history on the lizard trophic traits is stronger than previously thought. Specifically, niche conservatism rather than species interactions accounts for many trophic differences between the community components (Brooks & McLen- nan 2002; Webb et al. 2002; Vitt et al. 2003; Vitt & Pi- anka 2005; Mesquita et al. 2007; Espinoza et al. 2008). Within this context, lizards inhabiting small Mediterranean islands constitute an apparent paradigm of simplicity. On one hand, strong seasonality and impoverished trophic re- sources impose severe constraints to insular lizards (Pérez- Mellado & Corti 1993), higher than those in adjacent mainland, making lizard communities inhabiting Mediter- ranean islets extremely poor when compared to those on big islands or on the continent (Mylonas & Valakos 1990). ©OZFMK 112 Miguel A. Carretero et al. On the other hand, the exposition to less potential com- petitors and predators and subsequent increase in the con- specific density (Carretero 2004, 2006) may open new possibilities for enlarging trophic niche (Pérez-Mellado & Corti 1993; Carretero 2004). Nevertheless, evidence on lacertid lizards indicates that the ecological response to these shifted environmental pressures is not immediate and some evolutionary time is needed to develop profound trophic adaptations (Pérez-Mellado & Corti 1993; Car- retero 2004). Literature on the diet of Mediterranean lizards in small islands is abundant but usually focused on a single species (reviewed in Van Damme 1999; Pérez- Mellado & Traveset 1999; Carretero 2004), studies at mul- tispecies level being rare (Nouira 1983). Here, the diet composition of a microinsular community constituted by two divergent lizard species is analysed dur- ing spring-summer considering both inter- and intraspe- cific variation and compared to other populations of the same species. Moreover, the hypothesis of community structure at the trophic level is specifically tested against the null hypothesis of random trophic overlap. MATERIAL AND METHODS Study area Lampione (35°33’00”N—12°19’11”E) is a small islet lo- cated 17 km off the W coast of Lampedusa (Pelagian Is- lands) and 110 km off Tunisia, in the Channel of Sicily. The area is 0.021 km? and the maximum altitude is 36 m a.s.l. From a geological point of view, the islet is composed of dolomitised carbonates belonging to formations of the Tunisian offshore, and its definitive isolation from North Africa was since 18,000 years B.P. (Pasta 2002). The cli- mate is arid, characterised by strong drought periods in summer and by an average annual rainfall lower than 300 mm. The vegetation is mainly dominated by halo-ni- trophile perennial shrubs. The occurrence of a large colony of gulls causes a strong level of soil eutrophisation and nutrient imbalances, which allow the expansion of the ni- trophile biannual Malva veneta Soldano, Banfi & Galas- so, 2005 during the late spring on the top of the islet. Lam- pione is at present-day uninhabited, but late-Roman ru- ins document an early human presence, though probably only seasonal (Pasta & Masseti 2002). The invertebrate assemblage of the islet reflects several features typical of microinsular and arid environments, namely a low num- ber of species (about 30, excluding flying insects; Lo Cas- cio 2004, Lo Cascio unpublished), an over-representation of some groups (e.g., five species of Coleoptera Tenebri- onidae; Canzoneri 1972; Lo Cascio unpublished), some being found at extremely high densities. Bonn zoological Bulletin 57 (2): 111-118 Study lizards Two lizard species inhabit the islet: the Maltese wall lizard, Podarcis filfolensis (Bedriaga, 1876), (Squamata: Lacer- tidae) and the Ocellated skink, Chalcides ocellatus (Forskal, 1775) (Squamata: Scincidae). The first is a gen- uine insular species endemic to the Maltese Archipelago and two Pelagian islands, Linosa and Lampedusa, where it is said to be introduced in early or recent time (Capula 2006; Lo Cascio & Corti 2008). The Ocellated Skink, Chalcides ocellatus, is widely distributed on the Sindian- Mediterranean area and 1s recorded for all the Pelagian is- lands (Turrisi & Vaccaro 2006); the origin of the islet pop- ulation is probably related to the Pleistocene connections between Lampione and the nearby North-African main- land (see Grasso et al. 1985). The first data on the occur- rence of such species in Lampione were reported by Lan- za & Bruzzone (1961). Population density is extremely high for both species, only for Podarcis filfolensis being estimated using standard methods (7500-8000 individu- als/ha, see Lo Cascio et al. 2006). From field observations, the ratio of apparent abundance between this species and Chalcides ocellatus was 3:1 approximately (Lo Cascio un- published). Sampling and lab methods Field sampling was carried out during several visits in late spring/early summer of 2004 and 2005, when both species show the peak of annual activity (Corti & Lo Cascio 2002). Faecal pellets were obtained from adult Podarcis filfolensis and Chalcides ocellatus; all the specimens were measured (snout-vent length, SVL) to the nearest 0.1 mm using a digital calliper, sexed (in P. fi/folensis) and released back in the site of capture. Whereas adult P. filfolensis could be easily sexed in the field using sexual secondary characters (Corti & Lo Cascio 2002) and hemipenis ever- sion, the reduced external differences and the impossibil- ity for analysing of internal cloaca did not allow identi- fying the sexes of C. ocellatus in field (see Badir 1959; Capula & Luiselli 1994). The faecal contents were examined under stereoscopic mi- croscope (10-40X). Remains were identified to Opera- tional Taxonomy Units (OTUs) approximated here to the order/family level. Item counting was based on cephalic capsules, wings and legs, following the minimum num- bers criterion by sample. When possible, prey lengths were obtained measuring the remains with a micrometer eye- piece and calculated by using regression equations (H6- dar 1997) and then assigned to classes of 5 mm in length. ©OZFMK Diet of Lampione lizards 113 Statistical methods Three diet descriptors were used: the percentage of pel- lets containing an OTU (%P), the percentage of numeric abundance of each OTU (%N), and the use index (IU) (Jover 1989); the latter is preferred because combines %N and %P; the importance of a certain OTU in the diet be- ing estimated by calculation of the homogeneity of its con- sumption throughout all the individual contents (Carretero 2004). Brillouin’s index was used to estimate the diet di- versity according to Magurran (2004). For a given sam- ple, the average individual diversity (Hi) was obtained by averaging the diversity values of each pellet whereas the (asymptotic) population diversity (Hp) was estimated through jack-knife resampling (Jover 1989, Magurran, 2004), that is, recalculating the total diversity missing out each sample in turn and generating pseudovalues, which are normally distributed. Whereas Hi and Hp have stan- dard errors and allow statistical inference, the total accu- mulated diversity (Hz) of all pellets is a fixed value only provided for comparing with the literature (Ruiz and Jover 1981). Overlap between diets was evaluated by means of the Pi- anka’s index (Pianka 1973) applied on the IU values of OTUs and size classes (Carretero et al. 2006) using the Ecosym software (Gotelli & Entsminger 2004). Hypoth- esis of non-random similarity (Gotelli & Graves 1996) was tested using the RA2 (niche breadth relaxed / zero states retained) and RA3 (niche breadth retained / zero states reshuffled) Monte Carlo randomisation algorithms (Wine- miller & Pianka 1990) generating 1000 pseudomatrices considering each OTU equiprobable. Normality (Lilliefors test) and homoscedasticity (Fisher test) were assured prior to the application of parametric tests. Individual diversity and number of prey per pellet were compared using one-way ANOVA. Population diver- sity estimations obtained through jack-knife could not be compared using ANOVAs since the software provides on- ly mean+SE and diversity is non-additive (Carretero & Llorente 1993). Instead, t-tests corrected for multiple tests (using False Discovery Rate, FDR, Benjamini & Hochberg 1995) were applied. RESULTS Pellets were obtained from 45 C. ocellatus and 86 P. fil- folensis (58 males and 28 females). The SVLs in mm, mean+SE (range) of such specimens were 104.02+2.26 (62.0—140.0) for C. ocellatus, 65.45+0.56 (54.0—72.0) for male P. filfolensis, and 60.00+0.53 (44.5—67.0) for female P. filfolensis, The skinks were, in fact, much bigger than the wall lizards which displayed slight sexual size dimor- Bonn zoological Bulletin 57 (2): 111-118 phism favourable to males (ANOVA F, jy = 267.58, P < 10-6, Scheffé tests C. ocellatus-P. filfolensis males P< 10°, C. ocellatus-P. filfolensis females P< 10-6, P. fil- folensis males-females P = 0.05). The number of prey items by pellet (Table 1) was simi- lar between both species and between male and female P. filfolensis (ANOVA F j53 = 0.45, P = 0.64). However, the taxonomic composition of the diet (Table 1) showed sub- stantial interspecific differences, whereas intersexual dif- ferences within P. filfolensis were minor. Both species con- sumed important amounts of plant matter (IU = 41.61% in C. ocellatus and IU = 18.37% in P. filfolensis). With- in P. filfolensis, males (IU = 22.55%) used this resource more than females (IU = 9.69%). Moreover, C. ocellatus also consumed seeds and fruits (IU = 18.31%) but P. fil- folensis almost did not IU = 1.24%). Regarding the prey of animal origin (Table 1), the diet of C. ocellatus was strongly biased towards Coleoptera (IU = 24.95%) and only secondarily to insect larvae (IU = 7.79%). In contrast, the animal prey consumed by P. fil- folensis were more evenly distributed between Formici- dae (IU = 26.73%), Coleoptera (IU = 15.08%) and insect larvae (IU = 14.27%) with minimal differences between sexes. Interestingly, the diet of the Maltese wall lizard in- cluded some minor prey (Araneae, Pseudoscorpiones, Acarina, Homoptera, Malophaga) that were completely absent from the diet of the Ocellated skink. Overall, ani- mal diet was very similar between male and female P. fil- folensis, the latter consuming more Araneae and Het- eroptera (Table 1). It is worth noting that tails of juvenile P. filfolensis were found in adult conspecifics (two in males and two in females) and also in C. ocellatus (also two). Consequently, diet diversity (Table 2) was lower in C. ocellatus than in P. filfolensis, with no differences between males and females. This was true when considering either individuals (ANOVA F, 9g = 9.93, P < 10-4; Scheffé tests C. ocellatus - P. filfolensis males P = 0.0003, C. ocella- tus —P. filfolensis females P = 0.0005, P. filfolensis males - P. filfolensis females P = 0.99) or populations (C. ocel- latus — P. filfolensis males To, = 4.67, P = 5*10-6, Pepe <10-4; C. ocellatus — P. filfolensis females T7,; = 4.82, P = 4*10°6, Pepe < 104; P. filfolensis males — P. filfolensis females Tg, = 1.59, P = 0.06, Peper = 0.06). As to the size of the items consumed (Table 3), C. ocel- latus ate bigger prey than P. filfolensis but males and fe- males of the latter species did not differ (ANOVA F, 17 = 20.81, P< 10-6, Scheffé tests C. ocellatus-P. filfolensis males P < 10°, C. ocellatus-P. filfolensis females P = 5*10°, P. filfolensis males-females P = 0.94). The modal size class of C. ocellatus was 5-10 mm whereas ©ZFMK 114 Miguel A. Carretero et al. P. filfolensis was shifted to the 1-5 mm class. In fact, ex- cept in two females, pellets of P. filfolensis did not con- tain items larger than 10 mm. No significant correlation between prey and predator sizes was detected within each group although those P. filfolensis females eating the 10-15 mm prey were bigger than the rest (ANOVA Fj 35 = 5.35, P= 0.009; Scheffé tets: 1-5 mm — 5—10 mm P= 0.86, 1-5 mm — 10-15 mm P = 0.01, 5-10 mm — 10-15 mm P= 0.01). Finally, diet overlaps (Table 4) calculated from both tax- onomical and size classification of prey were very simi- lar, attaining moderate values between species but high values between male and female P. filfolensis. Pseudocom- munity analysis at species level revealed that taxonomi- cal overlap was higher than simulated in the RA3 matrix (niche breadth retained, P = 0.02) but similar to the RA2 matrix (zero states retained, P = 0.50). When considering the three classes (C. ocellatus, male and female P. filfolen- sis) none of the two algorithms indicated significant de- viations from random. No significant differences were ei- ther detected for the size overlap. DISCUSSION Differences in lizard diet arise from multiple factors in- cluding anatomy, sex, reproductive state, body condition, experience, prey availability, predation pressure, compe- tition and evolutionary history (Schoener 1974; Dunham 1980; Pianka 1986; Losos 1992; Vitt & Zani 1998; Vitt et al. 1999; Perry & Pianka 1999; Pitt & Ritchie 2002; Car- retero 2004). In Lampione, the manifest size differences between both species constitute the most obvious con- straint for the prey they consume. Within species, prey sizes tend to follow a logarithmic distribution, small in- dividuals simply not been able to consume the biggest items of the prey spectrum of the large individuals (Pi- anka 1986). However, between species this pattern can be altered if drastic divergence in anatomy or foraging tac- tics occurs (Carretero 2004). This seems to be the case, since C. ocellatus not only consumed large prey inacces- sible for P. filfolensis as expected, but also kept the same number of prey items but biased to medium sizes. This result suggests that both species may follow different for- aging strategies (Perry & Pianka 1999). In fact, C. ocel- latus is described a semi-fossorial, sit-and-wait forager in plant litter or under stones (Arnold 1984; Capula & Luisel- li 1994; Kalboussi & Nouira 2004; Lo Cascio et al. 2008) whereas P. filfolensis as most lacertids actively forages on the surface (Corti & Lo Cascio 2002; Bombi et al. 2005; Lo Cascio et al. 2006). Nevertheless, there is also evidence for anatomical constraints, since ocellated skinks con- sumed more hard prey (Coleoptera) than the wall lizards. In lacertids, large species tend to consume more Bonn zoological Bulletin 57 (2): 111-118 Coleoptera (Carretero et al. 2006) and there is experimen- tal evidence for inter- and intraspecific differences in bite force for prey crashing associated with the jaw muscle mass (Herrel et al. 1999, 2001). Nonetheless, sexually di- morphic lacertid heads, as intersexual differences in bite force, primarily derive from sexual selection (Herrel et al. 1999; Kaliontzopoulou et al. 2007), and dietary shifts (minimal in P. filfolensis) should be interpreted as a by- product. While divergent anatomy, foraging tactics and habitat use between both species accounted for a substantial part of the interspecific variation found, comparison with other populations indicates that other factors modified the tax- onomic composition of their diets. As other insular lacer- tids in the Mediterranean (Pérez-Mellado & Corti 1993; Carretero et al. 2001; Corti et al. 2008), P._filfolensis con- sumed great amounts of ants not only in Lampione but al- so in Linosa (Sorci 1990; Bombi et al. 2005) and Lampe- dusa (Lo Cascio & Corti 2008). Since only Podarcis pop- ulations inhabiting ancient Mediterranean islands (i.e. Balearics, Mylos) are myrmecophagous, this has been in- terpreted as a result of long term evolution in insularity (Pérez-Mellado & Corti 1993; Carretero 2004). The de- crease in predation pressure, the scarcity of alternative re- sources, together with the gregarious behaviour and sea- sonal stability of this prey may compensate for its low profitability and noxiousness (Carretero 2004). Apparent- ly, C. ocellatus has not been able to follow a similar strat- egy since neither continental nor insular populations are myrmecophagous (Capula & Luiselli 1994; Kalbousssi 2004; Lo Cascio & Corti 2008). Whether this is due to evolutionary constraints or to recent colonisation of Lam- pione currently remains under debate. Cannibalism and, in general, saurophagy seem also to in- crease in insular conditions due to the scarce resources and high lizard densities (Pérez-Mellado & Corti 1993; Car- retero et al. 2001) as is the case of P. filfolensis and C. ocel- latus (Scalera et al. 2004; Lo Cascio et al. 2006). How- ever, the predation of P. filfolensis by C. ocellatus consti- tutes not only an additional food source but also an in- stance of direct, asymmetric interaction between both species (Chase et al. 2002). As in the case of the ants, plants are also low profitable matter and their consumption seems to be restricted to old insular lacertid lineages which have developed behaviour- al and anatomical adaptations for herbivory (Pérez-Mel- lado & Corti 1993; Carretero 2004). Both P. filfolensis and C. ocellatus were partially herbivorous in Lampione. However, other populations of P. filfolensis studied also consumed substantial amounts of seeds, fruits and other plant remains (Sorci 1990; Bombi et al. 2005; Lo Cascio & Corti 2008) and even seed dispersal for some plant ©OZFMK Diet of Lampione lizards 115 species has been described in Linosa (Fici & Lo Valvo 2004). This suggests niche conservatism for herbivory in this species. In contrast, herbivory seems to be rare in C. ocellatus. Only two other microinsular populations, on Lampedusa and the Conigli islet (Lo Cascio & Corti 2008; Lo Cascio et al. 2008), were partially herbivorous, where- as, no or almost no plant remains were found in the pop- ulations from Sardinia (Capula & Luiselli 1994) and Tunisia (Kalboussi & Nouira 2004). Nevertheless, even continental ocellated skinks accept fruits in captivity (Schleich et al. 1996) suggesting certain exaptation for her- bivory in this species. Comparing the C. ocellatus popu- lation from Lampione with those from the other Pelagian islands (Lo Cascio & Corti 2008; Lo Cascio et al. 2008), there is an apparent trend for increasing the degree of her- bivory with isolation and for decreasing it with island area. The analysis of trophic diversity indicates that P. filfolen- sis is more euryphagous than C. oce//atus, with minimal intraspecific variation in the first. Remarkably, for both species, trophic diversity was much higher in populations than in individuals, indicating strong interindividual vari- ation typical of generalist predators (Carretero et al. 2006). In fact, C. ocellatus displayed even stronger differences (4x) than P. filfolensis (3x), which is accordance with its sit-and-wait trend. Finally, niche overlap summarises the trophic traits of the community previously exposed. Coincidence between tax- onomical and size overlaps and higher values between species than within species indicate that intrinsical mor- phological constraints constitute the main force for the or- ganisation of this microinsular community. Pseudocom- munity analysis does not support community structure but instead points to enlarged, more overlapped trophic nich- es. When compared to other conspecific populations, conver- gence in trophic strategies (herbivory, saurofagy) between both species due to insular conditions seems the more fea- sible hypothesis for explaining these results. Moreover, evolutionary history at both deep (foraging strategies, Vitt & Pianka 2005) and shallow (recent or ancient insular colonisation, see above) levels seems, rather than resource partitioning, responsible for the moderate trophic overlaps recorded and even may explain why both species coexist under the harsh conditions of this tiny islet. Nevertheless, coincidence of trophic overlap between two species with the direct predation of one on the other merits further analyses (see also Castilla 1995). Acknowledgements. We thank Giuseppina Nicolini, director of the Natural Reserve “Isola di Lampedusa’,, all the staff of Legam- biente, and Giuseppe Sorrentino of the Marine Protected Area of the Pelagie Islands for their irreplaceable assistance during Bonn zoological Bulletin 57 (2): 111-118 several visits at Lampione. We are also sincerely grateful to Damiano Sferlazzo, who help us significantly during the field work. The analysis of the collected samples has been done in the framework of the scientific research projects of the Associ- azione Nesos (Lipari). Analytical work was funded by the proj- ect PTDC/BIA-BDE/67678/2006 and PTDC/BIA- BEC/101256/2008 (to MAC) of Fundagao para a Ciéncia e a Tec- nologia, FCT (Portugal). 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Syst. 33: 475-505 Winemiller KO, Pianka ER (1990) Organization in natural as- semblages of desert lizards and tropical fishes. Ecological Monographs 60: 27—55 Received: 08. II. 2010 Accepted: 16.X. 2010 Table 1. Descriptors of the taxonomic composition of the diet for Chalcides ocellatus and Podarcis filfolensis from Lampione is- land. OTU: Operational taxonomical unit, %P: percentage of presence; %N: percentage of numerical abundance; IU: Resource use index: — not consumed; 0.00: consumed but index value next to zero. Chalcides ocellatus Podarcis filfolensis Podarcis filfolensis Podarcis filfolensis total total males females OTU %P %N IU %P %N IU %P %N IU %P %N IU Gastropoda 9.09 1.28 0.65 2.33 0.39 0.09 7.14 1.16 0.42 Isopoda 9.09 1.28 0.65 11.63 1.94 1.47 10.34 1.74 Hall 1429233" 170 Araneae — - - 12.79 2.13 1.68 Sally) 0.87 0.35 28.57 4.65 5.10 Pseudoscorpiones - — - LO eal (2204 52925 2-9) 2.24 7.14 2°33, 0:85 Acarina - — - 2733) 10:39) 10109 3.45 0.58 0.15 - ~ — Diptera SMS 2255 le 24.42 6.01 5.66 Zi59) 16198 6.51 17.86 4.07 3.20 Coleoptera 66.67 18.30 24.95 55.81 12.40 15.08 56.90 12:21 1498 53.57 12:79 16.48 Hymenoptera - — — 25.58 7.56 6.95 PES eH 6.41 21.43 814 7.04 Formicidae 18.18 4.26 3.19 60.47 23.45 26.73 58:62) 22538 25:18 ~64:29 25:58 3115 Homoptera ~ = — 6.98 1.74 0.97 10.34 2.62 1.60 - — = Heteroptera 6.06 0.85 0.27 17.44 3.49 2.90 1325/9) 233 eS) DOO” Bes, “5:25 Mallophaga — — = 930 1.55 1.06 10.34 = 1.74 ills) 7.14 1.16 0.42 insect larvae BOBO 8S" AS 37.21 14.15 14.27 34.48 15.70 14.93 42.86 11.05 13.26 Arth ind. 9.09 1.70 0.78 O30 ESS: 106 Sali 0.87 0.35 IASG 229i 24g Squamata 3.03 0.43 0.00 4.65 0.78 0.35 3.45 0.58 0.15 7.14 1.16 0.42 seeds, fruits MPN Poss) Itsesill 6.98 3.10 1.24 6.90 1.16 0.59 7.14 6.98 2.55 other plant matter 54.55 35.32 41.61 45.35 16.67 18.37 53.45 20.06 22.55 2857 9.88 9.69 Total (mean+SE) 45 pellets, 235 items 5.91+0.83 items/pellet 86 pellets, 516 items 5.19+0.34 items/pellet 58 pellets, 344 items 5.16+0.38 items/pellet 28 pellets, 178 items 5.25+0.70 items/pellet Bonn zoological Bulletin 57 (2): 111-118 ©ZFMK 118 Miguel A. Carretero et al. Table 2. Diet diversities of Chalcides ocellatus and Podarcis filfolensis from Lampione island. Numbers indicate mean+SE. Hi: individual diversity; Hp: population diversity; Hz: total accumulated diversity; all using Brillouin’s index. Species (class) N Hi Hp Hz Chalcides ocellatus (total) 45 0.61+0.03 2/201 2.44 Podarcis filfolensis (total) 86 0.98+0.02 3.33+0.08 B22 Podarcis filfolensis (males) 58 0.98+0.03 3.19+0.08 3.05 Podarcis filfolensis (females) 28 0.98+0.06 3.43+0.14 Bell 7 Table 3. Descriptors of the prey size composition of the diet for Chalcides ocellatus and Podarcis filfolensis from Lampione is- land. OTU: Operational taxonomical unit, %P: percentage of presence; %N: percentage of numerical abundance; IU: Resource use index: — not consumed; 0.00: consumed but index value next to zero. Chalcides ocellatus Podarcis filfolensis Podarcis filfolensis Podarcis filfolensis total total males females OTU %P %N IU %P %N IU %P %N IU %P %N IU 0-5 mm 32.14 24.62 19.09 80.00 67.24 68.92 73.68 64.10 64.41 100.00 73.68 78.66 5-10 mm 85.71 58.46 69.63 84.00 31.03 31.08 84.21 135.90) -35:59 83.33 DOS me 2ies 4 10-15 mm 17.86 7.69 5.31 4.00 7D, O00 16.67 5.26 0.00 > 15mm 17.86 9.23 5.96 = = - Total (mean+SE) 65 (of 235) items measured 116 (of 516) items measured 78 (of 344) items measured 38 (of 178) items measured 7.42+0.51 mm 4.22+0.24 mm 4.29+0.27 mm 4.0840.47 mm Table 4. Diet overlaps (Pianka’s index) between the lizard species and classes from Lampione island considering the taxonomi- cal and size composition of the prey consumed. taxonomical overlap size overlap C. ocellatus (total) — P. filfolensis (total) 0.54 0.63 C. ocellatus (total) — P. filfolensis (males) 0.54 0.64 C. ocellatus (total) — P. filfolensis (females) 0.53 0.50 P. filfolensis (males) — P. filfolensis (females) 0.97 0.97 Bonn zoological Bulletin 57 (2): 111-118 ©ZFMK Bonn zoological Bulletin Volume 57 Issue 2 | pp. 119-126 Bonn, November 2010 Evolutionary reproductive morphology of amphibians: an overview Susanne Kithnel!, Sandy Reinhard! & Alexander Kupfer!2* | Institut fiir Spezielle Zoologie und Evolutionsbiologie mit Phyletischem Museum, Friedrich-Schiller-Universitat Jena, Erbertstr. 1, D-07743 Jena, Germany 2 Biologie und Didaktik, Universitat Siegen, Adolf-Reichwein-Str. 2, D-57068 Siegen * Correspondence; E-mail: alexander.kupfer@uni-jena.de Abstract. Reproduction is a crucial trait in the life history of any organism, and vertebrates, whether aquatic or terres- trial, have evolved an extraordinary diversity of reproductive strategies and morphologies. Among tetrapods, the diver- sity of reproductive modes is exceptionally high in amphibians, who also show multiple trends towards terrestrialisation and internal fertilisation. Herein we give a brief overview of the diversity of amphibian reproductive morphology, with a special emphasis on the cloaca, for all three major lineages, 1.e., anurans, urodeles and caecilians. Key words. Reproduction, genital morphology, Amphibia. INTRODUCTION Reproduction is a crucial trait in the life history of any or- ganism and scientists have been intrigued and challenged by this event, and the structures associated with it, ever since the days of Darwin (1871). Both aquatic and terres- trial vertebrates have evolved an extraordinary diversity of reproductive strategies and morphologies, including va- rieties of oviparity and viviparity (Meisenheimer 1921; Lombardi 1998). Among tetrapods, the diversity of repro- ductive modes is exceptionally high in amphibians. In this group, we also see multiple trends towards terrestrialisa- tion and internal fertilisation (e.g. Haddad & Prado 2005). Reproductive modes such as viviparity have evolved in- dependently in all three lineages of modern amphibians (e.g. Noble 1927; Wake & Dickie 1998; Wells 2007). Herein we interpret a reproductive mode as a combina- tion of several reproductive traits, including oviposition site, clutch characteristics, stage and size of hatchling, and type of parental care (sensu Salthe 1969). Internal fertilization is a precondition for viviparity (Wourms & Lombardi 1992; Bohme & Ziegler 2008). It is associated with different strategies of sperm transfer, which have evolved within all three amphibian orders, ranging from cloacal apposition in anurans to a true cop- ula via a male intromittent organ in caecilians (Sever et al. 2001; Kupfer et al. 2006). Like most other tetrapods, amphibians have a cloaca, a chamber that receives prod- ucts from the kidneys, the intestine and the gonads, and opens to the outside through a cloacal opening or vent (Kardong 2006). Bonn zoological Bulletin 57 (2): 119-126 Below we review the diversity of amphibian cloacal mor- phologies involved in ensuring a secure direct sperm trans- fer and internal fertilization among anurans, urodeles and caecilians. REPRODUCTIVE MORPHOLOGY OF AMPHIBIANS Anura The majority of anurans, currently including almost 6000 species (AmphibiaWeb 2010), practice external fertiliza- tion, and thus have no special male cloacal arrangements facilitating direct sperm transfer (recently summarised by Wells 2007). During copulation, males grasp females firm- ly with their forearms (termed amplexus). In most cases, sperm is directly released on the eggs protruding from the female cloaca, but in some cases fertilisation takes place without amplexus (e.g. Crump 1974; Kunte 2004). Inter- nal fertilisation is rare among anurans, and mostly con- nected to viviparity or other complex parental care mech- anisms (e.g. Wake 1993; Beck 1998). Exceptionally, the phylogenetically basal tailed frogs As- caphus truei and A. montanus are the only anurans known to have evolved a true intromittent organ in males (see Figs 1A—B). During courtship they practise a combination of amplexus and copulation called “copulexus” (see Sever et al. 2001; Stephenson & Verrell 2003). The so-called ©ZFMK 120 Susanne Kiihnel et al. Fig. 1. | Reproductive morphology of anurans and salamanders. (A) Inguinal amplexus (“copulexus”) of Ascaphus truei. (B) ma- le Ascaphus truei. The “tail”, a cloacal extension, can be inserted into the cloaca of the female during amplexus, ventrolateral view. (C) male Mertensophryne micranotis (Anura: Bufonidae), left, dorsal view and its cloaca, right, caudal view (after Grandison 1980). (D) cephalic amplexus of Notophthalmus viridescens (Urodela: Salamandridae). The male grasps the females’s neck whilst fan- ning pheromones towards her nostrils. (E) cloacal region of lentic breeding Cynops pyrrhogaster (Urodela: Salamandridae). The male’s cloaca (left) is heavily swollen compared to that of the female (right). (F) cloacal region of lotic breeding Euproctus mon- tanus (Urodela: Salamandridae, after Brizzi et al. 1995). Males (left) possess a cloacal protuberance (cp) which bears a protusible pseudopenis (pp), whereas the female cloaca is slightly conical shaped and its opening is located ventrally (right). (G) amplecting pair of Calotriton arnoldi (Urodela: Salamandridae). The male grasps the female’s trunk with his tail. Bonn zoological Bulletin 57 (2): 119-126 ©ZFMK Amphibian reproductive morphology 121 “tail” resembles the posteriorly extended cloaca, proxi- mately attended by Nobelian rods and strengthened by vascularized tissue that is engorged with blood during cop- ulation. This gives the ventral cloacal surface a pinkish colour (Noble & Putnam 1931; Duellman & Trueb 1994). To insert the posterior pointing “tail” into the female vent, the male first flexes his pelvis at a right angle to the ver- tebral column. Contraction of the paired Musculi compres- sores cloacae (Duellman & Trueb 1994) bend the intro- mittent organ ventrally, with the male vent pointing ante- riorly (Slater 1931). Keratinised spines are present with- in the male cloacal orifice, but whether they function to enhance the attachment of the male to the female remains unclear (Noble & Putnam 1931; Metter 1964). Additionally, internal fertilisation including an amplexus and cloacal apposition occurs in a few anurans, such as several species of viviparous African dwarf toads Nec- tophrynoides (Wake 1980; Wake & Dickie 1998) and Nim- baphrynoides (Sandberger et al. 2010), and in two species of Caribbean Eleutherodactylus, the viviparous E. jasperi (Dewry & Kirkland 1976; Wake 1978) and the oviparous- direct developing E. coqui (Townsend et al. 1981). Mat- ing has only been observed in couples of E. coqui in a spe- cial amplectic position called reverse hind leg clasp, that is initiated by the female (Townsend & Stewart 1986). Males do not clasp, and the female rests her hind legs on top of the male’s legs. This behaviour might be correlat- ed with terrestrial reproduction and internal fertilization. It is also thought to be present in the viviparous E. jasperi but has not yet been observed (Wake 1978). Within the African Bufonidae, all species of Nectophrynoides (and also Altiphrynoides malcolmi and Nimbaphrynoides oc- cidentalis, former members of Nectophrynoides, see Frost et al. 2006) practice internal fertilization. A/tiphrynoides and Nimbaphrynoides both show a dimorphism of the male and female vent, and an inguinal amplexus in a unique belly-to-belly position has been reported as well (Grandison 1978). As in the internally fertilising Ascaphus ssp., males of the East African toad Mertensophryne micranotis (Bufonidae) exhibit modifications of the cloacal region (Duellman & Trueb 1994). They have small conical spines around the rim of the vent and at the entrance to the cloacal tube re- stricted to the ridges of the puckered vent (Grandison 1980, see also Fig 1C). Males and females keep a very tight cloacal contact during mating. Although the cloacal spines play a role to ensure a close apposition of the vents, to secure internal fertilisation, there is no evidence for a direct interlocking mechanism in the furrows of the female vent (Grandison & Ashe 1983). Another potential record of internal fertilisation is provid- ed for the neotropical Pumpkin Toadlet Brachycephalus Bonn zoological Bulletin 57 (2): 119-126 ephippium (Pombal et al. 1994). During mating, males shift from an inguinal to an axillary amplexus to optimal- ly allow positioning of the vents, and thus maximize fer- tilization of the relatively large eggs (5.1 to 5.3 mm). A further record of viviparity in fanged frogs (Limnonectes spec.) from Sulawesi probably also involves internal fer- tilisation(Emerson 2001). It can be hypothesised that (1) many terrestrially breed- ing species with large direct-developing clutches are in- ternal fertilizers and (2) if additional viviparous species are encountered they will also show internal fertilisation. Thus, internal fertilisation and viviparity in anurans might be more widespread than currently recognized (see also Wake 1978). Data on the reproductive biology, including the mating be- haviour, of many species is still lacking (Duellman & Trueb, 1994; Wells 2007). Life history data from around 23 % of the currently known species is missing, as listed in the data deficient category of the IUCN (Stuart et al. 2008). Urodela The majority of the 590 species of urodeles exhibit inter- nal fertilization, whilst only males of the basal families Hynobiidae, Cryptobranchidae, and presumably Sirenidae, fertilise eggs externally (summarised in Duellmann & Trueb 1994; Wells 2007). The complex and elaborate courtship behavior of most salamanders includes the dep- osition of a spermatophore by the male, which is subse- quently received by the female. A true intromittent organ in salamanders is lacking, although direct sperm transfer can be found in one species — the Corsican brook newt Euproctus montanus, a lotic breeding endemic of the is- land of Corsica. The cloaca of the male brook newt re- sembles a conical protuberance (Fig IF). The cloacal chamber hosts a “pseudopenis”, a broad conspicuous papilla, which can be evaginated during mating (Brizzi et al. 1995; Carranza & Amat 2005). The male grasps the female during amplexus, holding her tail with his jaws and wrapping his tail around her trunk, whilst his backward projecting cloaca is positioned close to that of the female. A deep groove along the ventral surface of the pseudope- nis, which is aligned with the cloacal tube, guarantees a guided, unidirectional flux of cloacal products. Thus, sperm mixed with secretory products is transferred direct- ly into the female’s cloaca. The Salamandroidea that prac- tice internal fertilization possess a distinct set of male cloa- cal glands necessary for spermatophore production (Sev- er 2002). The glands are hormonally controlled and hy- pertrophied during the breeding season, often causing a sexual dimorphism in cloacal shape. However, in Euproc- ©ZFMK tus montanus, cloacal glands are reduced or partly lack- ing (Brizzi et al. 1995; Sever 2002). Males of six salaman- drid genera possess a so-called “pseudopenis”, a projec- tion of the dorsal roof which nearly fills the entire ante- rior chamber of the cloaca. It is involved in shaping and expulsion of the spermatophore (Halliday 1998), but can- not be everted as in the Corsican brook newt (Brizzi et al. 1995; Carranza & Amat 2005). Sexes of most species, regardless of the mode of fertili- sation, show a sexual dimorphism in cloacal shape (Figs 1E—-F). Usually, the male cloaca appears larger and more swollen than the female one. This is caused by the activ- ity of the glands mentioned above (see also Sever 2002). Species that breed in the water and show elaborate courtship dances or walks, such as some members of the family Salamandridae, produce courtship pheromones, which are fanned towards the female using the tail. Sala- manders that mate terrestrially also use courtship pheromones secreted from specialised glands to attract fe- males. Pheromone-producing cloacal glands are therefore highly influenced by sexual selection (e.g. Sever 2002; Houck et al. 2008). Usually, female cloacae are less promi- nent, but they may also possess up to three types of cloa- cal glands in Salamandroidea, mainly accounting for sperm storage (spermathecae), a unique feature among vertebrates (Sever 1994). Females may retain and mix vi- able spermatozoa from multiple matings in the spermath- ecae for longer periods (e.g. Steinfartz et al. 2006). Fe- male Eurycea fertilise eggs from stored sperm up to eight months after insemination, female Notophthalmus viri- descens effectively store sperm for up to six months, and female Salamandra salamandra are reported to store sperm for up to two years (Sever et al. 1996; Stebbins & Cohen 1997; Sever & Brizzi 1998). Additionally, the shape of female cloacae can be adapted to a specific substrate for oviposition and type of water body. Females of stream-breeding species, such as Calotriton asper, sometimes have a conically shaped cloa- ca for egg deposition and safe attachment between stones and in crevices (e.g. Carranza & Amat 2005). Lotic breeders such as Calotriton ssp. often engage in an amplexus directly transferring the spermatophore into the female cloaca (Fig 1G). It ensures direct and rapid sper- matophore uptake, and thus reduces energy wasting, which can hardly be avoided during aquatic breeding where the male and the female often have no physical contact. Breed- ing patterns including an amplexus are common in sala- mandrids. Multiple ways of female capture are known, such as the cephalic capture of Notophthalmus ssp. (see Fig 1D), the dorsal capture of Zaricha, or the ventral cap- ture performed by fire salamanders (Stebbins & Cohen 1997). The mating amplexus may last up to several hours, Bonn zoological Bulletin 57 (2): 119-126 122 Susanne Kiihnel et al. depending on the species. Salamanders of the family Am- bystomatidae mate in the water, and the males guide fe- males to spermatophore-uptake using a “tail-nudging- walk”, except in Ambystoma gracile, A. laterale, A. jef- fersonarium and A. macrodactylum, which capture fe- males in an amplexus (Duellmann & Trueb 1994; Verrell & Krenz 1998). In contrast, some plethodontids perform a unique “tail-straddling-walk” behaviour (e.g. Arnold 1977). Gymnophiona In contrast to all salamanders (with the exception of Eu- proctus montanus) and frogs (with the exception of As- caphus ssp.), the male caecilian cloaca is evertible through the vent and operates as an intromittent organ or phallus, a unique structure among tetrapods (Tonutti 1931, see al- so Fig 2A). Presumably all ca. 190 caecilian species (oviparous and viviparous) practice internal fertilisation with the help of the phallodeum (Tonutti 1931; Wake 1972; Gower & Wilkinson 2002), which is inserted into the female vent during copulation (e.g. Kupfer et al. 2006a). The caecilian vent is simply surrounded by several folds, which are variably arranged among the groups and dis- play sexual dimorphism in some species, such as mem- bers of the Typhlonectidae (e.g. Taylor 1968; Kupfer 2007). In contrast, the cloaca is highly complex and di- verse. The male caecilian cloaca is an elongated tube di- vided into two distinct chambers. The cranial urodeum is rather simply built, bearing longitudinal ridges, and con- nects to the intestine and the urogenital ducts, which en- ter after performing a U-bend (Sawaya 1942; Gower & Wilkinson 2002). An extraordinary feature is the presence of Millerian ducts, which become glandular during repro- ductive activity, and secrete a fluid containing lipids and sugars necessary for sperm motility (e.g. Wake 1981). The caudal phallodeum is more broadly built and the inner structure is very different. The ridges are arranged in a more complex pattern (running transversely). In adults it is often equipped with tuberosities or crests, which give the phallodeum a characteristic morphology and gives rise to an extraordinary variation in shape (Wiedersheim 1879; Tonutti 1931, 1933; Wake 1972; Exbrayat 1991; Gower & Wilkinson 2002, see also Fig 2), that is impotant for caecilian systematics (Miller et al. 2005). East-African scolecomorphid caecilians even have cartilaginous spicules (Wake 1998). In many species, pouchy dorsolat- eral appendixes — so called “blind sacs” — extend anteri- or to the phallodeum. During eversion, the luminal sur- face of the phallodeum represents the outer structure of the phallus, with the urodeum lying in-between (see Tonut- ti 1931; Gower & Wilkinson 2002, see also Fig. C right). ©ZFMK Amphibian reproductive morphology 123 Smm 5mm 2mm caudal I 2mm 2mm Fig. 2. Genital morphology of caecilian amphibians. (A) male Chthonerpeton indistincum (Gymnophiona: Typhlonectidae) show- ing an everted phallus, MHNM 09323, right — detail. (B) Geotrypetes seraphini (Gymnophiona: Caeciliidae), lateral (left), dorsal (central) and ventral (right) view of the everted phallus, AK 01149. (C) Zyphlonectes natans (Gymnophiona: Typhlonectidae), SRuCT- Scan of the everted phallus. Right — virtual clipping, frontal view. (D) SRuwCT-Scan of female cloaca (/chthyophis cf. kohtaoen- sis). Dorsolateral view, virtual cut of cloacal sheath, cranial part and blind sacs, green - cloaca, violet — oviducts, yellow — blad- der. Abbreviations: MHNM = Museo Nacional de Historia Natural Montevideo Uruguay, AK = Alexander Kupfer collection. Bonn zoological Bulletin 57 (2): 119-126 ©ZFMK 124 Susanne Kihnel et al. To retract the cloaca within the body after copulation, cae- cilians possess a specific muscle (musculus retractor cloa- cae), which is also found in some females (Wilkinson 1990). The female cloaca of caecilians has received little atten- tion (e.g. Hypogeophis rostratus Tonutti 1931; Ty- phlonectes compressicauda, Exbrayat 2006), the only ded- icated morphological study was presented by Wake (1972), proposing a functional association between the specific male and female morphologies. The female cloa- ca is supposed to be non-eversible (Wilkinson 1990), therefore displaying a different morphology. Generally it is shorter than in males, and the urogenital ducts lack a copulatory loop (see Fig. 2D). There is also evidence for a bisection of the female cloaca (Exbrayat 1991; Kiihnel et al. submitted). The cranial chamber is homologous to the male urodeum. The caudal chamber is marked by a different arrangement of longitudinal cloacal folds most- ly lacking tuberosities, and therefore easily recognised. Nothing 1s at present known about how far the male phal- lus inserts into the female cloaca, and if special structures corresponding to the male ornamentation are present, help- ing in fixation during copulation. Copulations in caecilians have rarely been observed. Da- ta are available for two aquatic/semiaquatic species, the typhlonectids Typhlonectes compressicauda and Chthon- erpeton indistinctum. Pairs of C. indistinctum copulated for betwen 30 minutes and 5 hours (Barrio 1969) and those of 7. compressicauda for between 75 minutes and 3 hours (Murphy et al. 1977; Billo et al. 1985). Observations on copulations in terrestrial caecilians have, to the best of our knowledge, only been presented for the Indian ichthyophi- id Ichthyophis beddomei (Bhatta 1999) and Ichthyophis cf. kohtaoensis (Kupfer et al. 2006a). Bhatta reports on a copulation lasting for about 40 or 45 minutes, an obser- vation fitting well with the duration time of about 45 min- utes that was observed in Ichthyophis cf. kohtaoensis (Kupfer et al. 2006a). Caecilians show a remarkable diversity of reproductive modes associated with parental care (e.g. Wake 1977; Himstedt 1996; Wilkinson & Nussbaum 1998). Oviparous caecilians guarding egg clutches in terrestrial chambers (e.g. Sarasin & Sarasin 1887-1890) either have the pre- sumed ancestral amphibian life cycle with aquatic larvae, or show direct development of juveniles with no aquatic larval stage (e.g. Brauer 1897). Females of viviparous species retain fertilised eggs. Embryogenesis is complet- ed within the oviducts, and after hatching the foetuses feed mainly intrauterinely on the hypertrophied oviductal lin- ing (e.g. Parker 1956; Welsch et al. 1977). After a long gestation period, the females give birth to fully metamor- phosed, precocial young with the adult-type morphology Bonn zoological Bulletin 57 (2): 119-126 (e.g. Billo et al. 1985; Exbrayat & Delsol 1985). Recent- ly, a novel form of parental investment, maternal derma- totrophy, a.k.a. skin feeding, where altricial young feed externally on the mother’s hyperthrophied skin, has been described (Kupfer et al. 2006b; Wilkinson et al. 2008). SUMMARY AND PERSPECTIVES In addition to their remarkable diversity of reproductive modes, amphibians also show large variation in their re- productive morphology. Many morphological peculiari- ties are related to the evolution of internal fertilisation, and ultimately to viviparity. In relation to fertilisation and sperm transfer, different strategies have evolved within the three amphibian orders, ranging from cloacal apposition in anurans to a true copula via a highly complex male in- tromittent organ in caecilians. Amphibians offer a prime system for comparative studies of evolutionary reproduc- tive biology. Research on the reproductive or genital mor- phology should include modern methodology, such as 3D reconstruction and soft tissue synchrotron radiation based X-ray microtomography (SRuCT, see Fig. 2 C—D). Be- cause amphibian diversity is steadily increasing (although at the same time many species are declining or even go- ing extinct) we envisage that many more unexpected re- productive strategies and morphologies remain to be dis- covered. Acknowledgements. Phillipp Wagner is congratulated for or- ganising this magnificent Festschrift volume. Many thanks for giving us the opportunity to contribute. We would like to dedi- cate our contribution to Wolfgang Bohme for his continuous ef- fort in studying vertebrate, especially squamate genitalia and their evolution. Synchrotron radiation based x-ray microtomog- raphy (SRuCT) of caecilian genitalia was carried out at beam- line BW2 at the Deutschen Elektronen Synchroton (DESY, Ham- burg) under experimental projects I-20080054 and I-20090089. Travel of SK and AK has been generously supported by DESY. Felix Beckmann (DESY) provided initial 3D reconstructions and Frank Friedrich (University of Hamburg) and Thomas Kleinte- ich (University of Washington) aided in the successful process- ing of 3D models. Alexander Haas (University of Hamburg) gen- erously provided laboratory space and technical support for SK during her collections-based research at the Museum fur Zoolo- gie Hamburg (ZMH). SK is financially supported by the Volk- swagen Stiftung (grant initiative “evolutionary biology”). An anonymous referee, Lennart Olsson and Philipp Wagner gave valuable comments on earlier versions of the manuscript. REFERENCES Arnold SJ (1977) The courtship behavior of North American salamanders with some comments on Old World salamandrids. Pp. 141-183 in: Taylor D & Guttman S (eds.) The Reproduc- tive Biology of Amphibians. Plenum Press, New York Barrio A (1969) Observaciones sobre Chthonerpeton indistinc- tum (Gymnophiona, Caecilidae) y su reproduccion. 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American Zoologist 32: 276-293 Received: 25.1X.2010 Accepted: 16.X.2010 ©OZFMK Bonn zoological Bulletin Volume 57 | Issue 2 i: pp. 127-136 Bonn, November 2010 Updated checklist of the living monitor lizards of the world (Squamata: Varanidae) André Koch!, Mark Auliya? & Thomas Ziegler; ! Zoologisches Forschungsmuseum A. Koenig & Leibniz Institute for Animal Biodiversity, Department of Herpetology, Adenauerallee 160, D-53113 Bonn, Germany; E-mail: a.koch.zfmk@uni-bonn.de ? Helmholtz Centre for Environmental Research — UFZ, Department of Conservation Biology, Permoserstr. 15, D-04318 Leipzig; Germany; E-mail: mark.auliya@ufz.de 3 AG Zoologischer Garten K6In, Riehler Strae 173, D-50735 K6lIn, Germany; E-mail: ziegler@koelnerzoo.de Abstract. We provide an update of BOhme’s (2003) checklist of the living monitor lizards of the world. Since this con- tribution, ten new species and one new subspecies have been described. Ten of these taxa were described from the is- lands of the Indo-Australian Archipelago. One Soterosaurus taxon (macromaculatus) was revalidated to subspecies sta- tus, whereas the younger melanistic taxon komaini was synonymized with the former. In addition, five taxa (beccarii, cumingi, marmoratus, nuchalis, and togianus), that were formerly treated as subspecies, were re-elevated to species rank resulting in 73 extant species (including 21 subspecies). This represents a 20% increase of the world’s varanid diversity since 2003. In addition, ongoing taxonomic studies on V spinulosus from the Solomon Islands (formerly a member of the V. indicus species group) indicate that this species most likely represents a new subgenus. Therefore, this taxon is currently treated incertae sedis. In sum, taxonomic research in monitor lizards remains incomplete. Further studies must be initiated to fully understand diversity and distribution of these CITES-listed lizards next to implications for sustain- able conservation measures. Key words: Reptilia, Varanus, Philippinosaurus, Soterosaurus, Euprepiosaurus, Odatria, checklist. INTRODUCTION Monitor lizards are among the largest living squamates of the world. They inhabit Africa, the Arabian Peninsula, South and Southeast Asia as well as the Indo-Australian Archipelago including Australia and several Pacific island groups. Due to their often large body size and ecological role as top predators in most environments they inhabit, monitor lizards have always been a small reptile group comparable to large placental carnivores (Sweet & Pianka 2007). Nevertheless, the diversity of monitor lizards has been underestimated for many decades. One of the latest comprehensive listings of all extant mon- itor lizards was published by BOhme (2003), who listed 58 different species and 28 subspecies. This checklist was prepared at the request of the Nomenclature Committee of the Convention on International Trade in Endangered Species of Wild Flora and Fauna (CITES), because of the increase in monitor lizard diversity in the early 1990s. This caused considerable confusion within trade records, which made communication about these economically im- portant lizards in the CITES domain rather difficult. Bonn zoological Bulletin 57 (2): 127-136 Therefore, BOhme’s (2003) checklist was adopted as the standard reference for the Varanidae by the 12th Confer- ence of the Parties to CITES in November 2002. The need for an updated list only seven years after the last synopsis by B6hme (2003) derives from ongoing descrip- tions of new species. This is partly due to the fact, that the understanding of monitor lizards, their systematics, and the underlying concepts have been refined in recent years (e.g. Koch et al. 2009). In addition, the taxonomic status of several nominal taxa has changed. This involves either subspecies elevated to species rank or the subgeneric al- location of species. Also, knowledge of distribution ranges of some rare monitor lizard species has been advanced by the examination of new voucher specimens and investi- gations in the field. Next to taxonomic and phylogenetic studies on the Varanidae, those focusing on conservation particularly of the Indo-Australian realm remain scarce. A summary of threats monitor lizards are exposed to in this region, cur- ©OZFMK 128 André Koch et al. rent conservation studies and measures in place, and the conservation status of all Indo-Australian species will be outlined in detail elsewhere. METHODS In the present paper, we compiled all monitor lizard taxa that were published after BOhme (2003). This includes al- so such names where the taxonomic status has changed. We basically follow the format of this author, which has earlier been used by Mertens (1963). The synonymy list of each taxon starts with the original citation and is then arranged chronologically with the source of the respective name and its type locality. Chresonyms are generally not included with the excep- tion of names with a changed taxonomic status exempli- fied by subspecies names, that were elevated to species level or when a new species is separated from a long re- cognized species. In these cases, the taxon name and the author(s) are separated by a “—*. In addition, type speci- mens for valid taxa are provided, if available. Collection acronyms are as follows: BMNH = British Museum of Natural History, London, UK; KU = Kansas University, Museum of Natural History, Lawrence, USA; MNHN = Muséum national d’Histoire naturelle, Paris, France; MSNG = Museo Civico die Storia Naturale di Genova Gi- acomo Doria, Genova, Italy; MZB = Museum Zoologicum Bogoriense, Bogor, Indonesia; NMW = Naturhistorisches Museum Wien, Vienna, Austria; PNM = Philippine Na- tional Museum, Manila, Philippines; RMNH = National Natural History Museum Naturalis, Leiden, Netherlands; SMF = Naturmuseum Senckenberg, Frankfurt, Germany; USNM = National Museum of Natural History, Washing- ton, USA; WAM = Western Australian Museum, Perth, Australia, ZFMK = Zoologisches Forschungsmuseum Alexander Koenig, Bonn, Germany; ZMA = Zoological Museum, University of Amsterdam, Netherlands; ZMB = Museum fiir Naturkunde, Berlin, Germany; ZMUC = Zo- ological Museum, University of Copenhagen, Denmark. Update of the checklist of extant monitor lizards by Bohme (2003) Subgenus Philippinosaurus Mertens, 1959 Varanus bitatawa Welton, Siler, Bennett, Diesmos, Duya, Dugay, Rico, van Weerd & Brown, 2010 2010 Varanus bitatawa Welton, Siler, Bennett, Diesmos, Duya, Dugay, Rico, van Weerd & Brown, Biol. Lett., 6: 654. — Type locality: Base of the San Ildefonso Peninsu- Bonn zoological Bulletin 57 (2): 127-136 la, Sitio Casapsipan, Barangay Casiguran, Municipality of Casiguran, Aurora Province, Luzon Island, Philippines. 2008 Varanus olivaceus — Eidenmiller & Philippen (in part), Terralog, 6: 103. Type specimens: Holotype PNM 9719 (formerly KU 320000), paratypes KU 322188 and PNM 9008. Distribution: Northern Luzon, Philippines. Remark: This species was recently separated from V. oli- vaceus based on minor genetic variation, morphological differences and biogeographic evidence (Welton et al. 2010). Subgenus Soterosaurus Ziegler & Béhme, 1997 Varanus s. salvator (Laurenti, 1768) 1768 Stellio salvator Laurenti, Synops. Rept.: 56. — Type locality: Sri Lanka. 1758 Lacerta monitor part. Linnaeus, Syst. nat., 10 (1): 201. — Type locality: In Indiis (nomen rejectum accord- ing to ICZN 1959, Opinion 540). 1947 Varanus salvator kabaragoya Deraniyagala, Proc. 3rd ann. Sess. Ceylon Assoc. Sci., 2 (Abstr.): 12. — Type locality: Ceylon (= Sri Lanka). Type specimen: Neotype ZFMK 22092, designated by Koch et al. (2007). Distribution: Sri Lanka. Remark: Until recently the nominotypic subspecies had the widest distribution range within the widespread V. sal- vator complex. Due to the revalidation of the subspecies Vs. macromaculatus from continental Southeast Asia, the nominotypic subspecies is now restricted to Sri Lanka. Varanus salvator macromaculatus Deraniyagala, 1944 1944 Varanus salvator macromaculatus Derantyagala, Spol. Zeyl. 24: 60. — Type locality: Siam (= Thailand). 1802 Tupinambis elegans Daudin (in part), Hist. nat. Rept., 3: 36. — Type locality: Surinam. 1831 Tupinambis exilis Gray in Griffith, Anim. Kingd., 9: 25. — Type locality: India (nomen dubium, fide Koch et al. 2007). ©ZFMK Update of B6hme’s (2003) checklist of monitor lizards 129 1834 Varanus vittatus Lesson in Bélanger, Voyage Ind. Orient. Zool.: 307. — Type locality: Indian subcontinent and islands at the mouth of the Ganges River (nomen du- bium fide Koch et al. 2007). 1842 Varanus binotatus Blyth, J. asiat. Soc. Bengal, 11: 867 (Lapsus fide Mertens 1942). 1942 Lacertus tupinambis Mertens (in part, non Lacépede, 1788) Abh. Senckb. Naturf. Ges., 466: 245. — Type loca- lity: unknown (Lapsus fide Brygoo 1987). 1947 Varanus salvator nicobariensis Deraniyagala, Proc. 3rd ann. Sess. Ceylon Assoc, Sci., 2, Abstr.: 12. — Type locality: Tillanchong, Nicobar Islands. 1987 Varanus salvator komaini Nutphand, J. Thai. Zool. Center, 2 (15): 51. — Type locality: Sea shore areas and small islands in south western Thailand. 2007 Varanus salvator macromaculatus — Koch, Auliya, Schmitz, Kuch & Bohme, Mertensiella, 16: 136. Type specimens: Lectotype MNHN 871, paralectotype MNHN 1884.77, designated by Koch et al. (2007). Distribution: Thailand, Peninsula Malaysia, Vietnam, southern China, Hainan, Sumatra, and Borneo and small- er off-shore islands. Remark: This subspecies of V. salvator was recently res- urrected from the synonymy of the nominotypic sub- species which, due to differences in morphological char- acters and colour pattern, had to be restricted to Sri Lan- ka (Koch et al. 2007). At the same time, the melanistic tax- on komaini from Thailand was identified as a junior syn- onym of V. s. macromaculatus in the absence of morpho- logical differences except for the lack of a light colour pat- tern. Therefore, the remaining subspecies of V. salvator are: Vs. salvator, V. s. macromaculatus, V. s. andamanen- sis, and V. s. bivittatus (Koch et al. 2007). Varanus cumingi Martin, 1838 1838 Varanus cumingi Martin, Proc. Zool. Soc. London 1838: 69. — Type locality: Mindanao, Philippines. 1942 Varanus (Varanus) salvator cumingi— Mertens, Abh. Senckb. Naturf. Ges., 466: 256. 2007 Varanus (Soterosaurus) cumingi — Koch, Aultya, Schmitz, Kuch & Bohme, Mertensiella, 16: 168. Bonn zoological Bulletin 57 (2): 127-136 Distribution: Islands of the Greater Mindanao region (1.e., Mindanao, Samar, Leyte, and Bohol), Philippines. Remark: Recently, % cumingi was demonstrated to be specifically distinct from V. salvator (Koch et al. 2007). The species was also shown to be polytypic and a new sub- species was described from the northern islands within the species range (Koch et al. 2010). Varanus c. cumingi Martin, 1838 1838 Varanus cumingi Martin, Proc. Zool. Soc. London 1838: 69. — Type locality: Mindanao, Philippines. 1991 Varanus salvator cumingi — Gaulke (in part), Mer- tensiella, 2: 154. Type specimen: Lectotype BMNH 1946.8.31.5, designat- ed by Koch et al. (2007). Distribution: Restricted to Mindanao and off-shore islands, Philippines. Varanus cumingi samarensis Koch, Gaulke & Bohme, 2010 (Fig. 1D) 2010 Varanus cumingi samarensis Koch, Gaulke & Bohme, Zootaxa, 2440: 19 — Type locality: San Augustin near Gandara, Samar Island, Philippines. 1991 Varanus salvator cumingi — Gaulke (in part), Mer- tensiella, 2: 161. Type specimens: Holotype ZFMK 64713, paratype ZFMK 64712. Distribution: Samar, Bohol, and Leyte, Philippines. Varanus marmoratus (Wiegmann, 1834) 1834 Hydrosaurus marmoratus Wiegmann, in Meyen, Reise um die Erde, 3: 446. — Type locality: San Mat(h)eo village or Talim Island, Laguna Bay, near Manila, Luzon, Philippines. 1829 M[onitor| marmoratus Cuvier, Regne animal 2(2): 26. (nomen nudum fide Mertens 1942; Good et al. 1993). 1844 Monitor bivittatus philippensis Schlegel, Abb. Am- phib.: x. — Type locality: Manila, Luzon. ©ZFMK 130 André Koch et al. Fig. 1. Fig. 1: Some monitor lizards described after Bohme’s (2003) checklist and species where the taxonomic status has changed since. A: Varanus boehmei Jacobs, 2003 (photo T. Ziegler); B: V. nuchalis, revalidated species status (photo M. Gaulke); C: V. to- gianus, revalidated species status (photo A. Koch); D: V. cumingi samarensis Koch, Gaulke & Béhme, 2010 (photo M. Gaulke); E: V. rasmusseni Koch, Gaulke & Bohme, 2010, juvenile paratype ZFMK 89391 (photo A. Koch); F: V. lirungensis Koch, Arida, Schmitz, Bohme & Ziegler, 2009 (photo M. Auliya); G: V. palawanensis Koch, Gaulke & Béhme, 2010 (photo I. Langlotz). Bonn zoological Bulletin 57 (2): 127-136 ©ZFMK Update of BGhme’s (2003) checklist of monitor lizards 131 1876 Varanus manilensis von Martens, PreukB. Exped. Os- tas. Zool., 1: 196. (Lapsus fide Mertens 1942). 1942 Varanus (Varanus) salvator marmoratus — Mertens, Abh. Senckb. Naturf. Ges., 466: 254. 1944 Varanus salvator philippinensis Deraniyagala, Spol. Zeylan., 24: 61. — Type locality: Luzon. 1997 Varanus (Soterosaurus) salvator marmoratus — Ziegler & Boéhme, Mertensiella, 8: 177. 2007 Varanus (Soterosaurus) marmoratus — Koch, Auliya, Schmitz, Kuch & Boéhme, Mertensiella, 16: 161. Type specimen: Lectotype ZMB 470, designated by Mertens (1942). Distribution: Restricted to Luzon and some off-shore is- lands, Philippines. Remark: Originally, Wiegmann (1834) based his descrip- tion on two voucher specimens (Koch et al. 2007). The second larger syntype, however, which should have para- lectotype status, is missing (Good et al. 1993). Recently, V. marmoratus was shown to represent a collective species (Koch et al. 2010). The disjunct island populations of the Greater Palawan region and the Sulu Archipelago were al- located to two new species (see below). Varanus nuchalis (Giinther, 1872) (Fig. 1B) 1872 Hydrosaurus nuchalis Ginther, Proc. Zool. Soc. London, 1872: 145. — Type locality: Philippines. 1942 Varanus (Varanus) salvator nuchalis — Mertens, Abh. Senckb. Naturf. Ges., 466: 258. 1997 Varanus (Soterosaurus) salvator nuchalis — Ziegler & Bohme, Mertensiella, 8: 177. 2007 Varanus (Soterosaurus) nuchalis — Koch, Aultya, Schmitz, Kuch & Bohme, Mertensiella, 16: 165. Type specimen: Holotype BMNH 1946.9.1.17. Distribution: Islands of Negros, Panay, Masbate, Ticao, and Cebu, Philippines. Remark: Despite a high variation in colour pattern, a re- cent study could not document a correlation between colour pattern and distribution (Koch et al. 2010). Bonn zoological Bulletin 57 (2): 127-136 Varanus palawanensis Koch, Gaulke & Bohme, 2010 (Fig. 1G) 2010 Varanus palawanensis Koch, Gaulke & Boéhme, Zootaxa, 2446: 33.—Type locality: Tabon, Palawan Island, Philippines. 1942 Varanus (Varanus) salvator marmoratus — Mertens (in part), Abh. Senckb. Naturf. Ges., 466: 254. 1991 Varanus salvator marmoratus — Gaulke (in part), Mertensiella, 2: 154. 2007 Varanus (Soterosaurus) marmoratus — Koch, Aultya, Schmitz, Kuch & Bohme (in part), Mertensiella, 16: 161. Type specimens: Holotype SMF 73912, paratypes SMF 73914-15, BMNH 94.6.30.19, BMNH 94.6.30.20, MNHN 1884-187, ZMUC E78, and ZFMK 89691 (for- merly SMF 73913). Distribution: Islands of Greater Palawan (Palawan, Bal- abac and the Calamian Island group) and Sibutu Island within the Sulu Archipelago, Philippines. Remark: Traditionally, the populations of Palawan and ad- jacent islands were allocated to V. marmoratus, but recent investigations confirmed their morphological distinctness (Koch et al. 2010). Varanus rasmusseni Koch, Gaulke & Bohme, 2010 (Fig. 1E) 2010 Varanus rasmusseni Koch, Gaulke & Bohme, Zoo- taxa, 2446: 28. — Type locality: Tarawakan, north of Ba- tu-Batu, Tawi-Tawi Island, Sulu Archipelago, Philippines. 1992 Varanus salvator marmoratus — Gaulke (in part), Ha- madryad, 17: 21. 2007 Varanus (Soterosaurus) cf. marmoratus — Koch, Au- liya, Schmitz, Kuch & Bohme, Mertensiella, 16: 163. Type specimens: Holotype ZMUC R42151, paratype ZFMK 89391 (formerly ZMUC R42153). Distribution: Only known from the type locality, but prob- ably also on other islands of the Tawi-Tawi island group, Philippines. Remark: Recent morphological investigations demonstrat- ed the specific distinctness of the Tawi-Tawi population which was formerly assigned to X marmoratus (Koch et al. 2010). ©ZFMK 132 André Koch et al. Varanus togianus (Peters, 1872) (Fig. 1C) 1872 Monitor (Hydrosaurus) togianus Peters, Monatb. Kon. Preuss. Akad. Wiss., 1872: 582. — Type locality: Tim- otto, Togian (= Togean) Islands, Central Sulawesi, Indone- sia. 1942 Varanus (Varanus) salvator togianus — Mertens, Abh. Senckb. Naturf. Ges., 466: 253. 1997 Varanus (Soterosaurus) salvator togianus — Ziegler & Bohme, Mertensiella, 8: 177. 2007 Varanus (Soterosaurus) togianus — Koch, Auliya, Schmitz, Kuch & Bohme, Mertensiella, 16: 156. Type specimens: Lectotype ZMB 7388, paralectotype ZMB 7389, by designation of Mertens (1942). Distribution: Sulawesi, except the northern peninsula. Remark: Recent investigations revealed this endemic Su- lawesi taxon to be specifically distinct from V. salvator (Koch et al. 2007) and polytypic (Koch et al. unpubl. da- ta). Subgenus Euprepiosaurus Fitzinger, 1843 Varanus indicus species group Varanus lirungensis Koch, Arida, Schmitz, Bohme & Ziegler, 2009 (Fig. 1F) 2009 Varanus lirungensis Koch, Arida, Schmitz, Bohme & Ziegler, Austr. J. Zool., 57: 33. — Type locality: near Lirung, Salibabu Island, Talaud Islands, Indonesia. 1915 Varanus indicus — de Rooj (in part), Rept. Indo-Aus- tr. Arch., 1: 149. 1942 Varanus (Varanus) indicus indicus — Mertens (in part), Abh. Senckb. Naturf. Ges., 466: 263. Type specimens: Holotype MZB Lac. 5178, paratypes MZB Lac. 5176-77, 5179-80, ZFMK 87587 (formerly ZMA 15411a), ZMA 15411b. Distribution: Only known from the type locality. Remark: V. lirungensis represents the most north-western member of the V. indicus species group (Koch et al. 2009). Bonn zoological Bulletin 57 (2): 127-136 Varanus obor Weijola & Sweet, 2010 2010 Varanus obor Weijola & Sweet, Zootaxa, 2434: 18. — Type locality: Soela-Bési (= Sanana Island), Sula Is- lands, Moluccas, Indonesia. Type specimen: Holotype RMNH 7225. Distribution: Only known from the type locality. Remark: V. obor represents the latest discovery of a mem- ber of the V. indicus species group. Nothing is known about its conservation status (Weijola & Sweet 2010). Varanus rainerguentheri Ziegler, Bbhme & Schmitz, 2007 2007 Varanus rainerguentheri Ziegler, Bohme & Schmitz, Mitt. Mus. Nat.kd. Berlin, Suppl. 83: 110. — Type locali- ty: Jailolo, Halmahera Island, Moluccas, Indonesia. 2005 Varanus cf. indicus — Bohme & Ziegler, Salaman- dra, 41: 57. Type specimens: Holotype ZFMK 85404, paratype USNM 237438. Distribution: Northern Moluccan islands of Halmahera, Ternate, Tidore, Morotai, Bacan, Gebe and Obi. Remark: Originally, V. rainerguentheri was only known from its type locality (Ziegler et al. 2007a), but recent field studies showed that this species occurs over a wider range in the Moluccas (Weijola 2010). Varanus zugorum Bohme & Ziegler, 2005 2005 Varanus zugorum Bohme & Ziegler, Salamandra, 41(1/2): 52. — Type locality: Kampung Pasir Putih, Jailo- lo district, Halmahera Island, Moluccas, Indonesia. Type specimen: Holotype USNM 237439. Distribution: Only known from the type locality. Remark: V. zugorum appears to be the rarest or at least known monitor lizard of all species described, known on- ly from the holotype specimen. Recent field work on Halmahera Island failed to record this secretive species (Setiadi & Hamidy 2006; Weijola 2010; Awal Riyanto, Bogor, pers. comm. viii.2010). Only one putative photo- graph of a live specimen exists (see BOhme & Ziegler 2005). ©ZFMK Update of B6hme’s (2003) checklist of monitor lizards 13 Varanus prasinus species group Varanus beccarii (Doria, 1874) 1874 Monitor Beccarii Doria, Ann. Mus. Civ. Stor. Nat. Genova, 6: 331.—Type locality: Wokam, Aru Islands, In- donesia. 1942 Varanus (Odatria) prasinus beccarii — Mertens, Abh. Senckb. Naturf. Ges., 466: 296. 2003 Varanus (Euprepiosaurus) prasinus beccarii — Bohme, Zool. Verh., 341: 25. 2007 Varanus (Euprepiosaurus) beccarii — Ziegler, Schmitz, Koch & Bohme, Zootaxa, 1472: 15. Type specimens: Syntypes ZMB 7993, MSNG 28723. Distribution: Restricted to the Aru Islands. Remark: In the past, the taxon beccarii was considered a subspecies of V. prasinus (e.g. Mertens 1942; Ziegler & Bohme 1997). Recently, Ziegler et al. (2007b) demonstrat- ed that V. beccarii is distinct from the latter species. The species is potentially threatened by the international trade in live specimens. Varanus boehmei Jacobs, 2003 (Fig. 1A) 2003 Varanus boehmei Jacobs, Salamandra, 39(2): 66. — Type locality: Waigeo Island, West Papua, Indonesia. Type specimens: Holotype ZFMK 77837, paratypes ZFMK 82826, ZFMK 84000, ZMA 21702 (formerly ZFMK 79122) and three further specimens which were still alive and will be deposited in ZFMK after their de- mise. Distribution: Only known from the type locality. Remark: Due to its restricted distribution range and its ex- ploitation for the international pet trade, Vv boehmei must be considered threatened. Varanus reisingeri Eidenmiller & Wicker, 2005 2005 Varanus reisingeri Eidenmiller & Wicker, Sauria, 27(1): 4. — Type locality: Insel Misol (= Misool Island) off the west coast of West Papua, New Guinea, Indone- sia. Bonn zoological Bulletin 57 (2): 127-136 Ww 1942 Varanus (Odatria) prasinus prasinus — Mertens (in part), Abh. Senckb. Naturf. Ges., 466: 292. Type specimens: Holotype SMF 83679, the two paratypes are still alive and will be deposited in SMF after their de- mise (Bernd Eidenmiller, Frankfurt, pers. comm. x11.2010). Distribution: Only known from the type locality. Remark: The taxonomic validity of this species remains uncertain because diagnostic morphological characters largely overlap with V. prasinus from New Guinea. As the former species, V. reisingeri is also potentially threatened by exploitation for the pet trade. Subgenus Odatria Gray, 1838 Varanus bushi Aplin, Fitch & King, 2006 2006 Varanus bushi Aplin, Fitch & King, Zootaxa, 1313: 24. — Type locality: Marandoo, Western Australia (22° S75 S08 4E): 1980 Varanus caudolineatus — Storr (in part), Rec. West. Austr. Mus., 8: 250. Type specimens: Holotype WAM R108999, paratypes WAM R54230, WAM R56834, and WAM R62171. Distribution: Endemic to the Pilbara region of Western Australia. Remark: V. bushi was described as morphologically and genetically distinct from its closest relatives V. caudoline- atus and V. gilleni. All three Australian dwarf monitor lizards display complex patterns of sexual dimorphism (Aplin et al. 2006). Subgenus: Incertae sedis Varanus spinulosus Mertens, 1941 1941 Varanus indicus spinulosus Mertens, Senckenber- giana, 23: 269. — Type locality: Georgs-Insel (= St. George Island or San Jorge Island), near Santa Isabel (= Ysabel) Island, Solomon Islands. 1942 Varanus (Varanus) indicus spinulosus — Mertens, Abh. Senckb. Naturf. Ges., 466: 271. 1994 Varanus spinulosus — Sprackland, Herpetofauna, 24 (2): 34. ©ZFMK 134 André Koch et al. 1997 Varanus (Euprepiosaurus) spinulosus — Ziegler & Bohme, Mertensiella, 8: 14. 2007 Varanus (subgen. inc. sed.) spinulosus — Bohme & Ziegler, Mertensiella, 16: 105. Type specimen: Holotype NMW 23387 (formerly NMW 3709). Distribution: San Jorge and Santa Isabel Islands, Solomon Islands, and Bougainville Island, Papua New Guinea. Remark: For almost 50 years, this rare monitor lizard species was only known from the holotype (Sprackland 1993). The former collection number of the type was orig- inally assigned by Mertens (1941, 1942) and still applied by de Lisle (2009). Tiedemann et al. (1994) and B6hme & Koch (2010) provided the current number. Sprackland (1994) elevated spinulosus to full species status. Recent- ly, the distribution range of V. spinulosus was extended, when the species was newly recorded from Bougainville Island and its occurrence was confirmed on the island of Santa Isabel (BOhme & Ziegler 2007; Dwyer 2008). VK. spinulosus was formerly allocated to the V. indicus species group of the subgenus Euprepiosaurus (see Ziegler & Bohme 1997), but a new monotypic subgenus is dis- cussed based on new genital morphological findings (Bohme & Ziegler 2007). DISCUSSION In total, ten new species and one new subspecies were in- troduced to science since BOhme’s (2003) checklist. Nine of the species (1.e., 90%) and the subspecies were de- scribed from islands of the Indo-Australian Archipelago. Only one new species, V. bushi, was recently identified from Western Australia (Aplin et al. 2006). In addition, five taxa (beccarii, cumingi, marmoratus, nuchalis, and togianus) were re-elevated to full species status due to morphological (e.g., scale counts, morphometrics, colour pattern) and/or genetical idiosyncrasies, thus bringing the global diversity to 73 (including 21 subspecies). This re- presents an increase in species diversity of 20% since 2003. Particularly 2010 has been a very productive year for the increase of monitor lizard diversity. Four new species and one new subspecies were described from In- donesia and the Philippines within the first half of 2010 (Koch et al. 2010; Weijola & Sweet 2010; Welton et al. 2010). Descriptions of new monitor lizard species since 2003 mainly refer to two different taxonomic groups, the South- Bonn zoological Bulletin 57 (2): 127-136 east Asian and Indo-Australian subgenera Soferosaurus and Euprepiosaurus, with two and six new species, respec- tively. In addition, the taxonomic status of several mem- bers of the subgenus Soferosaurus has changed. While this subgenus was hitherto considered monotypic with V. sal- vator being the only, albeit polytypic species with eight recognized subspecies (B6hme 2003), the Philippine sub- species cumingi, marmoratus, and nuchalis, and togianus from Sulawesi were re-elevated to their original species status (Koch et al. 2007), thus resulting in a species com- plex of closely related allies. Additionally, one subspecies (V. salvator macromaculatus) of the Southeast Asian wa- ter monitor lizard was revalidated, whereas the younger melanistic taxon komaini was synonymized with the for- mer (Koch et al. 2007). With currently 22 recognized species, the subgenus Eu- prepiosaurus has displaced the Australian Odatria as the most species-rich subgenus of varanids. Within Eupre- piosaurus, the new species descriptions are unevenly dis- tributed over the two species groups involved, viz. the Pa- cific monitors around V. indicus and the tree monitors around V. prasinus, respectively. The latter group experi- enced only two new species descriptions (1.e., v boehmei and V. reisingeri) and currently comprises nine allopatric species from New Guinea and its offshore islands. On the other hand, four new species were added to the V. indicus species group leading to a total of 13 recognized species, at least four of which occur in sympatry on New Guinea and Halmahera in the northern Moluccas. Among these recently described Pacific monitor lizard species, next to morphologically cryptic taxa, such as V. lirungensis or V. rainerguentheri, there are also strikingly different species with idiosyncratic features in morphology and colour pat- tern, such as the melanistic V. obor and the silver-coloured V. zugorum. Five years after its formal description and de- spite repeated field trips to the northern Moluccas (Setia- di & Hamidy 2006; Weijola 2010; Awal Riyanto, Bogor, pers. comm. viti.2010), the latter species is still only known from the holotype specimen and has thus to be re- garded the rarest and at least known varanid species. Bohme (2003) already recognized a taxonomic trend to- wards a reduction of polytypic monitor lizard species by the elevation of nominal subspecies to species rank. Cer- tainly, this trend still continues as seen in the V. salvator complex (Koch et al. 2007). The description of a new sub- species of V. cumingi from the Philippines (Koch et al. 2010), however, demonstrates that a distinction is still made between the degree of morphological (e.g., morpho- metrics and scalation features) and molecular differenti- ation (i.e. characteristics of the full species category) and mere geographically correlated differences in colour pat- tern (i.e. diagnostic features of subspecies). ©ZFMK Update of B6hme’s (2003) checklist of monitor lizards 135 CONCLUSIONS Because taxonomy is a dynamic discipline, further changes and additions to the list of extant monitor lizards are to be expected in the future. This will include new species descriptions — either real discoveries or by the splitting of already recognized species — as well as a change of the taxonomic status. Therefore, we are aware that this updated checklist can only represent the latest state of art and may already be outdated by the time of publication. In terms of conservation purposes, it is essen- tial to refer to these most recent checklists, and with the increase of monitor lizard diversity especially in the In- do-Australian realm, there is definitely a need to estab- lish user-friendly identification tools for a vertebrate group globally sought after within the international pet and rep- tile leather trade. We do wish that our contribution will serve as a useful supplement to the checklist of the living monitor lizards of the world by B6hme (2003) for all those who are in- terested in or concerned with the diversity of monitor lizards. Acknowledgments. The fact, that Wolfgang Bohme was in- volved in most of the recent taxonomic changes and additions to the global checklist of varanids together with the naming of one new species, viz. V. boehmei, after him, reflect his interna- tionally renowned competence and eminent position in syste- matic monitor lizard research. On the occasion of his retirement, we hereby wish to dedicate this contribution to Professor Dr. Wolfgang Bohme, Vice Director, Head of the Vertebrate Depart- ment and Curator of Herpetology for 39 years at the Zoologi- sches Forschungsmuseum Alexander Koenig in Bonn, Germany. We are deeply indebted to Wolfgang Béhme, who initiated and supervised our careers related to varanid research. Furthermore, we thank Gunter KG6hler (SMF, Frankfurt), Bernd Eidenmiiler (Frankfurt), and Manfred Reisinger (Landshut), for information on preserved and live type specimens in their care. Awal Riyanto (MZB, Bogor) kindly shared his observations on monitor lizards from the field. The helpful comments of two anonymous referees improved an earlier version of this contri- bution. REFERENCES Aplin KP, Fitch AJ, King DJ (2006) A new species of Varanus Merrem (Squamata: Varanidae) from the Pilbara region of Western Australia, with observations on sexual dimorphism in closely related species. 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VII.2010 Accepted: 21.X.2010 ©ZFMK Bonn zoological Bulletin Volume 57 Issue 2 pp. 137-147 Bonn, November 2010 New discoveries of amphibians and reptiles from Vietnam Thomas Ziegler! & Truong Quang Nguyen23 1 Cologne Zoo, Riehler StraBe 173, D-50735 Cologne, Germany; E-mail: ziegler@koelnerzoo.de 2 Institute of Ecology and Biological Resources, 18 Hoang Quoc Viet, Hanoi, Vietnam; E-mail: nqt2@yahoo.com 3 Zoologisches Forschungsmuseum Alexander Koenig, Adenauerallee 160, D-53113 Bonn, Germany Abstract. We provide a list of 21 new amphibian and reptilian species and subspecies discoveries from Vietnam, includ- ing one new snake genus, published after the comprehensive overview by Nguyen et al. (2009). The new herpetofauna representatives are introduced inclusive of the original description, type locality, English and Vietnamese names, as well as current distribution. Key words. Vietnam, herpetofauna, new species. INTRODUCTION Although Vietnam has one of the world’s richest amphib- ian and reptilian fauna, as revealed particularly through surveys by Vietnamese scientists and their international collaborators during the past quarter century, the study of its herpetofauna was long overshadowed by research in India, China, and the East Indies (Adler 2009). The first significant summary of the Vietnamese herpetofauna was written by Morice (1875), in which 13 amphibians and 114 species of reptiles including the marine species were list- ed. Ten years later, Tirant (1885) published a 104-pages book containing 166 species of herpetofauna from Viet- nam and Cambodia. Subsequently, the following staff members or associates of the Muséum d’Histoire Na- turelle, Paris, published specific studies of Vietnamese am- phibians and reptiles: Léon Vaillant (1834-1914), Francois Mocquard (1834-1917), Jacques Pellegrin (1873-1944), Paul Chabanaud (1876-1959), Fernand Angel (1881-1950), and René Bourret (1884-1957), who was the only one of the afore mentioned zoologists who ever set foot in Vietnam (Adler 2009). It was René Bourret, originally a geologist, who became the leading authority on Vietnamese herpetofauna. From 1927 until 1947 he published a series of papers and books Bonn zoological Bulletin 57 (2): 137-147 of Indochinese and specifically on Vietnamese herpetol- ogy. Besides the publication of several identification man- uals, he is most famous for a substantial series of mono- graphs of Indochinese herpetology, which are to date the most important background on the subject (Bourret 1936a, b, 1941, 1942). In total, René Bourret reported of 177 lizard taxa (1.e., species and subspecies), 245 snake taxa, 45 turtle taxa and 171 amphibian taxa for the Indochinese region (Nguyen 2006). Because of the French Indochina War, no remarkable her- petological studies were undertaken in the period from 1946 to 1954. In 1954, when northern Vietnam attained independence from France, Vietnamese herpetologists be- gan to conduct herpetological field surveys predominant- ly in northern Vietnam, and the first lists and keys to the species of Vietnamese amphibians and reptiles were com- piled by Dao (e.g., 1977, 1978, 1979, 1981, 1982), includ- ing 363 species in total, but with inaccuracies. Another noteworthy key to snakes of southern Vietnam was writ- ten during the Second Indochina War, better known in the West as Vietnam War, by Campden-Main (1970), who had served several years as a medic with American forces sta- tioned in Vietnam. ©ZFMK 138 Thomas Ziegler & Truong Quang Nguyen The end of the Vietnam War in 1975 marked the begin- ning of another period of biodiversity research in Vietnam (Sterling et al. 2006). Thereafter, an increasing engage- ment in herpetological field work mainly by Russian in- stitutions was observable, which is still persistent. Even- tually, in the last 30 years many international herpetolog- ical cooperations emerged, which led to an enormous in- crease in new records and species descriptions from Viet- nam (see overview in Nguyen 2006). As a consequence, the updated checklist by Nguyen et al. (2005) comprised 458 species, viz., 162 species of amphibians and 296 species of reptiles, which included more than 100 addi- tional species compared with the previous checklist of the herpetofauna of Vietnam by Nguyen & Ho (1996). Fur- thermore, the most actual list (Nguyen et al. 2009) cov- ers 12 additional species of amphibians and 64 addition- al species of reptiles compared with Nguyen et al. (2005). On the occasion of this Proceedings Volume addicted to Professor Dr. Wolfgang Béhme, Vice Director, Head of the Vertebrate Department and Curator of Herpetology at the Zoological Research Museum Alexander Koenig, Bonn, Germany, we provide a list of new amphibian and reptilian discoveries from Vietnam that were published subsequent to the comprehensive overview provided by Nguyen et al. (2009). We would like to dedicate this pa- per to Wolfgang Bohme, who supervised the PhD theses of both authors (T. Ziegler: 1997-2000; T.Q. Nguyen 2007-2011) and thus decisively brought forward herpeto- diversity research in Vietnam. MATERIAL AND METHODS We herein compiled species’ descriptions that were for- mally published after the appearance of Nguyen et al. (2009). We therefore followed the style and taxonomic arrangement provided by the latter authors. Abbreviations are as follows: AMNH = American Muse- um of Natural History, New York, USA; AMS = Australian Museum, Sydney, Australia; IEBR = Institute of Ecolo- gy and Biological Resources, Hanoi, Vietnam; ITBCZ = Institute of Tropical Biology, Collection of Zoology, Ho Chi Minh City, Vietnam; LSUHC = La Sierra Universi- ty, Herpetological Collection, La Sierra University, River- side, California, USA; UNS = Zoological Collection of the University of Natural Sciences, Ho Chi Minh City, Vietnam; VNMN = Vietnam National Museum of Nature, Hanoi, Vietnam; ZFMK = Zoologisches Forschungsmu- seum Alexander Koenig, Bonn, Germany; ZISP = Zoo- logical Institute, St. Petersburg, Russia; a.s.]. = above sea level. Bonn zoological Bulletin 57 (2): 137-147 LIST OF NEW SPECIES AND SUBSPECIES SINCE NGUYEN ET AL. (2009) Amphibia Anura Megophryidae Leptolalax applebyi Rowley & Cao, 2009 Leptolalax applebyi J.J.L. Rowley & T.T. Cao, 2009, Zoo- taxa 2198: 52. Holotype: AMS R171703. Type locality: Song Thanh Proposed Nature Reserve, Phouc Son (Phuoc Son) District, Quang Nam Province, Vietnam, 1,402 ma.s.l. English name: Appleby’s Asian Toad. Vietnamese name: Coc may ap-li-bai. Distribution: This species is currently known only from the type locality. Ranidae Odorrana geminata Bain, Stuart, Nguyen, Che & Rao, 2009 Odorrana geminata R.H. Bain, B.L. Stuart, T.Q. Nguyen, J. Che & D.-Q. Rao, 2009, Copeia 2: 355. Holotype: AMNH 163782. Type locality: Mount Tay Con Linh II, Cao Bo Commune, Vi Xuyen District, Ha Giang Province, Vietnam, 1,420 m a.s.l. English name: Geminated Cascade Frog. Vietnamese name: Ech bam da hoa. Distribution: This species (Fig. 1a) is currently known on- ly from montane areas in northeastern Vietnam (Ha Gi- ang and Cao Bang provinces) and southeastern Yunnan Province, China. ©ZFMK New amphibians and reptiles from Vietnam 139 Fig. 1. Rhacophoridae Theloderma lateriticum Bain, Nguyen & Doan, 2009 Theloderma lateriticum R.H. Bain, T.Q. Nguyen & K.V. Doan, 2009, Zootaxa 2191: 60. Holotype: AMNH 168757/IEBR A. 0860. Type locality: Nam Tha Commune, Van Ban District, Lao Cai Province, Vietnam, 1,300—1,400 m a.s.l. English name: Brick-red Bug-eyed Frog. Vietnamese name: Ech cay san do. Distribution: This species (Fig. 1b) is currently known on- ly from the type locality. Bonn zoological Bulletin 57 (2): 137-147 a) Odorrana geminata from Ha Giang Province, Photo T.Q. Nguyen; b) Theloderma lateriticum from Lao Cai Province, Photo T.Q. Nguyen; c) Leiolepis ngovantrii from Ba Ria—Vung Tau Province, Photo L.L. Grismer; and d) Pseudocalotes ziegleri from Kon Tum Province, Photo C.T. Ho. Reptilia Squamata Sauria Agamidae Leiolepis ngovantrii Grismer & Grismer, 2010 Leiolepis ngovantrii J.L. Grismer & L.L. Grismer, 2010, Zootaxa 2433: 52. Holotype: LSUHC 9234. Type locality: Binh Chau—Phuoc Buu Nature Reserve, Xuyen Moc District, Ba Ria—Vung Tau Province, Vietnam, 30 ma.s.l. ©OZFMK 140 Thomas Ziegler & Truong Quang Nguyen Fig. 2. English name: Ngovantri’s Butterfly Lizard. Vietnamese name: Nhong cat ngo van tri. Distribution: This species (Fig. 1c) is currently known on- ly from Vietnam (Ba Ria—Vung Tau Province). Pseudocalotes ziegleri Hallermann, Nguyen, Orlov & Ananjeva, 2010 Pseudocalotes ziegleri J. Hallermann, T.Q. Nguyen, N. Orlov & N. Ananjeva, 2010, Russ. J. Herpetol. 17(1): 32. Holotype: IEBR 330. Type locality: Nuoc Ka forest, near Mang Canh, Kon Plong District, Kon Tum Province, Vietnam, ca. 1,200 m a.s.l. English name: Ziegler’s Tree Lizard. Vietnamese name: Nhong zig-lo. Distribution: This species (Fig. 1d) is currently known on- ly from Vietnam (Kon Tum Province). Bonn zoological Bulletin 57 (2): 137-147 a) Cnemaspis psychedelica from Ca Mau Province, Photo L.L. Grismer; b) Cyrtodactylus cattienensis from Dong Nai Province, Photo P. Geissler; c) Cyrtodactylus roesleri from Quang Binh Province, Photo T. Ziegler; and d) Dixonius aaronbaueri from Ninh Thuan Province, Photo T.V. Ngo. Remarks: Specimens identified as Pseudocalotes floweri from Kon Tum Province (Bain et al. 2007) were subse- quently re-identified as P. ziegleri by Hallermann et al. (2010). Gekkonidae Cnemaspis psychedelica Grismer, Ngo & Grismer, 2010 Cnemaspis psychedelica L.L. Grismer, T.V. Ngo & J.L. Grismer, 2010, Zootaxa 2352: 48. Holotype: UNS 0444. Type locality: Hon Khoai Island, Ngoc Hien District, Ca Mau Province, Vietnam. English name: Psychedelic Gecko. Vietnamese name: Tac ke duoi vang. Distribution: This species (Fig. 2a) is currently known on- ly from the type locality. ©OZFMK New amphibians and reptiles from Vietnam 14] Cyrtodactylus cattienensis Geissler, Nazarov, Orlov, Bohme, Phung, Nguyen & Ziegler, 2009 Cyrtodactylus cattienensis P. Geissler, R. Nazarov, N.L. Orlov, W. Bohme, T.M. Phung, T.Q. Nguyen & T. Ziegler, 2009, Zootaxa 2161, 21. Holotype: IEBR A.0856. Type locality: Cat Tien National Park, Dong Nai Province, Vietnam, 120 m a.s.1. English name: Cattien Bent-toed Gecko. Vietnamese name: Thach sung ngon cat tien. Distribution: This species (Fig. 2b) is currently known on- ly from Vietnam (Ba Ria-Vung Tau and Dong Nai provinces). Cyrtodactylus roesleri Ziegler, Nazarov, Orlov, Nguyen, Vu, Dang, Dinh & Schmitz, 2010 Cyrtodactylus roesleri T. Ziegler, R. Nazarov, N. Orlov, T.Q. Nguyen, T.N. Vu, K.N. Dang, T.-H. Dinh & A. Schmitz, 2010, Zootaxa, 2413: 24. Holotype: ZFMK 89377. Type locality: Phong Nha—Ke Bang National Park, Minh Hoa District, Quang Binh Province, Vietnam. English name: Roesler’s Bent-toed Gecko. Vietnamese name: Thach sung ngon ro-x-lo. Distribution: This species (Fig. 2c) is currently known on- ly from the type locality. Cyrtodactylus yangbayensis Ngo & Chan, 2010 Cyrtodactylus yangbayensis T.V. Ngo & K.O. Chan, Zootaxa, 2504: 48. Holotype: UNS 0476. Type locality: Yang Bay Waterfall, Dien Khanh District, Khanh Hoa Province, southern Vietnam, 500-600 m a.s.1. English name: Yangbay Bent-toed Gecko. Vietnamese name: Than lan chan ngon yang bay. Bonn zoological Bulletin 57 (2): 137-147 Distribution: This species is currently known only from the type locality in Khanh Hoa Province. Dixonius aaronbaueri Ngo & Ziegler, 2009 Dixonius aaronbaueri T.V. Ngo & T. Ziegler, 2009, Zoosyst. Evol., 85(1): 119. Holotype: UNS 0284. Type locality: Binh Tien Forest Station, Ninh Hai District, Nui Chua National Park, Ninh Thuan Province, southern Vietnam, 4-5 m a.s.1. English name: Aaron Bauer’s Leaf-toed Gecko. Vietnamese name: Than lan chan la a-ron-bau-o. Distribution: This species (Fig. 2d) is currently known on- ly from the type locality. Gekko canhi Rosler, Nguyen, Doan, Ho, Nguyen & Ziegler, 2010 Gekko canhi H. Résler, T.Q. Nguyen, K.V. Doan, C.T. Ho, T.T. Nguyen & T. Ziegler, 2009, Zootaxa 2329: 57. Holotype: IEBR A.0910. Type locality: Huu Lien, Huu Lung, Lang Son Province, North Vietnam. English name: Canh’s Gecko. Vietnamese name: Tac ke canh. Distribution: This species (Fig. 3a) is currently known on- ly from northern Vietnam (Lang Son and Lao Cai provinces). Gekko russelltraini Ngo, Bauer, Wood & Grismer, 2009 Gekko russelltraini T.V. Ngo, A.M. Bauer, P.L. Jr. Wood & J.L. Grismer, 2009, Zootaxa 2238: 34. Holotype: UNS 0293. Type locality: Chua Chan Mountain, Suoi Cat Commune, Xuan Loc District, Dong Nai Province, Vietnam, ca. 100 m a.s.l. English name: Russell Train’s Marble Gecko. ©ZFMK 142 Thomas Ziegler & Truong Quang Nguyen Fig. 3. a) Portrait of preserved Gekko canhi from Lang Son Province, Photo T. Ziegler; b) Gekko russelltraini from Dong Nai Province, Photo T.V. Ngo; c) Gekko takouensis from Binh Thuan Province, Photo S.N. Nguyen; and d) Gekko vietnamensis from An Giang Province, Photo S.N. Nguyen. Vietnamese name: Than lan da ru-xen-tren. Distribution: This species (Fig. 3b) is currently known on- ly from Vietnam (Dong Nai Province). Gekko takouensis Ngo & Gamble, 2010 Gekko takouensis T.V. Ngo & T. Gamble, 2010, Zootaxa 2346: 18. Holotype: UNS 0491. Type locality: Ta Kou Mountain, Ham Thuan Nam Dis- trict, Binh Thuan Province, Vietnam, 425 m a.s.l. English name: Takou Marbled Gecko. Vietnamese name: Than lan da ta kou. Distribution: This species (Fig. 3c) is currently known on- ly from the type locality. Bonn zoological Bulletin 57 (2): 137-147 Gekko vietnamensis Nguyen, 2010 Gekko vietnamensis S.N. Nguyen, 2010, Zootaxa 2501: 55: Holotype: ITBCZ 667. Type locality: Tuc Dup Hill, An Giang Province, south- ern Vietnam, 43 m a.s.1. English name: Vietnam Gecko Vietnamese name: Tac ke viet nam. Distribution: This species (Fig. 3d) is currently known on- ly from the type locality. Scincidae Scincella apraefrontalis Nguyen, Nguyen, BOhme & Ziegler 2010 Scincella apraefrontalis T.Q. Nguyen, S.V. Nguyen, W. Bohme & T. Ziegler, 2010, Folia. Zool., 59(2): 116. ©ZFMK New amphibians and reptiles from Vietnam 143 Holotype: IEBR A.0832. Type locality: Huu Lien Nature Reserve, Huu Lung Dis- trict, Lang Son Province, Vietnam, ca. 200 m a.s.]. English name: Huulien Ground Skink. Vietnamese name: Than lan co huu lien. Distribution: This species is only known from Lang Son Province, northeastern Vietnam. Tropidophorus boehmei Nguyen, Nguyen, Schmitz, Orlov & Ziegler, 2010 Tropidophorus boehmei T.Q. Nguyen, T.T. Nguyen, A. Schmitz, N.L. Orlov & T. Ziegler, 2010, Zootaxa 2439: D7, Holotype: VNMN 822. Type locality: Hoang Lien Mountain, near Ban Khoang, Sa Pa District, Lao Cai Province, northern Vietnam, 1,200—1,300 m a.s.l. English name: Boehme’s Water Skink Vietnamese name: Than lan tai boe-me Distribution: This species (Fig. 4a) is currently known on- ly from Hoang Lien Mountain in Sa Pa and Van Ban dis- tricts, Lao Cai Province, Vietnam. Serpentes Colubridae Calamaria gialaiensis Ziegler, Nguyen & Nguyen, 2008 Calamaria gialaiensis T. Ziegler, S.V. Nguyen & T.Q. Nguyen, 2008, Current Herpetol., 27(2): 72. Holotype: IEBR A.0714. Type locality: Kon Ka Kinh, K Bang District, Gia Lai Province, Vietnam, 1300 m a.s.l. English name: Gialai Reed Snake. Vietnamese name: Ran mai gam gia lai. Distribution: This species (Fig. 4b) is currently known on- ly from Gia Lai Province, Vietnam. Bonn zoological Bulletin 57 (2): 137-147 Calamaria sangi Nguyen, Koch & Ziegler, 2010 (2009) Calamaria sangi T.Q. Nguyen, A. Koch & T. Ziegler, 2010 “2009”, Hamadryad 34(1): 2. Holotype: IEBR 360. Type locality: Mang Canh Commune, Kon Plong District, Kon Tum Province, Vietnam, 1,200 m a.s.1. English name: Sang’s Reed Snake. Vietnamese name: Ran mai gam sang. Distribution: This species is currently known only from Vietnam. Colubroelaps nguyenvansangi Orlov, Kharin, Ananje- va, Nguyen & Nguyen, 2009 Colubroelaps nguyenvansangi N.L. Orlov, V.E. Kharin, N.B. Ananjeva, T.T. Nguyen & T.Q. Nguyen, 2009, Russ. J- Herpetol. 16(3): 235. Holotype: ZISP/IEBR 25682. Type locality: Loc Bac Forest Enterprise, Lam Dong Province, Vietnam, ca. 720 m a.s.]. English name: Nguyenvansang’s Snake. Vietnamese name: Ran nguyen van sang. Distribution: The second record of this species was report- ed by N. Poyarkov from Bu Gia Map National Park, Binh Phuoc Province (Fig. 5). Therefore Colubroelaps nguyen- vansangi 1s currently known from Lam Dong and Binh Phuoc provinces, Vietnam. Lycodon ruhstrati abditus Vogel, David, Pauwels, Sumontha, Norval, Hendrix, Vu & Ziegler, 2009 Lycodon ruhstrati abditus G. Vogel, P. David, O.S.G. Pauwels, M. Sumontha, G. Norval, R. Hendrix, T.N. Vu & T. Ziegler, 2009, Tropical Zoology 22(2): 144. Holotype: ZFMK 86451. Type locality: U Bo region, Phong Nha — Ke Bang Na- tional Park, Quang Binh Province, Vietnam. English name: Hidden Mountain Wolf Snake. Vietnamese name: Ran khuyet an. ©ZFMK 144 Thomas Ziegler & Truong Quang Nguyen Fig. 4. a) Tropidophorus boehmei from Lao Cai Province, Photo T.T. Nguyen; b) Portrait of preserved Calamaria gialaiensis from Gia Lai Province, Photo T. Ziegler; and c) Protobothrops trungkhanhensis from Cao Bang Province, Photo T.T. Nguyen. SSS Osi a ee ae ~~ Fig. 5. | Colubroelaps nguyenvansangi from Binh Phuoc Province, Photo N. Poyarkov. Bonn zoological Bulletin 57 (2): 137-147 ©ZFMK New amphibians and reptiles from Vietnam 145 Dao (1977- 1982) Fb (1996) = (2005) Fig. 6. Species diversity of the herpetofauna of Vietnam. Distribution: This subspecies is currently known from Quang Binh and Vinh Phuc provinces in Vietnam, and from Fujian, Anhui, Zhejiang, Guangdong, Yunnan, Sichuan and Gansu provinces in China. Because it has of- ten been confused with other taxa, its range is probably much wider, both in northern Vietnam and China (prob- ably also present in Guangxi Province). Viperidae Protobothrops trungkhanhensis Orlov, Ryabov & Nguyen, 2009 Protobothrops trungkhanhensis N.L. Orloy, S.A. Ryabov & T.T. Nguyen, 2009, Russ. J. Herpetol. 16(1): 71. Holotype: ZISP 25351. Type locality: Trung Khanh Nature Reserve, Trung Khanh District, Cao Bang Province, Vietnam, 600 m a.s.1. English name: Trungkhanh Pitviper. Vietnamese name: Ran luc trung khanh. Distribution: This species (Fig. 4c) is currently known on- ly from the type locality. Bonn zoological Bulletin 57 (2): 137-147 ® Anphibians Navan & Nanandal Navandal July 210 (2009) DISCUSSION After the publication of the “Herpetofauna of Vietnam” by Nguyen et al. (2009) 20 new amphibian and reptilian species, one new subspecies, and a new snake genus have been described from Vietnam by June 2010. Among them there were three new amphibians (1 Megophryidae, 1 Ranidae, | Rhacophoridae) and 18 new reptilian taxa (2 Agamidae, 9 Gekkonidae, 2 Scincidae, 4 Colubridae, and 1 Viperidae). In contrast, two species which were listed as valid and occurring in Vietnam in Nguyen et al. (2009) were synonymized meanwhile: Gekko ulikovskii Darevsky & Orlov, 1994 was regarded as a junior synonym of Gekko badenii Szczerbak & Nekrasova, 1994 by Nguyen et al. (2010d) and the specimen previously identified as Pseudocalotes floweri from Kon Tum Province was re- identified as P. ziegleri by Hallermann et al. (2010). Most of the recent species’ discoveries affected lizards, with geckos clearly being the predominant group. In addition to these new species descriptions, three new country records were published after the book of Nguyen et al. (2009): one amphibian species, the megophryid anuran Leptobrachium promustache, the scincid lizard Scincella monticola, and the colubrid snake Amphiesmoides ornat- iceps (Bain et al. 2009b, Nguyen et al. 2010a, b). The results of this paper clearly exemplify that even af- ter the comprehensive book provided by Nguyen et al. ©ZFMK 146 Thomas Ziegler & Truong Quang Nguyen (2009) much research is needed to describe Vietnam’s rich herpetodiversity. In particular because not only cryptic or inconspicuous species were discovered and formally de- scribed in the past months, but also striking and colour- ful species like Cnemaspis psychedelica (Grismer et al. 2010) or even new genera, as was recently shown by the description of Colubroelaps (Orlov et al. 2009a). Current- ly, the herpetofauna of Vietnam comprises 181 species of amphibians and 385 species of reptiles (Fig. 6). Howev- er, diversity research and species inventories are only the first steps, which must be followed by investigations of the natural history and specific adaptations, which final- ly are prerequisites for adequate conservation measures. Acknowledgements. We are indebted to our colleagues Peter Geissler (Bonn), L. 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Excursions et Reconnaissance, Saigon 20 Vogel G, David P, Pauwels OSG, Sumontha M, Norval G, Hen- drix R, Vu NT, Ziegler T (2009) A revision of Lycodon ruh- strati (Fischer 1886) auctorum (Squamata Colubridae), with the description of a new species from Thailand and a new sub- species from the Asian mainland. Tropical Zoology 22(2): 131-182 Ziegler T, Nazarov R, Orlov N, Nguyen QT, Vu NT, Dang NK, Dinh HT, Schmitz A (2010) A third new Cyrtodactylus (Squa- mata: Gekkonidae) from Phong Nha-Ke Bang National Park, Truong Son Range, Vietnam. Zootaxa 2413: 20-36 Ziegler T, Nguyen VS, Nguyen QT (2008) A new reed snake of the genus Calamaria Boie (Squamata: Colubridae) from Viet- nam. Current Herpetology 27(2): 71-80 Received: 09. VI.2010 Accepted: 24. VIII.2010 ©ZFMK L Bonn zoological Bulletin Volume 57 Issue 2 pp. 149-171 Bonn, November 2010 The distinction between family-series and class-series nomina in zoological nomenclature, with emphasis on the nomina created by Batsch (1788, 1789) and on the higher nomenclature of turtles Alain Dubois & Roger Bour Reptiles & Amphibiens, UMR 7205 OSEB, Département de Systématique & Evolution, Muséum national d’ Histoire naturelle, CP 30, 25 rue Cuvier, F-75005 Paris, France; E-mails: sapo421@gmail.com, bour@mnhn. fr Abstract. The Code only regulates the scientific names or nomina of zoological taxa from the rank subspecies to the rank superfamily, but not those of taxa at ranks above the latter (class-series nomina). It is shown here that its current Rules are somewhat ambiguous regarding the availability of family-series nomina and the distinction between the latter and class-series nomina, and it is again suggested that it should be improved in this respect. It should cover the whole nomenclatural hierarchy in zoology, which requires to expand it in order to incorporate Rules for the nomenclature of higher taxa. A detailed analysis is devoted to the poorly known work of Batsch (1788, 1789), and it is shown that 17 fam- ily-series nomina, 16 of which have remained unnoticed until now, are available and should replace the homonymous junior nomina currently considered valid in zootaxonomy. Particular attention is given to the higher nomenclature of tur- tles, and it is shown that the nomen 7éSTUDINES Batsch, 1788 is a family-series, not a class-series nomen. This nomen is therefore the valid one, as TESTUDINIDAE Batsch, 1788, of the family including the genus Jestudo Linnaeus, 1758, but cannot apply to the order of turtles, tortoises and terrapins. Key words. Zoological nomenclature, Code, availability, class, family, Batsch, turtles, TESTUDINES, TESTUDINIDAE. ABBREVIATIONS AND PRINTING CONVEN- TIONS In this paper, “ICZN” designates the International Com- mission on Zoological Nomenclature and “the Code” the edition currently in force of the /nternational Code of Zo- ological Nomenclature (Anonymous 1999). For reasons explained in details elsewhere (Dubois 2000, 2006a), some terms of the Code are here substituted by other terms, as follows (in the order of their first appearance in the text, indicated there by an asterisk*): nomen (plural nomina) for “scientific name” (Dubois 2000); nominal-series for “groups of names” (Dubois 2000), with four such series (“groups”) being discussed here: the species-, genus-, fam- ily- and class-series (Dubois 2000); anoplonym for a “nomenclaturally unavailable name” (Dubois 2000); so- zonym for a nomen that has had a universal or significant use in non-systematic literature after 31 December 1899 (Dubois 2005a—b); distagmonym for a nomen that has not had such a use (Dubois 2005a—b); onomatophore for “type” or “name-bearing type” (Simpson 1940); nucle- ogenus (plural nucleogenera) for “type genus” (Dubois 2005b); nucleospecies for “type species” (Dubois 2005b); Bonn zoological Bulletin 57 (2): 149-171 monophory for “monotypy” (Dubois 2005b); neonym for “new replacement name” or “nomen novum” (Dubois 2000); autoneonym for a neonym being an “unjustified emendation” (Dubois 2000); archaeonym for the nomen replaced by a neonym (Dubois 2005a); hyponymous for “nominotypical” (Dubois 2006c). The nomina belonging to the species-series and genus-series are printed, as usu- al, in lower case italics, whereas nomina of higher-ranked taxa are printed in small capitals, with the following dis- tinction: family-series nomina are in /74LICS, whereas class-series nomina are in BOLD. Anoplonyms are print- ed “between quotation marks”. FAMILY-SERIES AND CLASS-SERIES NOMEN- CLATURE IN ZOOLOGY The Code regulates the nomina* of zoological taxa from the rank subspecies to the rank superfamily, but not those of taxa at ranks above the latter. Therefore the use and al- ©OZFMK 150 Alain Dubois & Roger Bour location of nomina of taxa referred to the higher ranks of zoological nomenclature (order, class, phylum, etc.) are left to the freedom and opinions of individual zoologists, as no Rules exist in the Code for their availability, allo- cation to taxa and validity, the three basic steps of the nomenclatural process (Dubois 2005a—c, 2006a). The nomina covered by the Code are distributed in three nominal-series*: the species-*, genus-* and family-se- ries*. Any nomen, to be recognized as nomenclaturally available, must first be explicitly or implicitly referred to one of these nominal-series. No difficulty usually arises regarding the allocation of nomina to the species- and genus-series, but, in some cases, problems may be encoun- tered to know whether a given nomen belongs in the fam- ily-series, and thus is governed by the nomenclatural Rules of the Code, or to a rank above the family-series, there- fore in the class-series* and thus is outside the Rules of the Code. The Code is not fully clear regarding the conditions of availability of family-series nomina. Only two conditions are clear for all nomina, concerning the stem of the nomen and the reference to a suprageneric taxon. Article 11.7.1.1 states that, to be available in its original publication, a fam- ily-series nomen must “be a noun in the nominative plu- ral formed from the stem of an available generic name’, which is then its nucleogenus* (type-genus). Therefore, any higher taxon nomen not based on an available gener- ic nomen is unavailable for a family-series nomen, but may under certain conditions be available for a class-se- ries nomen. Article 11.7.1.2 adds that the new nomen must “be clearly used as a scientific name to denote a supra- generic taxon and not merely as a plural noun or adjec- tive referring to the members of a genus”. Therefore, the explicit use of the rank family, or of another traditional rank of the family-series (subfamily, superfamily, tribe, subtribe, etc.), is not required for availability of nomina in the family-series. Two additional clear conditions, ap- plying only to nomina published after 1999, are given in Articles 16.1 (the nomen “must be explicitly indicated as intentionally new”) and 16.2 (the nomen “must be accom- panied by citation of the name of the type genus’’). Ac- cording to Article 11.7.1.1, before 2000, the type-genus may be indicated “either by express reference to the gener- ic name or by reference to its stem”, 1.e., by implicit ety- mological designation (Dubois 1984). Three conditions are unclear in the current Code regard- ing the status of new family-series nomina: (C1) the date; (C2) the requirement for validity of the nomen of the nu- cleogenus; and (C3) the distinction between family-series and class-series nomina. Bonn zoological Bulletin 57 (2): 149-171 (C1) No starting date is given in the Code for the use of family-series nomina in zoological nomenclature. How- ever, the rank family and related ones (superfamily, sub- family, tribe, subtribe, etc.) were not recognized by Lin- naeus (1758, 1761, 1764, 1766, 1767), although this au- thor made use of no less than seven ranks above the rank genus (Dubois 2007). Some authors of the 18 century used the ranks family and tribe, but not always for taxa above the rank genus and below the rank order, with fam- ily as a rank above tribe (Dubois 2006a). For example, some authors (e.g., De Geer 1778; Goeze & Donndorff 1797) used family as a rank below the rank genus, where- as others, including some quite recently, used tribe as a rank above the rank order (e.g., Scopoli 1777; de Blainville 1816; Huene 1952) or below the rank order but above the rank family (e.g., Oken 1821, 1833; Fitzinger 1826, 1843; Swainson 1835; Hogg 1841; Bonaparte 1845; de Blainville 1847; Stannius 1856). In zoological taxon- omy, the first authors that are traditionally credited with the creation of family-series nomina for taxa above the rank genus are authors who published their works in the early 19th century: e.g., Lamarck (1801), Latreille (1802, 1824, 1825), Oppel (181la—b), Rafinesque-Schmaltz (1814a—d), Rafinesque (1815), Vieillot (1816), Fischer (1817), Goldfuss (1820), Gray (1825) or Vigors (1825). However, a few authors in the second half of the 18" cen- tury already used the rank family for taxa at ranks between genus and order. This is the case of Batsch (1788, 1789), in a rather poorly known work discussed in detail below. Inasmuch as the familial nomina created by these authors were clearly based on the stems of available generic nom- ina considered valid by these authors, there is no reason for not crediting these authors with the creation of these familial nomina, even if this was ignored by most subse- quent authors until now (Dubois 2010: 25). (C2) Regarding the requirement for validity of the gener- ic nomen used as stem (nucleogenus), Article 11.7.1.1 states that “the generic name must be a name then used as valid in the new family-group taxon [Arts. 63, 64] (use of the stem alone in forming the name is accepted as ev- idence that the author used the generic name as valid in the new family-group taxon unless there is evidence to the contrary)”. There are several questions with this unclear formulation. First, what does “then” mean in this context? This word would have a clear sense only if it meant “in the work where the new family-series nomen is created”, but then why not write it in full words? If it meant “‘at the period of this work”, this would be difficult to apply, first because it is unclear how long the period to be considered should be (preferably it seems that it should not include more than ten or 20 years around the creation of the new family-series nomen), and second because at any given pe- ©ZFMK Family- and class-series nomina in zoology 151] riod of taxonomy the same nomen may be accepted as valid by part of the authors then active, and invalid by oth- ers, as will be illustrated below with the example of the nomina Lacerta Linnaeus, 1758 and LAcer7T4E Batsch, 1788. Furthermore, the words “used as valid in the new family-group taxon” show that this condition cannot ap- ply to works published before the creation of the latter tax- on! Therefore, this part of Article 11.7.1.1 would be made clearer by choosing between the two following formula- tions: (Fl) “the generic nomen must be used as valid in the new family-group taxon in the work where its nomen is created’; (F2) “the generic nomen must be used as valid by all active taxonomists in the 10 years before and after creation of the new family-group nomen” (or another span). Until this choice is made by the ICZN, this Article is not fully operational, as will be exemplified below. The French version of Article 11.7.1.1 in the current Code is not strictly equivalent to its English version, which is prob- lematic as these two texts are deemed to be “equivalent in force and meaning” (Anonymous 1999: xiii). As a mat- ter of fact, the French version of this Article ignores the term “then” (“alors”). In the previous edition of the Code (Anonymous 1985: 25), Article 11(f)(i)(1) wrote “then used as valid for a genus contained in that family-group taxon’. These elements suggest that formulation (F1) above corresponds to the real meaning of this article, and we follow this interpretation below. (C3) Regarding the distinction between family-series and class-series nomina, it is unambiguous in the Code only in the case of suprageneric nomina that are not based on available generic nomina, which are unavailable in the family-series, but may be available in the class-series, at least in some cases (see below). But what is the status of nomina based on the stem of available generic nomina cre- ated for taxa at ranks clearly above the family-series (or- der, class, etc.), or for taxa of unusual ranks, not clearly referable to the family- or class-series (such as phalanx, cohort, gens, etc.), or for taxa of unspecified ranks? The Code does not provide any clue for decision in such cas- es, all the more that, as reminded above, the explicit use of the rank family, or of another rank of the family-series, is not required for availability of nomina in the family- series. A few clear situations exist: (1) when a nomen is created for a taxon that is explicitly originally referred to a rank higher than superfamily, or than order, class or an- other rank traditionally referred for the class-series in zo- ology, whatever this rank is, such a nomen belongs in the class-series; (2) in contrast, when a nomen is created for a suprageneric taxon of rank lower than superfamily or than any other traditional rank in the family-series (fam- ily, subfamily, tribe, etc.), and is based on the stem of a nucleogenus, it belongs in the family-series. But when- ever a nomen is proposed for a taxon of any rank above the rank genus, and without clear hierarchical relationships Bonn zoological Bulletin 57 (2): 149-171 with other taxa of ranks unambiguously referable either to the family- or to the class-series, it may be treated e1- ther as a class-series nomen (this is the case for example of all suprageneric nomina created by Linnaeus: see Dubois 2007) or as a family-series nomen. In such cases, the etymology of the nomen may be a help for the deci- sion: 1f the nomen is based on the stem of an available generic nomen, it may be treated as a family-series nomen, otherwise as a class-series nomen. Another matter ignored by the Code is what could be called the consistency problem. In some publications of the 18th, 19th and even 20 centuries, some authors were not consistent regarding the mode of formation of their new familial nomina: some were based on the stem of available generic nomina, whereas others were not, being descriptive or geographical terms, terms based on the names of persons, etc. In such cases, the nomina of the first category could be accepted as available both as fam- ily-series and class-series nomina, but those of the second category can be considered available only in the class-se- ries. However, a choice has to be made between these two nominal-series for a// the nomina created together with the same rank, as it is not logical and conceivable to admit that the same author, in the same publication, created both family-series and class-series nomina for taxa of same rank (for details, see Dubois 2008b). Dubois (2006a: 178) pro- posed that, in such cases, for reasons of consistency in the taxonomic hierarchy, all these nomina be referred to the family-series, but that those which are incorrectly formed (not being based on available generic nomina, or formed through addition of a complex suffix unacceptable as a family-series suffix according to the Code), be considered nomenclaturally unavailable. These are of two kinds (Dubois 2006a: 178): arhizonyms are family-series nom- ina not based on generic nomina, and caconyms are fam- ily-series nomina based on generic nomina but with a com- plex suffix (such as -forma, -morpha, etc.). Examples of arhizonyms include “BATRACINIA”, “GYMNODERMIA” and ‘“PHRYNACINIA”, coined by Rafinesque (1815) for taxa of ranks family or subfamily, along with available family- series nomina like HYLARINIA, RANARINIA and TRITONIA. Ex- amples of caconyms include “RANIFORMES”, “HYLAE- FORMES”, “BUFONIFORMES” and “PIPAEFORMES’’, coined by Dumeéril & Bibron (1841) for taxa of rank family, along with available family-series nomina like CECILIOIDES, SALAMANDRIDES, AMPHIUMIDES and PROTEIDES. In his study of class-series nomenclature in zoology, Dubois (2006a: 228), after a detailed discussion of the problems mentioned above and others, proposed two new Rules to clarify this situation and to distinguish between family-series and class-series nomina in a simple, objec- tive and automatic manner: ©ZFMK 152 Alain Dubois & Roger Bour “(R4) Allocation of nomina to the family-series or to the class-series. Whenever a single new suprageneric nomen of a given taxonomic rank was established in a publica- tion, this nomen must be referred to the family-series if both following conditions are fulfilled: (A) it was proposed for a taxon of a rank usual within the family-series or of an unusual rank but clearly presented as being hierarchi- cally subordinate to a usual rank of that series although above the genus; and (B) it was coined by addition of a simple suffix denoting the plural to the stem of an avail- able genus-series nomen. In all other cases, the nomen must be referred to the class-series. Whenever several new suprageneric nomina of the same rank were established in a publication, they must all be referred to the same nom- inal-series, if they were treated heterogeneously with re- gard to the criteria above, they must follow the Rule of Taxonomic Consistency (R5). (R5) Rule of Taxonomic Consistency. All suprageneric nomina created in the same publication for taxa that were afforded the same taxonomic rank must be referred to the same nominal-series. In case of conflict between their al- location to nominal-series according to Rule (R4), the fam- ily-series takes precedence over the class-series, and nom- ina that, being incorrectly formed (arhizonyms or ca- conyms), cannot be considered as belonging to that se- ries, must be treated as nomenclaturally unavailable (anoplonyms*).” These proposed Rules should be studied carefully by the ICZN and incorporated into the Code, or others Rules should be proposed, but until this is done, ambiguity will exist and decisions regarding the status of some nomina of higher taxa will remain unclear, and will have to rely on arbitrary decisions on the part of some zoologists, as will now be shown. In what follows, these general questions will be concrete- ly studied in one zoological group: we will examine the status of the nomina used by the authors until now for (1) the order of reptiles including the turtles and (2) the fam- ily of turtles including the genus 7estudo Linnaeus, 1758. THE HIGHER NOMENCLATURE OF TURTLES Despite various works dealing with it, the higher nomen- clature of turtles is not yet stabilized. The nomenclatural chaos is clearly emphasized by the use of different and incompatible nomenclatures over very short periods of times, not only by different authors, but sometimes by the same one (e.g., Vetter 2002, 2004; Vetter & van Dijk 2006). The last publications in this respect, by Rhodin et al. (2008, 2009), are not reliable references, as they dis- play ignorance of several basic nomenclatural Rules of the Bonn zoological Bulletin 57 (2): 149-171 Code. For example, they do not follow the Code’s Prin- ciple of Coordination for superfamilies, which are cred- ited to authors and dates different from those of their hy- ponymous* families (e.g., KINOSTERNIDAE Agassiz, 1857 and KINOSTERNOIDEA Joyce, Parham & Gauthier, 2004) and sometimes given incorrect endings (7RIONyCHIA Hummel, 1929). An important nomenclatural flaw, discussed in de- tail below, is to refer the same nomen (7ESTUDINES Batsch, 1788) both to the family- and the class-series. Table | (in Appendix 1) provides a survey of various nom- ina, with their authors and dates when they were speci- fied, that have been used until now by a number of zool- ogists for the order of turtles and the family including the genus Zestudo Linnaeus, 1758. Several problems can be identified in this table. First, al- though the family including the genus 7estudo has almost always been known as TESTUDINIDAE, the author and date of the latter nomen has not been consensual. Some authors (e.g., Hunt 1958: 150; Iverson 1992: 3; Xianrui 1994: 4) have credited a nomen “Testudines” to Linnaeus (1758: 194, 198). However, it is fully clear that, in this and oth- er works of Linnaeus, the term Jestudines was a plural noun referring to the members of the genus Jestudo, not a family-series or class-series nomen (Article 11.7.1.2; Bour & Dubois 1985). This is stressed by the fact that Lin- naeus (1758: 198-199) also mentioned this word as 7és- tudine and Testudinibus. Others have credited the famil- ial nomen JESTUDINIDAE to Gray (1825), until Bour & Dubois (1985) drew the attention to the fact that the nomen TESTUDINES, coined by Batsch (1788: 437) for a family in- cluding the single genus Zestudo Linnaeus, 1758, was doubtless available in the family-series, where it has pri- ority over all subsequent nomina coined on the basis of the stem of this generic nomen (including 7EsTuUDIA Rafinesque-Schmaltz, 1814c, a nomen ignored by most authors until now). Following the Code, this nomen must simply be emended to TESTUDINIDAE Batsch, 1788 if used for a taxon of rank family, to TESTUDINOIDEA Batsch, 1788 for a taxon of rank superfamily, 7ESTUDININAE Batsch, 1788 for a subfamily, TESTUDININI Batsch, 1788 for a tribe and TESTUDININA Batsch, 1788 for a subtribe. Still more confusion has been exhibited by the authors re- garding the nomen of the order of turtles. The nomen TEs- TUDINES was used for this purpose, credited either to Lin- naeus (1758) or to Batsch (1788), which is incorrect in both cases for the reasons given above (the former being a generic nomen in the plural, the latter a family-series nomen). The first valid creation of a nomen TESTUDINES for an order was by Wagler (1830: 130, 133), but this is subsequent to the other nomina discussed below. As a mat- ter of fact, three other nomina were also widely used for the order, CHELONI, CHELONIA and TESTUDINATA. OZFMK Family- and class-series nomina in zoology 153 Both CHELON and CHELONIA are just subsequent la- tinizations of CHELONIENS Brongniart, 1800a. The spelling CHELONIA was first used by Ross & Macartney (and not Macartney alone, as wrongly stated by Loveridge 1957 or Romer 1966) in their 1802 translation of the work of Cuvier (1800). This latinization was posterior to that in CHELONI by Latreille (1800), used by many subsequent authors in the 19 century (Bour & Dubois 1985: 79) and resurrected by Bour (1981). Although the Code provides no guidelines for the authorship and date of class-series nomina, for reasons discussed in detail by Dubois (2006a, 2009), by simple consistency and parallelism with the Rules of the Code concerning family-series and genus-se- ries nomina, it is justified to credit a class-series nomen published first in a non-latinized form to the author of this original nomen, so in this case to Brongniart (1800a). The spelling CHELONII being anterior to CHELONIA, and the latter being open to confusion because of hemihomonymy (Starobogatov 1991) with the generic nomen Chelonia Brongniart, 1800b, the use of CHELONII was supported by Bour (1981) and Bour & Dubois (1985), who noted that this nomen had priority over TESTUDINATA, an ordinal nomen coined by Oppel (1811b). In conclusion, Bour & Dubois (1985) proposed to use the nomen CHELONII Brongniart, 1800a for the order of turtles, a suggestion adopted by various subsequent authors (see Table 1). As the Code provides no Rules or even guidelines for class-series nomenclature, this suggestion was based on the use of the Principle of Onomatophores* (so-called “Principle of Typification”) in a way similar to its use in the three lower nominal-series recognized by the Code, a method explicitly presented by Dubois (1984). However, as was later shown by Dubois (2004, 2005a—b, 2006a—b, 2009; Dubois & Ohler 2009), because no Principle of Co- ordination is in force in class-series nomenclature, such a practice does not allow unambiguous allocation of a class-series nomen to a taxon as soon as several hierar- chically subordinated taxa have the same onomatophore, so that more complete Rules had to be devised (Dubois 2006a). For the precise allocation of nomina to higher taxa, this system uses both the originally included genera or conucleogenera of the newly established taxon, and the genera originally expressly excluded from it, its alieno- genera. Until these proposed Rules, or others, are incor- porated into the Code in order to regulate class-series nomenclature, the latter will remain chaotic and left to “freedom” and “opinions” of individual zoologists, which will be a permanent nuisance for proper and unambigu- ous communication among all biologists. This problem is made worse by the ambiguity, discussed above, regarding the distinction between class-series and family-series nomina in the Code. Although Batsch (1788) had clearly referred his new taxon TESTUDINES to the rank Bonn zoological Bulletin 57 (2): 149-171 family, there is nothing in the Code that imposes to refer this nomen to the family-series, all the more that Batsch (1788, 1789) was not consistent in his use of etymology for his familial nomina, some only being based on the stems of generic nomina he considered valid (see below). Because of this ambiguity of the Code, it would be pos- sible to refer the nomen 7ESTUDINES Batsch, 1788 either to the family-series (which clearly has our preference) or to the class-series. But it is fully unacceptable and impos- sible to refer it to both! This would be similar to accept- ing that a genus-series nomen, such as Ranoidea Tschu- di, 1838 for example, can be considered available both as the nomen of a genus and of a superfamily! This is how- ever what has been done by Fritz & Havas (2006, 2007), followed by Vetter & van Dijk (2006) and Rhodin et al. (2008, 2009), who recognized, in the same classification, an order TESTUDINES Batsch, 1788 and a family 7ésTU- DINIDAE Batsch, 1788, although both nomina are based on the one and single appearance of the nomen TESTUDINES in page 437 of Batsch (1788)! The fact that such incred- ible nomenclatural treatments can be accepted as valid by several contemporaneous taxonomists and periodicals points to the poor interest granted by many colleagues nowadays to nomenclatural Rules and to the chaotic sit- uation created in zoological nomenclature by the incom- pleteness and ambiguity of the Code. This exemplary case prompted us to undertake a detailed and complete survey of all suprageneric nomina created by Batsch (1788, 1789), which fully exemplifies these problems and allows to propose solutions to them. BATSCH’S (1788, 1789) SUPRAGENERIC NOMINA IN ZOOLOGY Batsch (1788, 1789) was one of the authors who, in the late 18' century, proposed a comprehensive classification of the animal kingdom and tried to improve the scheme of Linnaeus (1758, 1766, 1767) in this respect. In this clas- sification, he used four ranks above the rank genus: fam- ily, order, class and an upper unnamed rank that we treat here for more simplicity as “superclass”. This classifica- tion is summarized here in our Table 2 (in Appendix 1). Batsch (1788) was the first author to divide the animal kingdom in two main groups, his “superclasses” OSSEA and CRUSTACEA, which exactly correspond to the distinc- tion between “animaux a vertébres” and “animaux sans vertébres” first proposed by Lamarck in his lectures (which were not published until 1801), and which Cuvier (1800) was the first author to formally name in a publication as VERTEBRES (VERTEBRATA) and IN- VERTEBRES (INVERTEBRATA). Although Batsch’s (1788) ©ZFMK 154 Alain Dubois & Roger Bour OssEaA has priority over VERTEBRATA, it would be inap- propriate to replace the latter, which has been used mil- lions of times in the scientific literature and therefore qual- ifies as a sozonym*, by the former, which has been ignored and which is therefore a distagmonym* (Dubois 200S5a: 86, 2005b: 412). In his OsskA, Batsch (1788) recognized four classes, MAMMALIA, AVES, AMPHIBIA and PISCES, whereas in his CRUSTACEA he recognized two classes, INSECTA and VER- MES. Although the nomina of these six classes are iden- tical to those of the six zoological classes of Linnaeus (1758, 1766, 1767), their content is not always exactly the same. For example, Batsch (1789) removed from his VER- MES the genus Myxine Linnaeus, 1758 placed in this clas- sis by Linnaeus, and which is in fact a chordate. There- fore, the nomina used by Batsch for these classes should be credited to him, not to Linnaeus. They are junior homonyms of Linnaeus’ identical nomina (see Dubois 2006a). All genera in Batsch (1788, 1789) are referred to fami- lies. Families are referred to orders and then to the class only in one class (MAMMALIA). The nomina of the orders of mammals also are in part borrowed from Linnaeus, but here also sometimes with a slightly different content, which requires to consider them as distinct, junior homonymous nomina. In the other five classes, the only rank used above genus is that of family. Because the rank family is expressly used by Batsch, is situated in the nomenclatural hierarchy above the rank genus and below the ranks class and order (when available), and because some at least of these nomina are coined by addition of an ending indicating plural to the stem of an available generic nomen considered valid by Batsch (1788, 1789), we hereby consider the nomina of Batsch’s “families” to be indeed family-series nomina, some of which only are nomenclaturally available. The available family-series nomina in Batsch (1788, 1789), that appear in Table 2, are the 17 familial nomina in his work based on available generic nomina listed by him as valid among the genera of the family. This is for example the case of TESTUDINES Batsch, 1788, a taxon ex- pressly mentioned as including the genus Jestudo Lin- naeus, 1758. As shown in Table 2, there are two categories of unavail- able family-series nomina in Batsch (1788, 1789). The first one consists of arhizonyms, 1.e., family-series that were not based on any then available zoological generic nomen. The second one consists of nomina that were indeed based on then available zoological generic nomina, but these nomina not being listed by Batsch (1788, 1789) as valid Bonn zoological Bulletin 57 (2): 149-171 among the members of the family, being presumably con- sidered invalid synonyms of nomina used by Batsch as valid. As we here adopted the formulation (F1) above of Article 11.7.1.1 of the Code, these nomina must be con- sidered as nomenclaturally unavailable, but if interpreta- tion (F2) had to be followed these nomina would have to be treated as available. This small doubt is one of the con- sequences of the ambiguous writing of Article 11.7.1.1 in the current version of the Code. The Code is silent about the nomenclatural status of fa- milial nomina such as those created by Batsch (1788, 1789) but shown above to be unavailable in the family- series. In contrast, under the Rules proposed by Dubois (2006a) for class-series nomenclature, these nomina be- long unambiguously in the family-series and are therefore clearly unavailable in the class-series as well, because of the Rule of Taxonomic Consistency presented above. Except three, all the generic nomina listed by Batsch (1788, 1789) in his classification of the animal kingdom had previously been made available in zoological nomen- clature by Linnaeus (1758) and in subsequent publications between 1758 and 1790. The only three exceptions are the nomina Cylindrus Batsch, 1789, Hydrocantharus Batsch, 1789 and Turris Batsch, 1789. The status of these three nomina is discussed below in Appendix 2. Table 3 (in Appendix 1) lists the 17 familial nomina made nomenclaturally available in zoological nomenclature by Batsch (1788, 1789). Until now, only one (TESTUDINIDAE) has been credited to Batsch (1788), and the other 16 are traditionally credited to other authors at subsequent dates, but should now be credited to Batsch. This poses no prob- lem of “nomenclatural stability”, as none of these 16 fa- milial nomina has to change, the change concerning on- ly their author and date. The familial nomen LAceRTIDAE, that had previously (Dubois 2004; Dubois & Bour 2010) been credited to Batsch (1788), does not appear in Table 3. This is because this nomen could be considered available only under in- terpretation (F2) of Article 11.7.1.1. The genus Lacerta Linnaeus, 1758 was recognized by most authors of the end of the 18 century, but not by Laurenti (1768) who had split it into several genera and had not retained the nomen Lacerta for any of them (in contrast for what he had done in other cases, e.g. for Rana). He was apparently followed in this by Batsch (1788), who did not recognize or even mention the genus Lacerta. As we here adopted interpre- tation (F1) of Article 11.7.1.1, the family nomen L4cER- TIDAE cannot be credited to Batsch (1788). It must there- fore be credited to the first subsequent author who used a family nomen based on the generic nomen Lacerta for ©ZFMK Family- and class-series nomina in zoology 155 a family where the latter generic nomen was considered valid. This happens to be Oppel (1811b: 16). Establishing the proper nomen for the order of turtles (or “turtles, tortoises and terrapins’’), i.e., including all recent turtles as well as a few additional Triassic groups, is be- yond the scope of the present paper, and we just provide here a few comments in this respect. As discussed above, the nomen 7ESTUDINES Batsch, 1788, being available in the family-series, is not available in the class-series and can- not be used for an order. Under the nomenclatural Rules proposed by Dubois (2006a), the nomina CHELONII Brongniart, 1800a and TESTUDINATA Oppel, 1811b are available in the class-series. However, they do not apply to the order of turtles, but to still higher taxa. Under these Rules, the nomen CHELONI Brongniart, 1800a applies to the most inclusive class-series taxon con- taining the genera Che/onia Brongniart, 1800b and Tes- tudo Linnaeus, 1758, and excluding the 19 nominal gen- era referred by Brongniart (1800b) to his orders BATRA- CHIA, OPHIDIA and SAURIA. As for the nomen TESTUDINATA Oppel, 1811b, it applies to the most inclusive class-series taxon containing the gen- era Chelonia Brongniart, 1800b, Chelys Oppel, 1811b, Emys Dumeril, 1806, TJestudo Linnaeus, 1758 and Trionyx Geoffroy Saint-Hilaire, 1809, and excluding the 48 nom- inal genera referred by Oppel (1811b) to his orders SQua- MATA and NUDA. Oppel (1811b) credited the nomina of his orders TESTU- DINATA and NupDaA to Klein (1751), a work which, being anterior to 1758, is not nomenclaturally available. How- ever, Joyce et al. (2004: 998) recently drew the attention to Behn’s (1760) translation and adaptation of Klein’s (1751) book, which includes all the taxa and nomina of the latter work. These post-1758 nomina would be avail- able, with the authorship “Klein in Behn, 1760”, if this book was nomenclaturally available, but, for reasons ex- plained in detail in our Appendix 2 below, we consider that it should not be considered so. Several other class-series nomina applying to turtles and related groups have been published after the works just mentioned. Establishing the class-series taxa to which these nomina apply under Dubois’s (2006a) proposed Rules requires a long and detailed survey that would take us far beyond the purpose of the present paper and will be presented elsewhere. For the time being, this work is not urgent, as the phylogenetic relationships among these groups, and with the other tetrapods, are currently high- ly controversial (e.g., Werneburg & Sanchez-Villagra 2009, and included references), and it will be possible to Bonn zoological Bulletin 57 (2): 149-171 settle a robust nomenclature of these groups only when some consensus emerges on these questions. CONCLUSION The analysis presented above and the examples studied show that the current Rules of the Code are ambiguous regarding the allocation of nomina of higher zoological taxa to either the family-series or the class-series of nom- ina, and regarding the conditions of availability of fami- ly-series nomina. These Rules should be improved through modifications of Article 11.7.1.1 as suggested above, and mostly through incorporation of Rules for class-series nomina, as proposed in detail by Dubois (2006a). A detailed study of all suprageneric nomina in the work of Batsch (1788, 1789) shows that this author proposed many family-series nomina, which belong in three cate- gories: (C1) nomina clearly based on the stems of avail- able generic nomina that were considered valid in this work: such nomina are available in the family-series; (C2) nomina apparently based on the stems of generic nomina nomenclaturally available at that time, but not treated as valid in this work: such nomina are unavailable both in the family-series and in the class-series; (C3) arhizonyms, i.e., nomina not based on the stems of any generic nomen nomenclaturally available at that time: such nomina are also unavailable both in the family-series and in the class- series. Nomina of the categories (C2) and (C3) are defi- nitely unavailable and will never have to be used as valid in zoological nomenclature. But the nomina of category (C1) compete for priority with all other family-series sub- sequently proposed in zoological nomenclature. It so hap- pens that these 17 nomina are identical with family-series nomina coined later on and based on the same nucleogen- era. Therefore they must replace them, which entails no change in the nomina themselves (and therefore no dis- ruption of nomenclatural stability) but only modifications regarding their authors and dates. These changes, listed in Table 3, should be implemented without delay in the respective zoological groups where they belong. This analysis contributes to a clarification of the higher nomenclature of turtles. The nomen 7ESTUDINES Batsch, 1788 is not a class-series, but a family-series nomen. It cannot be used for the order of turtles, but is the valid nomen, under the spelling 7esTUDINIDAE, of the family in- cluding the genus 7estudo Linnaeus, 1758 and of all oth- er coordinate taxa as recognized in any given classifica- tion. As for the order of turtles, establishing the valid nomen of this taxon and of its superordinate taxa under the Rules proposed by Dubois (2006a) is beyond the scope ©OZFMK 156 Alain Dubois & Roger Bour of the present paper, but it is shown here that neither TES- TUDINES Linnaeus, 1758, nor TESTUDINATA Klein in Behn, 1760, nor TESTUDINES Batsch, 1788, nor CHELONII Brongniart, 1800a, nor TESTUDINATA Oppel, 1811b apply to this taxon. As long as the Code does not provide for- mal Rules for the nomenclature of class-series taxa, the higher nomenclature of turtles (as well as that of all oth- er zoological groups) will remain a matter of personal or collective tastes, opinions and arbitrary decisions of zootaxonomists. 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Online at http://www.animalbase.uni-goettingen.de /zooweb/servlet/AnimalBase/home/genustaxon?id=4111 last accessed 08-02-2010 Winckworth R (1945) The types of the Boltenian genera. Jour- nal of molluscan Studies 26: 136-148 Xianrui Y (1994) Fossil and recent turtles of China. Science Press, Beijing Yeh HK (1963) Fossil turtles of China. Paleontologica sinica 150 (N.S.C. 18): i-iv + 1-112, pl. 1-21 Zhao E, Adler K (1993) Herpetology of China. Contributions to Herpetology, Nr. 10. Society for the Study of Amphibians and Reptiles, Athens (OH) Zimmermann EAW von (1780) Geographische Geschichte des Menschen, und der vierftiBigen Thiere. Zweiter Band. Wey- gandschen Buchhandlung, Leipzig Received: 10. VII.2010 Accepted: 25.1X.2010 ©ZFMK 162 APPENDIX 1 Table 1. Alain Dubois & Roger Bour Chronological presentation of the family-series and class-series nomina used in various publications for the order of turtles and for the family including the genus Zestudo Linnaeus, 1758. The authors and dates are mentioned below only when they were so in the works cited. Nomina connected by the sign <> are allelonyms (Dubois 2006a), i.e., alternative nomina proposed or used by an author in the same publication for the same taxon (same content and taxonomic rank), without choosing between them. Reference Nomen used for the order (or for a class-series taxon of another rank) of turtles Nomen used for the family including the genus Zestudo Linnaeus, 1758 Batsch 1788: 437 Brongniart 1800a: 81 Latreille 1800: xi Ross & Macartney in Cuvier 1802: tab. 3 Dumeéril 1806: 76 Oppel 1811b: 4, 6 Rafinesque-Schmaltz 1814c: 66 Rafinesque 1815: 74 Merrem 1820: 6, 7, 12, 17 Latreille 1825: 91 Gray 1825: 194, 210 Fitzinger 1826: 5 Ritgen 1828: 269, 270 Wagler 1828: 861 Wagler 1830: 130 Bonaparte 1831: 63, 68 Griffith & Pidgeon 1831: 4, 6 Gray 183la: 2 Gray 1831b: 3, 7 Duméril & Bibron 1834: 346, 352 Dumeéril & Bibron 1835: 1 Fitzinger 1835: 107 Fitzinger 1843: 29 Ruippel 1843: 297 Wiegmann & Ruthe 1843: 166, 168 Gray 1844: 3 Bonaparte 1845: 3 Bonaparte 1850: pl. Gray 1855: title page, 1, 4 Agassiz 1857: 235, 249 Gunther 1859: 379 Peters 1862: 271 Strauch 1862: 19, 20 Gunther 1864: x, 1, 3 Strauch 1865: 205 Troschel 1866: 182 Fitzinger 1867: 85 Gray 1873: iv, 1 Cope 1875: 50, 54 Peters 1882: 2 Baur 1887: 96, 101 Boulenger 1889: 4, 48 Strauch 1890: 9, 10 Baur 1892: 419, 420 Boettger 1893: 2, 3 Barboza du Bocage 1895: | Steyneger 1907: 483, 488 Siebenrock 1909: 429 De Rooij 1915: 285, 288 Stejneger & Barbour 1917: 113 Boulenger 1923: 42 CHELONIENS CHELONII CHELONIA CHELONII TESTUDINATA Klein, 1751 PEROSTIA PEROSTIA TESTUDINATA Oppel, 1811 CHELONII CHELONII Latreille, 1800 Monopnoa [including tribe TestupINaATA Klein, 1751] CHELONII <> STERRICHROTES CHELYNAE TESTUDINES CHELONII CHELONIA TESTUDINATA CHELONII CHELONIENS Brongniart, 1800 CHELONIENS Brongniart, 1800 MONOPNOA TYLOPODA Wagler, 1828 CHELONII CHELONII CHELONIA CHELONII CHELONII TESTUDINATA Oppel, 1811 <* CHELONIA Gray, 1835 [sic] TESTUDINATA Klein, 1751 CHELONII CHELONII CHELONIA CHELONIA CHELONII CHELONII TESTUDINATA TESTUDINATA CHELONI <> TESTUDINATA TESTUDINATA CHELONIA Brongniart, 1800 CHELONIA TESTUDINATA CHELONIA CHELONIA TESTUDINATA Oppel, 1811 TESTUDINATA Oppel, 1811 CHELONIA TESTUDINATA Oppel, 1811 CHELONIA TESTUDINES AMYDAE TESTUDIA TESTUDIA CRYPTOPODI TESTUDINIDAE TESTUDINOIDEA CHERSOCHELONES <*> DYSMYDAE TYLOPODAE HEDRAEOGLOSSAE TESTUDINIDAE TESTUDINIDAE TESTUDINIDAE Gray, 1825 “CHERSITES” “CHERSITES” TYLOPODA TESTUDINES CHERSINAE TESTUDINIDAE TESTUDINIDAE TESTUDINIDAE TESTUDINIDAE TESTUDININA Bonaparte, 1831 TESTUDINIDAE TESTUDINIDA TESTUDINIDAE TESTUDINIDA TESTUDINIDA CHERSINAE TESTUDINIDAE TESTUDINIDAE TESTUDININA TESTUDINIDAE TESTUDINIDAE TESTUDINIDA TESTUDINIDAE TESTUDINIDAE TESTUDINIDAE TESTUDINIDAE TESTUDINIDAE Gray 1825 TESTUDINIDAE TESTUDINIDAE TESTUDINIDAE Bonn zoological Bulletin 57 (2): 149-171 ©ZFMK Reference Family- and class-series nomina in zoology Nomen used for the order (or for a class-series taxon of another rank) of turtles 163 Nomen used for the family including the genus Zestudo Linnaeus, 1758 Mertens & Miller 1928: 20 Smith 1933: 49, 136 Freiberg 1938: 7, 9 Terente’v & Chernov 1949: 88, 95 Smith & Taylor 1950: 12, 27 Schmidt 1953: 86, 104 Bergounioux 1955: 187, 508 Mertens & Wermuth 1955: 333, 370 Romer 1956: 495, 504 Loveridge 1957: 163 Loveridge & Williams 1957: 175, 181 Hunt 1958: 150 Wermuth & Mertens 1961: 1, 171 Fuhn & Vancea 1961: 157, 158 Goin & Goin 1962: 73, 254 Yeh, 1963: 7, 27 Romer 1966: 365 Kuhn 1967: 114 Pritchard 1967: 27 Ckhikvazde 1970: 245 Auffenberg 1974: 140 Gaffney 1975: 423 Mtynarski 1976: 6 Webb et al. 1978: vii Nutaphand 1979: 13, 55 Bour 1981: 133 De Broin 1982: 897 Welch 1982: 206, 207 Pritchard & Trebbau 1984: 11, 197 Bour & Dubois 1985: 78 Alderton 1988: 108 Dundee 1989: 403 Ernst & Barbour 1989: 3, 227 King & Burke 1989: 16, 69 Jiufa & Ting, 1992: 1, 4 Iverson, 1992: 3, Zhao & Adler, 1993: 164, 171 David 1994: 16, 18 Xianrui 1994: 4, 62 Richard 1999: 85 Boycott & Bourquin 2000: 32 De Lapparent de Broin 2001: 166, 187 Noriega et al. 2000: 321 Kuzmin 2002: 17, 84 Vetter 2002: 3, 5 Mickoleit 2004: 282, 294 Pough et al. 2004: 97, 109 Vetter 2004: 3, 8 Vanni & Nistri 2006: 23 Fritz & Havas 2006: 10, 122 Vetter & van Dijk 2006: 3, 8 Fritz & Havas 2007: 163, 265 Pritchard 2007: 46 Abbazzi et al. 2008: 123 Rhodin et al. 2008: 2, 12 Rhodin et al. 2009: 42, 52 TESTUDINATA Oppel, 1811 TESTUDINES Batsch, 1788 TESTUDINATA Oppel, 1811 CHELONIA [in subclass TESTUDINES | TESTUDINES Batsch, 1788 CHELONIA CHELONIA [in subclass TESTUDINATA ] TESTUDINES CHELONIA <> TESTUDINATA TESTUDINATA Oppel, 1811 TESTUDINATA TESTUDINES Linnaeus, 1758 TESTUDINES TESTUDINES Batsch, 1788 TESTUDINATA CHELONIA CHELONIA Macartney, 1802 TESTUDINES Batsch, 1788 CHELONIA CHELONIA TESTUDINATA Shaw, 1802 TESTUDINES Batsch, 1788 TESTUDINES Batsch, 1788 CHELONIA CHELONIA <> TESTUDINES CHELONII CHELONII Brongniart, 1800 CHELONIA TESTUDINES CHELONII Brongniart, 1800 CHELONIA TESTUDINES TESTUDINES TESTUDINES Batsch, 1788 TESTUDINATA TESTUDINES Linnaeus, 1758 TESTUDINES CHELONI Brongniart, 1800 TESTUDINES Linnaeus, 1758 CHELONI Brongniart, 1800 CHELONIA <> TESTUDINES CHELONI Brongniart, 1800 CHELONII TESTUDINES TESTUDINES Linnaeus, 1758 CHELONIA «> TESTUDINES TESTUDINES <> CHELONIA TESTUDINES Linnaeus, 1758 CHELONI Brongniart, 1800 TESTUDINES Batsch, 1788 TESTUDINES Batsch, 1788 TESTUDINES Batsch, 1788 CHELONII Latreille, 1800 <* CHELONIA Macartney, 1802 <> TESTUDINES [neither Linnaeus, 1758, nor Batsch, 1788] CHELONH Brongniart, 1800 TESTUDINES Batsch, 1788 TESTUDINES Batsch, 1788 TESTUDINIDAE TESTUDINIDAE Gray, 1825 TESTUDINIDAE Gray, 1825 TESTUDINIDAE TESTUDINIDAE Gray, 1825 TESTUDINIDAE TESTUDINIDAE TESTUDINIDAE TESTUDINIDAE TESTUDINIDAE TESTUDINIDAE Gray, 1825 TESTUDINIDAE Gray, 1825 TESTUDINIDAE TESTUDINIDAE Gray, 1825 TESTUDINIDAE TESTUDINIDAE TESTUDINIDAE TESTUDINIDAE Gray, 1825 TESTUDINIDAE TESTUDINIDAE TESTUDINIDAE Gray, 1825 TESTUDINIDAE Gray, 1825 TESTUDINIDAE Gray, 1825 TESTUDINIDAE TESTUDINIDAE TESTUDINIDAE Batsch, 1788 TESTUDINIDAE TESTUDINIDAE TESTUDINIDAE Gray, 1825 TESTUDINIDAE Batsch, 1788 TESTUDINIDAE TESTUDINIDAE TESTUDINIDAE Gray, 1825 TESTUDINIDAE TESTUDINIDAE Batsch, 1788 TESTUDINIDAE TESTUDINIDAE Batsch, 1788 TESTUDINIDAE Gray, 1825 TESTUDINIDAE TESTUDINIDAE TESTUDINIDAE Batsch, 1788 TESTUDINIDAE TESTUDINIDAE Batsch, 1788 TESTUDINIDAE Batsch, 1788 TESTUDINIDAE TESTUDINIDAE TESTUDINIDAE Rafinesque, 1815 TESTUDINIDAE Batsch, 1788 TESTUDINIDAE Batsch, 1788 TESTUDINIDAE Batsch, 1788 TESTUDINIDAE Batsch, 1788 TESTUDINIDAE Batsch, 1788 TESTUDINIDAE Batsch, 1788 TESTUDINIDAE Batsch, 1788 Bonn zoological Bulletin 57 (2): 149-171 ©ZFMK 164 Alain Dubois & Roger Bour Table 2. The supraspecific taxa of animals listed in Batsch (1788, 1789). The animals are distributed in two class-series taxa, OsseA and Crustacea, for which no ranks are given in this book; they are here referred to the rank “superclassis”. All other ranks are mentioned expressly in Batsch (1788, 1789). Nomina connected by the sign <> are allelonyms (Dubois 2006a), i.e., alternative nomina proposed by an author in the same publication for the same taxon (same content, onomatophore and taxonomic rank), with- out choosing between them. The generic nomina are given here under their original spelling (protonym; Dubois 2000), with men- tion between parenthesis of the subsequent spelling (aponym; Dubois 2000) used by Batsch, whenever relevant. All these generic nomina had been created by Linnaeus (1758) or in subsequent works published before those of Batsch, except three, followed here by the sign +, which were made nomenclaturally available by Batsch (1789), and the status of which is discussed below in Appen- dix 1. This appendix also discusses the status of three post-Linnean generic nomina, followed by the sign ‘¢, which we consider here nomenclaturally unavailable. The familial nomina created by Batsch (1788, 1789) are of three kinds: (1) a familial nomen un- derlined in this Table was clearly based on the nomen (also underlined) of a genus expressly referred by Batsch to the familia as a valid nomen, which is therefore its nucleogenus (type-genus) by implicit etymological designation (Dubois 1984), thus making this family-series nomen available under Art. 11.7.1.1; (2) a familial nomen followed by an asterisk * can be considered derived from the nomen of a genus traditionally referred to the same taxonomic group, but not used as valid by Batsch, being probably considered a synonym of another nomen; this generic nomen is listed between square brackets, also followed by *, after the list of the valid genera of the family according to Batsch; such a family-series nomen, being based on a generic nomen considered in- valid by Batsch, is unavailable under Art. 11.7.1.1, thus shown “between quotation marks”; (3) a familial nomen followed by the sign ° is an arhizonym (Dubois 2006a: 178), 1.e., cannot be construed as being based on a then available generic nomen and is therefore unavailable under Art. 11.7.1.1, thus also shown “between quotation marks”. “Superclassis” OssEA Batsch, 1788: 81. Classis MAMMALIA Batsch, 1788: 87. Ordo Bruta Batsch, 1788: 103. Familia “Cozossr’° Batsch, 1788: 107. Genera (2): Elephas Linnaeus, 1758: 18; Rhinoceros Linnaeus, 1758: 19. Familia “CaT4PHrAcTA’* Batsch, 1788: 107. Genera (2): Dasypus Linnaeus, 1758: 18; Manis Linnaeus, 1758: 18. [Cataphractus* Brisson, 1762: 12-13]. Familia BraDyPopa Batsch, 1788: 108. Genera (2): Bradypus Linnaeus, 1758: 18; Myrmecophaga Linnaeus, 1758: 18. Ordo Pecora Batsch, 1788: 103. Familia “Ovina’’* Batsch, 1788: 105. Genera (2): Camelus Linnaeus, 1758: 19; Capra Linnaeus, 1758: 19. [Ovis* Linnaeus, 1758: 19]. Familia CERVINA Batsch, 1788: 105. Genera (4): Antilope Pallas, 1766b: 232; Bos Linnaeus, 1758: 19; Cervus Linnaeus, 1758: 19; Moschus Linnaeus, 1758: 19. Ordo GLIRES Batsch, 1788: 103. Familia Murina Batsch, 1788: 115. Genus (1): Mus Linnaeus, 1758: 19. Familia LEPORINA Batsch, 1788: 115. Genera (4): Cavia Pallas, 1766b: 30; Lepus Linnaeus, 1758: 19; Marmota Blumenbach, 1779: 79; Spalax Gueldenstaedt, 1770: 409. Familia SC/URINA Batsch, 1788: 115. Genera (3): Dipus Zimmermann, 1780: 354; Glis Brisson, 1762: 13, 113; Sciurus Linnaeus, 1758: 19. Familia C4s7orREA Batsch, 1788: 115. Genera (2): Castor Linnaeus, 1758: 19; Hystrix Linnaeus, 1758: 19. Ordo Primates Batsch, 1788: 103. Familia “PrimaTes’’° Batsch, 1788: 108. Genera (3): Homo Linnaeus, 1758: 18; Lemur Linnaeus, 1758: 18; Simia Linnaeus, 1758: 18. Ordo FERAE Batsch, 1788: 103. Familia FELINA Batsch, 1788: 110. Genus (1): Fe/is Linnaeus, 1758: 18. Familia Canina Batsch, 1788: 110. Genera (2): Canis Linnaeus, 1758: 18; Hyaena Brisson, 1762: 13, 168. Familia Ursivé Batsch, 1788: 110. Genus (1): Ursus Linnaeus, 1758: 18. Familia MUSTELINA Batsch, 1788: 110. Genera (3): Lutra Brisson, 1762: 13, 201; Mustela Linnaeus, 1758: 18; Viverra Linnaeus, 1758: 18. Ordo BELLUAE Batsch, 1788: 103. Familia “BELLUAE”® Batsch, 1788: 105. Genera (4): Equus Linnaeus, 1758: 19; Hippopotamus Linnaeus, 1758: 19; Hydrochoerus Brisson, 1762: 12, 80 (as Hydrochae- rus); Sus Linnaeus, 1758: 18. Ordo Rosores Batsch, 1788: 103. Familia Z4LPinA Batsch, 1788: 113. Genera (3): Erinaceus Linnaeus, 1758: 18; Sorex Linnaeus, 1758: 18; Za/pa Linnaeus, 1758: 18. Familia “PTEROPoDA’* Batsch, 1788: 105. Genus (1): Vespertilio Linnaeus, 1758: 18. [Pteropus* Brisson, 1762: 13, 153]. Bonn zoological Bulletin 57 (2): 149-171 ©ZFMK Family- and class-series nomina in zoology 165 Familia “MarsuPidALes”* Batsch, 1788: 105. Genus (1): Didelphis Linnaeus, 1758: 18 (as Didelphys). [“Marsupiale”*: Edward in Catesby, 1771]. Ordo PINNIPEDA Batsch, 1788: 103. Familia “P/NN/PEDA’’° Batsch, 1788: 116. Genera (3): Phoca Linnaeus, 1758: 18; Rosmarus Briinnichius, 1771: 34; Trichechus Linnaeus, 1758: 18 (as Trichecus). Ordo CETACEA Batsch, 1788: 103. Familia “Ce74cea”* Batsch, 1788: 116. Genera (4): Balaena Linnaeus, 1758: 19; Del/phinus Linnaeus, 1758: 19; Monodon Linnaeus, 1758: 19; Physeter Linnaeus, 1758: 19. [Cetus* Brisson, 1762: 225]. Classis AVES Batsch, 1788: 88. Familia “ANSERES”’* Batsch, 1788: 276. Genera (11): A/ca Linnaeus, 1758: 84; Anas Linnaeus, 1758: 84; Colymbus Linnaeus, 1758: 135; Diomedea Linnaeus, 1758: 84; Larus Linnaeus, 1758: 84; Mergus Linnaeus, 1758: 84; Pelecanus Linnaeus, 1758: 84; Phaeton Linnaeus, 1758: 84; Procel- laria Linnaeus, 1758: 84; Rynchops Linnaeus, 1758: 138 (as Rhynchops); Sterna Linnaeus, 1758: 84. [Anser* Brisson, 1760: 262]. Familia “GrALLAe”* Batsch, 1788: 276. Genera (11): Ardea Linnaeus, 1758: 84; Charadrius Linnaeus, 1758: 85; Fulica Linnaeus, 1758: 84; Haematopus Linnaeus, 1758: 85; Phoenicopterus Linnaeus, 1758: 84; Platalea Linnaeus, 1758: 84; Rallus Linnaeus, 1758: 84; Recurvirostra Linnaeus, 1758: 84; Scolopax Linnaeus, 1758: 84; Tantalus Linnaeus, 1758: 84; Tringa Linnaeus, 1758: 84. [“Gralla”+: Eberling in Sonnerat, 1777]. Familia STRUTHIONES Batsch, 1788: 276. Genera (3): Didus Linnaeus, 1766: 119; Otis Linnaeus, 1758: 85; Struthio Linnaeus, 1758: 85. Familia “7ENUIROSTRES’”° Batsch, 1788: 276. Genera (4): Certhia Linnaeus, 1758: 83; Merops Linnaeus, 1758: 83; Trochilus Linnaeus, 1758: 83; Upupa Linnaeus, 1758: 83. Familia “CUNEIROSTRES’’° Batsch, 1788: 276. Genera (2): Alcedo Linnaeus, 1758: 83; Picus Linnaeus, 1758: 83. Familia “G4LLINAE’* Batsch, 1788: 276. Genera (7): Columba Linnaeus, 1758: 85; Crax Linnaeus, 1758: 85; Meleagris Linnaeus, 1758: 85; Numida Linnaeus, 1764: 27; Pavo Linnaeus, 1758: 85; Phasianus Linnaeus, 1758: 85; Tetrao Linnaeus, 1758: 85. [Gallus* Brisson, 1760: 45]. Familia “AccipiTres’’* Batsch, 1788: 277. Genera (3): Falco Linnaeus, 1758: 83; Strix Linnaeus, 1758: 83 (as Stryx); Vultur Linnaeus, 1758: 83. [Accipiter* Brisson, 1760: 310]. Familia “LEVIROSTRES’° Batsch, 1788: 27. Genera (4): Buceros Linnaeus, 1758: 83; Crotophaga Linnaeus, 1758: 83; Psittacus Linnaeus, 1758: 83; Ramphastos Linnaeus, 1758: 83. Familia Cordces Batsch, 1788: 277 <> “P4SSERES”’° Batsch, 1788: 277. Genera (20): Alauda Linnaeus, 1758: 85; Ampelis Linnaeus, 1766: 119; Caprimulgus Linnaeus, 1758: 85; Coracias Linnaeus, 1758: 83; Corvus Linnaeus, 1758: 83; Cuculus Linnaeus, 1758: 83; Emberiza Linnaeus, 1758: 85; Fringilla Linnaeus, 1758: 85; Gracula Linnaeus, 1758: 83; Hirundo Linnaeus, 1758: 85; Jynx Linnaeus, 1758: 83 (as Jynx); Lanius Linnaeus, 1758: 83; Loxia Linnaeus, 1758: 85; Motacilla Linnaeus, 1758: 85; Oriolus Linnaeus, 1766: 117; Paradisaea Linnaeus, 1758: 110; Parus Linnaeus, 1758: 85; Sitta Linnaeus, 1758: 83; Sturnus Linnaeus, 1758: 85; Turdus Linnaeus, 1758: 85. Classis AMPHIBIA Batsch, 1788: 88. Familia TESTUDINES Batsch, 1788: 437. Genus (1): Zestudo Linnaeus, 1758: 196. Familia “Barracur’? Batsch, 1788: 437. Genera (4): Bufo Laurenti, 1768: 25; Hy/a Laurenti, 1768: 32; Pipa Laurenti, 1768: 24; Rana Linnaeus, 1758: 196. Familia “LACERTAE”* Batsch, 1788: 437. Genera (13): Basiliscus Laurenti, 1768: 50; Caudiverbera Laurenti, 1768: 43; Chamaeleo Laurenti, 1768: 45 (as Chamaeleon); Cordylus Laurenti, 1768: 51; Crocodylus Laurenti, 1768: 53; Draco Linnaeus, 1758: 196; Gekko Laurenti, 1768: 43; [Iguana Laurenti, 1768: 47; Salamandra Laurenti, 1768: 41; Scincus Laurenti, 1768: 55; Seps Laurenti, 1768: 58; Ste//io Laurent, 1768: 56; Triton Laurenti, 1768: 37. [Lacerta* Linnaeus, 1758: 196]. Familia “SERPENTES’* Batsch, 1788: 437. Genera (16): Amphisbaena Linnaeus, 1758: 196; Anguis Linnaeus, 1758: 196; Aspis Laurenti, 1768: 105; Boa Linnaeus, 1758: 196; Caecilia Linnaeus, 1758: 229; Caudisona Laurenti, 1768: 92; Cerastes Laurenti, 1768: 81; Cobra Laurenti, 1768: 103; Coluber Linnaeus, 1758: 196; Constrictor Laurenti, 1768: 106; Coronella Laurenti, 1768: 84; Dipsas Laurenti, 1768: 89; Laticauda Laurenti, 1768: 109; Naja Laurenti, 1768: 90; Natrix Laurenti, 1768: 73; Vipera Laurenti, 1768: 99. [Serpens* Garsault, 1764: pl. 667]. Classis Pisces Batsch, 1788: 88. Familia “Muzrirorr’° Batsch, 1788: 483. Genera (3): Petromyzon Linnaeus, 1758: 196; Raja Linnaeus, 1758: 196; Squalus Linnaeus, 1758: 196. Familia “Monsrrosr’° Batsch, 1788: 483. Genera (2): Chimaera Linnaeus, 1758: 196; Lophius Linnaeus, 1758: 196. Familia “Gzosarr’° Batsch, 1788: 484. Genera (3): Diodon Linnaeus, 1758: 243; Ostracion Linnaeus, 1758: 243; Tetrodon Linnaeus, 1758: 243. Familia “ArTicuLATr’? Batsch, 1788: 484. Bonn zoological Bulletin 57 (2): 149-171 ©OZFMK 166 Alain Dubois & Roger Bour Genera (3): Fistularia Linnaeus, 1758: 243; Pegasus Linnaeus, 1758: 243; Syngnathus Linnaeus, 1758: 243. Familia “Lorica7r’® Batsch, 1788: 484. Genera (4): Acipenser Linnaeus, 1758: 196; Centriscus Linnaeus, 1758: 243; Cyclopterus Linnaeus, 1758: 242; Loricaria Lin- naeus, 1758: 243. Familia “SPECULARES”’° Batsch, 1788: 484. Genera (7): Callionymus Linnaeus, 1758: 242 (as Callyonimus); Cottus Linnaeus, 1758: 242; Gobius Linnaeus, 1758: 242; Scor- paena Linnaeus, 1758: 242; Trachinus Linnaeus, 1758: 242; Uranoscopus Linnaeus, 1758: 242; Zeus Linnaeus, 1758: 242. Familia “SoveaTr’* Batsch, 1788: 484. Genera (3): Balistes Linnaeus, 1758: 243; Chaetodon Linnaeus, 1758: 242; Pleuronectes Linnaeus, 1758: 242. [“Soleas: Ed- wards in Catesby, 1771]. Familia “Ferr’° Batsch, 1788: 485. Genera (11): Coryphaena Linnaeus, 1758: 242; Esox Linnaeus, 1758: 243; Gasterosteus Linnaeus, 1758: 242; Labrus Linnaeus, 1758: 242: Mullus Linnaeus, 1758: 243; Perca Linnaeus, 1758: 242; Salmo Linnaeus, 1758: 243; Sciaena Linnaeus, 1758: 242; Scomber Linnaeus, 1758: 243; Sparus Linnaeus, 1758: 242; Trigla Linnaeus, 1758: 243. Familia “BRACTEATI’® Batsch, 1788: 485. Genera (5): C/upea Linnaeus, 1758: 243; Cyprinus Linnaeus, 1758: 243; Exocoetus Linnaeus, 1758: 243; Mugil Linnaeus, 1758: 243; Polynemus Linnaeus, 1758: 243. Familia “Nupr’° Batsch, 1788: 485. Genera (7): Anarhichas Linnaeus, 1758: 242; Blennius Linnaeus, 1758: 242; Cobitis Linnaeus, 1758: 243; Echeneis Linnaeus, 1758: 242; Gadus Linnaeus, 1758: 242; Silurus Linnaeus, 1758: 243; Xiphias Linnaeus, 1758: 242. Familia “SERPENTINI’° Batsch, 1788: 485. Genera (4): Ammodytes Linnaeus, 1758: 242; Gymnotus Linnaeus, 1758: 242; Muraena Linnaeus, 1758: 242; Trichiurus Lin- naeus, 1758: 242. “Superclassis” CRUSTACEA Batsch, 1788: 84. Classis INSECTA Batsch, 1788: 89. Familia “COLEOPTERA’®° Batsch, 1789: 539. Genera (21): Attelabus Linnaeus, 1758: 342; Buprestis Linnaeus, 1758: 342; Byrrhus Linnaeus, 1766: 537; Cantharis Linnaeus, 1758: 342; Carabus Linnaeus, 1758: 342; Cassida Linnaeus, 1758: 342; Cerambyx Linnaeus, 1758: 342; Chrysomela Lin- naeus, 1758: 342; Cicindela Linnaeus, 1758: 342; Coccinella Linnaeus, 1758: 342; Curculio Linnaeus, 1758: 342; Dermestes Linnaeus, 1758: 342; Elater Linnaeus, 1758: 342; Hydrocantharus+ Batsch, 1789: 550; Lampyris Geoffroy, 1762: 165; Mordel- la Linnaeus, 1758: 342; Necydalis Linnaeus, 1758: 342; Nicrophorus Fabricius, 1775: 71; Scarabaeus Linnaeus, 1758: 342; Silpha Linnaeus, 1758: 342; Tenebrio Linnaeus, 1758: 342. Familia “HEM/pPTERA’® Batsch, 1789: 539. Genera (5): Blatta Linnaeus, 1758: 342; Forficula Linnaeus, 1758: 342; Gryllus Linnaeus, 1758: 342; Meloe Linnaeus, 1758: 342; Staphylinus Linnaeus, 1758: 342. Familia “NEvRoPTERA’® Batsch, 1789: 539. Genera (7): Ephemera Linnaeus, 1758: 343; Hemerobius Linnaeus, 1758: 343; Libellula Linnaeus, 1758: 543; Myrmeleon Lin- naeus, 1767: 539 (as Myrmeleo); Panorpa Linnaeus, 1758: 343; Phryganea Linnaeus, 1758: 343; Raphidia Linnaeus, 1758: 343. Familia “Hymenoprera’® Batsch, 1789: 540. Genera (9): Apis Linnaeus, 1758: 343; Chrysis Linnaeus, 1761: xlii; Cynips Linnaeus, 1758: 343; Formica Linnaeus, 1758: 343; Ichneumon Linnaeus, 1758: 343; Sirex Linnaeus, 1761: xli; Sphex Linnaeus, 1758: 343; Tenthredo Linnaeus, 1758: 343; Ves- pa Linnaeus, 1758: 343. Familia “Diprera’® Batsch, 1789: 540. Genera (10): Asi/us Linnaeus, 1758: 344 (as Asylus); Bombylius Linnaeus, 1758: 344; Conops Linnaeus, 1758: 344; Culex Lin- naeus, 1758: 344; Empis Linnaeus, 1758: 344; Hippobosca Linnaeus, 1758: 344; Musca Linnaeus, 1758: 344; Oestrus Lin- naeus, 1758: 344; Tabanus Linnaeus, 1758: 344; Tipula Linnaeus, 1758: 344. Familia CivicariA Batsch, 1789: 540. Genera (3): Cimex Linnaeus, 1758: 343; Nepa Linnaeus, 1758: 343; Notonecta Linnaeus, 1758: 343. Familia CicabiNa Batsch, 1789: 540. Genera (6): Aphis Linnaeus, 1758: 343; Chermes Linnaeus, 1758: 343; Cicada Linnaeus, 1758: 343; Coccus Linnaeus, 1758: 343; Fulgora Linnaeus, 1766: 538; Thrips Linnaeus, 1758: 343. Familia “LEPIDOPTERA”? Batsch, 1789: 540. Genera (10): Alucita Linnaeus, 1758: 496; Bombyx Linnaeus, 1758: 495; Geometra Linnaeus, 1758: 496; Papilio Linnaeus, 1758: 343; Phalaena Linnaeus, 1758: 343; Pyralis Linnaeus, 1758: 496; Sphinx Linnaeus, 1758: 343 (as Sphynx); Tinea Linnaeus, 1758: 496; Tortrix Linnaeus, 1758: 496; Zygaena Fabricius, 1775: 550. Familia “Hexv4Popa”’”° Batsch, 1789: 540. Genera (4): Lepisma Linnaeus, 1758: 344; Pediculus Linnaeus, 1758: 344; Podura Linnaeus, 1758: 344; Pulex Linnaeus, 1758: 344. Familia “PoLypopa”’® Batsch, 1789: 540. Genera (10): Acarus Linnaeus, 1758: 344; Aranea Linnaeus, 1758: 344; Cancer Linnaeus, 1758: 344; Gammarus Fabricius, 1775: 418; Julus Linnaeus, 1758: 344 (as ulus); Monoculus Linnaeus, 1758: 344; Oniscus Linnaeus, 1758: 344; Phalangium Lin- naeus, 1758: 344; Scolopendra Linnaeus, 1758: 344; Scorpio Linnaeus, 1758: 344. Classis VERMES Batsch, 1788: 89. Familia “/ivrestina’’° Batsch, 1789: 664. Bonn zoological Bulletin 57 (2): 149-171 ©ZFMK Family- and class-series nomina in zoology 167 Genera (8): Ascaris Linnaeus, 1758: 644; Cucullanus Miller, 1777: pl. 38 fig. 1-7; Echinorynchus Muller, 1776: 214 (as Echi- norhynchus),; Gordius Linnaeus, 1758: 644; Hirudo Linnaeus, 1758: 644; Hydatigena Goeze, 1782: 192; Taenia Linnaeus, 1758: 646; Trichuris Roederer, 1761: 243. Familia “SETIPEDA”° Batsch, 1789: 664. Genera (4): Aphrodita Linnaeus, 1758: 644; Lumbricus Linnaeus, 1758: 644; Nais Miller, 1771: 6; Nereis Linnaeus, 1758: 644. Familia “UBERES”° Batsch, 1789: 665. Genera (6): Argonauta Linnaeus, 1758: 645; Clio Linnaeus, 1767: 1072; Lernaea Linnaeus, 1758: 644; Nautilus Linnaeus, 1758: 645; Scyllaea Linnaeus, 1758: 644; Sepia Linnaeus, 1758: 644. Familia Limacina Batsch, 1789: 665. Genera (25): Aplysia Linnaeus, 1767: 1072 (as Laplysia); Buccinum Linnaeus, 1758: 645; Bulla Linnaeus, 1758: 645; Cassis Scopoli, 1777: 393; Chiton Linnaeus, 1758: 645; Conus Linnaeus, 1758: 645; Cylindrus+ Batsch, 1789: 692; Cymbium Mendes da Costa, 1776: 182; Cypraea Linnaeus, 1758: 645; Doris Linnaeus, 1758: 644; Fasciola Linnaeus, 1758: 644; Haliotis Lin- naeus, 1758: 645; Helix Linnaeus, 1758: 645; Limax Linnaeus, 1758: 644; Murex Linnaeus, 1758: 645; Nerita Linnaeus, 1758: 645; Orthoceras Bruguiére, 1789: xvi; Patella Linnaeus, 1758: 645; Purpura Bruguiére, 1789: xv; Serpula Linnaeus, 1758: 645; Strombus Linnaeus, 1758: 645; Tethys Linnaeus, 1758: 644; Turbo Linnaeus, 1758: 645; Turris+ Batsch, 1789: 691; Voluta Linnaeus, 1758: 645. Familia “SyPHoNATA’’° Batsch, 1789: 665. Genera (17): Anomia Linnaeus, 1758: 645; Arca Linnaeus, 1758: 645; Ascidia Linnaeus, 1767: 1072; Cardium Linnaeus, 1758: 645; Chama Linnaeus, 1758: 645; Mactra Linnaeus, 1767: 1073; Mya Linnaeus, 1758: 670; Mytilus Linnaeus, 1758: 645; Ostrea Linnaeus, 1758: 645; Pecten Miller, 1776: 248; Perna Philipsson, 1788: 20; Pholas Linnaeus, 1758: 645; Pinna Lin- naeus, 1758: 645; Solen Linnaeus, 1758: 645; Spondylus Linnaeus, 1758: 645; Te/lina Linnaeus, 1758: 645; Venus Linnaeus, 1758: 645. Familia “Cris74T4”’° Batsch, 1789: 665. Genera (6): Actinia Pallas, 1766b: 152; Balanus Mendes da Costa, 1778: 249; Holothuria Linnaeus, 1758: 644; Lepas Linnaeus, 1758: 645; Medusa Linnaeus, 1758: 644; Triton Linnaeus, 1758: 644. Familia “Crustosa”° Batsch, 1789: 665. Genera (2): Asterias Linnaeus, 1758: 644; Echinus Linnaeus, 1758: 644. Familia “FRoNDOSA’’? Batsch, 1789: 665. Genera (2): Astrophyton Schultze, 1760: 53; Pennatula Linnaeus, 1758: 646. Familia “Potypina’’® Batsch, 1789: 666. Genera (10): Alcyonium Linnaeus, 1758: 646; Eschara Linnaeus, 1758: 646; Gorgonia Linnaeus, 1758: 646; Hydra Linnaeus, 1758: 646; Isis Linnaeus, 1758: 646; Madrepora Linnaeus, 1758: 646; Millepora Linnaeus, 1758: 646; Sertularia Linnaeus, 1758: 646; Spongia Linnaeus, 1759: 1317; Tubularia Linnaeus, 1758: 646. Familia “FyveriATa’’° Batsch, 1789: 666. Genera (3): Brachyonus Pallas, 1766a: 89; Trichoda Miller, 1773: 71; Vorticella Linnaeus, 1767: 1074. Familia “CHa4orica’* Batsch, 1789: 666. Genera (10): Burfaria Miller, 1773: 62; Cercaria Miller, 1773: 64; Cyclidium Miller, 1773: 49; Enchelis Miller, 1773: 33; Go- nium Miller, 1773: 60; Kolpoda Miller, 1773: 56; Monas Miller, 1773: 25; Paramaecium Miller, 1773: 54; Vibrio Miller, 1773: 39; Volvox Linnaeus, 1758: 646. [Chaos* Linnaeus, 1767: 1074]. Bonn zoological Bulletin 57 (2): 149-171 ©ZFMK VUOAINAVD “VIIVNNV]A “VLVGNOH) SANIGALSAL “VdISdOUNVS ‘VLVGNOHD VHdYOWNOAOVNINY ‘VITVAINVIA, “VLVGXOHD SAWUOAINOIHLAULS ‘VAISAOUAVS ‘VLVGYOHD VILNAGOY ‘VIIVAWVIA, “VLVCXOHD VUOAINAV ‘VITVINNINV]A] “VLIVGNOHD VILNAGOY ‘VIIVAWVIA, “VLVGNOHD VIVNOWT1Ndg ‘VdOdOULSV) ‘VOSATIO[L VHdaYOWODV’T ‘VITVININVIA, “VLVGQdOH) Alain Dubois & Roger Bour VUOAINUVD ‘VITVNWVIA, ‘VLVCaXOHD SAWNUOALOVUOD “VAISdOUNVS ‘VIVAUOHD VuALdINAH ‘VLOASN] ‘VdOdOUHLYY VuaLdINaH{ ‘VLOASNY ‘VaGOdOUHLNY VIALOVGOILYY ‘VITVAWV]] ‘VIVQXOHD VILNAGOY ‘VITVAWV]A] ‘VIVGYOHD VUOAINAVD ‘VITVAINV]A] ‘VLVGaYOHD VSOTIg ‘VIIVANV]A] ‘VLVQaOHD S8LI ‘Yyosieg gvaisyQ S8LI ‘YOsweg IVCINIGNLST QSLI ‘YOsweg IVCId TKI B8LI ‘Yosieg FVGINOIHINULS S8LI Yosieg arvariniog S8LI ‘Yosweg FraITALSAW S8LI ‘Yosieg arannpy 68L1 “YOseg APGIOVITT S8LI “Yosieg @raroddT S8LI “Yyosyeg araITAY B8LI ‘Yosieg aVAIOKYOD 68L1 “Yosweg FFAIOINID 68LI “Yosieg IralaVIID S8LI Yosieg grdiAN7D SBI “Yosieg JVARIOLSVD S8LI ‘Yosieg FrdINKD S8LI ‘Yosieg avaidodsarag TLE ‘LISI “JOYOSIY INISU) LEv ‘88L1 Yosweg SINIdALSAL TLE LIST IOYOSIA INTEL JOP ‘STR ‘SIOSIA, FVCINOIHLNULS TLE ‘LIST BOYosty mag ZLE LIST ‘IOyosty INITILSAW OF “SIT TOSI] KNIZAY ZO (ORL “oteule’ 7] SYFIOKNIT TLE (LIST “JOYOSIY INIIOdTT TLE ‘LIST FOYOsty INITIY 66€ :ST8I “SIOBIA VNFIOFYOD OFT :ZOST “ANT SIGIOINID CTH ‘OPI “POOMISSM FVAKILD PLe ‘O78 ‘SSNIplOH KNIANTD €€ (OTSI “Yotduiay VNIYOLSKD TLE ‘LIST AOydsty ININKD POE FIT8I ‘ABI avaidsarag LIST *tULoYypleAA op IOyOSly,, JVGISYA «88LI ‘YOsieg FVCINIGALSAL LIST *WHSYpIeAA eP JoyOSly,, IVCId TKI LIST *.WULOYpleA\ ep JOYOSIy,, JVCINOIHLNULS LIST ‘.WHoypreM op JOYOsly,, IFAMAIIS LIST *.WHoypleM op JOYOSIJ,, FAITALSAP SIS] TOs] gran LOST ‘yoreue] avCIOKNIT LIST *..WuIoUpleM op JOIST, TVCIMOdTT LIST *.CuloypleA ep JOYOSIy,, IVCITI CTZ8I ‘SIOSIA FVAIOKYOD TOSI ‘ay[lo-ne] AVGIOINID OFS I ‘POOMISAM FFAIGKIID OZ8I “SSIYP[OH avaiuuTD OZ8I “Youduiay FraIVOLsKD LIST *.CuloypleA ep JOyoSty,, JVCINVD ITS ‘AvIQ avaidodsarag QSL] ‘sneeuury snsuQ BBL ‘yosyeg FNISUAQ SSL ‘SnaeuUry opnisal Bgl ‘Yosweg SINIGALST SSL] ‘snoeuury vdjny QSLT YOsweg FNIdTKT BSL] ‘snoeuUry O17n.15 B8/ | “YOSIeY SINOIHLAULS SSL ‘snoeuury snuniog == ST “Yosieg FNIZAIDS SSLI Yoseg FNITILSAW S8LI ‘Yosieg FNIZQW 68L1 “YOsyeg PNIOKWIT SLI ‘sneeuury vjajsnpy SSL] ‘ShevuuryT snpy QSL ‘snoeuury xpuT SSLI ‘snevuury sndaT — gL { “yosIeg VNIYOdTT S8LI Yosteg rPNITAY S8LI “Yosieg s7ory0D 68LI Yosweg FrRIFOINID 68LI “Yosreg FNIGKOID S8LT “Yosieg KNIAYTD S8LI ‘Yosieg rAYOLSKD SSL ‘sneeuury] say SSL] ‘sheeuury sp1on410) SLI ‘Snovuury xaw1y SSL] ‘snovuury vppo1g SCL] ‘snevuury siasay SCL] ‘snaeuury 1ojspy QSL ‘snovuuryT sip) S8LI ‘Yyosieg FNINKD QSL ‘snevuury snddpoig g8/ | ‘yosieg raodsarag Opso ‘sisseyo ‘unyAyd :exe) JOYysTy PeUONIpPeLL A]MuUey SI JO UOWIOU pIeA ATW} SI} 1OJ pasn A][euoMIpe.y UdWIOU JO WIAUOJOIg Awe} SITY} 1OF pasn A]JeuOn pel) USWION (68L1 ‘88Z1) yos}eg ul snuasosjonN Suriwodde uawiou Ayrwey OZFMK ‘(RQQOT SlOgnd) SWeU Sty} JopuN pojId aq prnoys pue ,.JOYOSTy,, MWe a]SuIs ay} Aq pousdis AT[eNjIOw SI WIOYP]eAA YP JOYOSIJ,, 0} JIS UOSTUIOU -OXxB], OY} Ul poyposd uOneKo[gnd sy “(/PpIM/S1o'erpadryimus//:dyy) erpodryl Ay OS OY} Ul PoZIUBOda1 SsOU} SAIS OM ‘,,YSLIO}Se UL Aq PIYACUI ‘dJIS STU} UL UDATS JOU de dJep pue diys -IOYyINe YSIYM JOJ suo ATUO 9} IO ‘(/[UAWOUOXe)] UOSTWOUOXe}//:dyY) UOSTWIOUOXRT, SY} JO IIS IY} Ul PAZIUBOIaI dso} 91B SULUNIOS JseT] PUL PITY] OY} UI USATS exe} JOYSIY pue soiql -WUJ JO PUTWIOU [BUOTIPe.Y sy], “AWOUOXk}00Z UI WY} 0} Popo1d AT[RUOTIPe.Y Sayep PUR SIOYING OY} YIM “(68/T “VSL 1) YOsie Ur o[quyileae opeUl PUILUIOU SdTIds-ATIWIR] OY, *E BTQUL, 168 Bonn zoological Bulletin 57 (2): 149-171 Family- and class-series nomina in zoology 169 APPENDIX 2 THE NOMENCLATURAL STATUS OF A FEW PROBLEMATIC ZOOLOGICAL NOMINA THE ZOOLOGICAL NOMINA CREATED IN THE BOOK OF BEHN (1760) Klein (1751) published a comprehensive classification of his “QUADRUPEDIA”, 1.e., roughly, the tetrapods without the cecilians, snakes, birds and whales. This book in Latin be- ing pre-1758, the nomina it contains are nomenclaturally unavailable. Joyce et al. (2004) pointed out the existence of Behn’s (1760) German translation and adaptation of Klein’s (1751) book, where all the taxa and nomina of the latter work are reproduced. These post-1758 nomina would be available, with the authorship “Klein in Behn, 1760”, if this book had to be considered nomenclatural- ly available, but it should not. The nomenclatural hierar- chy used in this book is unclear and inconsistent. It in- cludes the ranks ordo (Ordnung) and familia (Familie), the taxa at these ranks being designated by uninomina (nom- ina consisting of a single term). The ranks used below the rank Familie are denominated in German Geschlecht, then Art, then Gattung. Considering their hierarchy and con- tent, they could be construed to correspond respectively to the ranks tribe, genus and species, but this would prob- ably be misleading. Each of these ranks can contain a var- ious numbers of unnamed subranks, and the number of terms used to designate taxa is variable, from one to two and more, some of these nomina being plurinominal di- agnoses borrowed without change from various pre-1758 works. This work clearly does not comply with the re- quirement of Article 11.4 of the Code for the availability of species-, genus- and family-series nomina. However, this might not preclude considering the class-series nom- ina in this work, or some of them, as available, since Ar- ticle 11.4 implicitly states that “this Article does not ap- ply to the availability of names of taxa above the family group”. If it was possible to establish objectively where lays the separation between the family-series and the class-series nomenclature in Behn (1760), and if all these nomina were uninomina, it could be possible to recognize as available the class-series nomina proposed in this work, but this is difficult if not impossible. The nomina of the three orders of “QUADRUPEDIA” rec- ognized in Behn (1760) are plurinomina, as follows: (O1) “Pilosa et Ungulata (vivipara) sive “Zaotoxa”’; (O02) “Pi- losa et Digitata sive sint tota coriacea, sive cataphracta; omnia vivipara”’; (O03) “Depilata, sive tecta, sive nuda, nequicquam pilosa, omnia ovipara, sive “Qotoxa”. Such designations are in fact diagnoses, and cannot qualify as Bonn zoological Bulletin 57 (2): 149-171 nomina of zoological taxa. They are unavailable in zoo- logical nomenclature. It can be noted that, in the original text of Klein (1751), the same taxa were designated by uni- nomina (“UNGULATA”, “DIGITATA” and “DEPILATA’’), but as this text is pre-Linnaean, these nomina also are unavail- able. In contrast, the nomina of the 13 “families” recognized by Behn (1760) are all uninomina. They are distributed as follows in the three orders: (O1) ‘“MONOCHELON”, “DICHELON”, “TRICHELON”’, ““TETRACHELON” and “PEN- TACHELON”; (O2) “DIDACTYLON”, “TRIDACTYLON”, “TETRADACTYLON’, “PENTADACTYLON” and “ANOMALA- PES” (instead of “ANOMALOPES” in Klein, 1751); (O3) ‘“TESTUDINATA ”, “CATAPHRACTA” and “NUDA”. Except pos- sibly for one, these nomina are not based on the stems of included nominal genera. “TESTUDINATA” could be con- strued to be based on the stem of the only included genus of the family, Zestudo Linnaeus, 1758, but this is highly improbable. The other twelve familial nomina are clear- ly based on characters that are considered diagnostic for the taxa they designate, and the nomen “TESTUDINATA” can also be understood as based on the Latin adjective testu- dinatus, meaning “of turtle, vaulted, arched”. Therefore, all nomina of “families” in Behn (1760) appear to be arhi- zonyms. Under the Rules of Dubois (2006a), such nom- ina cannot be accepted as family-series nomina and qual- ify as class-series nomina. This case is not unique. Other examples were discussed by Dubois (2006a, 2009) and Dubois & Ohler (2009): for example, the nomina of “fam- ilies” in Ritgen (1828), which are also arhizonyms, must be treated as available class-series nomina. However, in the case of the new familial nomina appear- ing in Behn (1760), difficulties would arise if they were to be treated as available class-series nomina. In the sys- tem of Dubois (2006a), the allocation of class-series nom- ina to taxa is made through their included (conucleogen- era) and excluded (alienogenera) nominal genera, and to be usable in this respect, conucleogenera and alienogen- era must be nomenclaturally available. If all the nomina of taxa just below the rank family in Behn (1760), desig- nating taxa of rank “Geschlecht”, were considered to be genus-series nomina, part of them could not be used for taxonomic allocation of their nomina, because they are un- available in Behn’s (1760) work. In his order (O3), cor- responding to the traditional amphibians and reptiles, on- ly three generic nomina then available are mentioned as valid nomina: 7estudo Linnaeus, 1758 for a “Geschlecht” of his family “TESTUDINATA”; Lacerta Linnaeus, 1758 for a “Geschlecht” of his family “NupA”; Rana Linnaeus, 1758 for an “Art” of his “Geschlecht” “Batrachus” (then an unavailable nomen) of his family “NuDA”’; and none in his family “CATAPHRACTA”. In order to allocate the nomen “NuDA” to a class-series taxon, one would have ©ZFMK 170 Alain Dubois & Roger Bour to take an arbitrary decision, considering that either the rank “Geschlecht” or the rank “Art” corresponds to the rank genus in the current Code. If the rank “Geschlecht” was considered to correspond to the rank genus, and “Art” to the rank species, the nomen “NUDA” would apply, in a modern classification, to the most inclusive taxon includ- ing the genus Lacerta and excluding the genus Jestudo. But if the rank “Geschlecht” was considered to correspond to the rank tribe, and “Art” to the rank genus, the nomen “NuDA” would apply, in a modern classification, to the most inclusive taxon including the genus Rana and exclud- ing all the mammalian genera, bearing then available Lin- naean generic nomina, mentioned by Behn (1760) in his orders (O1) and (O2). Therefore, according to the arbitrary decision taken, the same nomen could apply to widely dis- tinct higher taxa. Because of these uncertainties, many other examples of which could be given, we here argue that Behn’s (1760) should not be considered as an available work in zoolog- ical nomenclature, even for class-series nomina. We sug- gest that this book should be invalidated as a whole by the ICZN, and that all the new nomina it contains should be considered unavailable in zoological nomenclature. *Marsupiale” Edwards in Catesby, 1771 According to Sherborn (1902: 593), there exists a genus Marsupiale, based on the following reference: “G. Ed- wards in M. Catesby, Carol. I. 1771, xxix”. Actually this refers to Catesby (177 1a: xxix), in “An account...” added by George Edwards, where the binomen Marsupiale amer- icanum appears, with a diagnosis. However, this item fol- lows another one entitled Vulpi affinis americana and many others where the nomenclature is not consistently binominal. Consequently the ICZN (Anonymous 1954) has suppressed the whole work (Catesby 1771a-b) for nomenclatural purposes, except for the nomina employed by Edwards in accordance with the Linnean system in his “Catalogue of the Animals and Plants” (i.e., Catesby 1771a: 1-2, 1771b: 1-2), usually referred as George Ed- wards’ “Appendix”. “Solea” Edwards in Catesby, 1771 According to Sherborn (1902: 593), there exists a genus Solea, based on the following reference: “G. Edwards in M. Catesby, Carol. II. 1771, 27”. Actually this refers to Catesby (1771b: 27), where appears the combination Solea lunata et punctata, with a diagnosis and a plate; howev- er, this is not a binomen, and therefore it has no status in nomenclature. The ICZN (Anonymous 1954) has sup- pressed the whole work (Catesby 1771a-b) for nomenclat- Bonn zoological Bulletin 57 (2): 149-171 ural purposes, except for the nomina employed by Ed- wards in accordance with the Linnean system in his “Cat- alogue of the Animals and Plants” (i.e., Catesby 1771a: 1-2, 1771b: 1-2), usually referred as George Edwards’ “Appendix”. Edwards (in Catesby 1771b: 1) linked this description with the binomen Pleuronectes lunatus Lin- naeus, 1758. “Gralla” Eberling in Sonnerat, 1777 According to Sherborn (1902: 431), there exists a genus Gralla, based on the following reference: “J. P. Ebeling in Sonnerat, Reise Neuguinea, 1777, 31”. Actually this refers to Sonnerat (1777: 31 [and 45]), where appears the combinations gralla parra and gralla fulica. Wieland (2010) admitted the nomenclatural availability of both, which he treated as binomina, and also of the genus Gral- la Sonnerat, 1777, but with this comment: “The basic da- ta of this taxon were not entered consulting the original description, but from secondary sources”. On the other hand, The Richmond Index, published by the Division of Birds at the National Museum of Natural History, Wash- ington, D.C (Anonymous 2010), states that Gralla Ebel- ing in Sonnerat is not nomenclaturally a valid generic name: “Gralla fulica p. 45; Gralla parra p. 31, Ebeling, in Sonnerat, Reise Neu Guinea, 1777. These have no stand- ing! being simply Ebeling ’s way of writing Order Gral- le, Genus Fulica + Parra/!’’. Actually Ebeling (in Son- nerat 1777) put a capital at the start of the generic name of his binomina, but neither at gralla parra nor at gralla fulica. We follow here The Richmond Index statement and do not recognize the nominal genus “Gralla Ebeling in Sonnerat, 1777”. Cylindrus Batsch, 1789: 692 Three homonymous nominal genera Cylindrus are avail- able in zoological nomenclature: Cylindrus Batsch, 1789: 692; Cylindrus Deshayes, 1824: 236; and Cylindrus Fitzinger, 1833: 107. Cylindrus Batsch, 1789 has apparently been ignored by all authors until now. It was introduced with a diagnosis that makes it nomenclaturally available and that clearly points to marine cone shells. Cylindrus Deshayes, 1824 is an autoneonym* (unjustified emendation) of Cylinder Denys de Montfort, 1810: 390, a nomen established for a genus of marine cone shells. Its nucleospecies* (type-species) is Conus textile Linnaeus, 1758: 717, by original designation. The original nomen of this genus was preceded in zoological nomenclature by Cylinder Voet, 1793 and Cylinder Voet, 1806, but both ©ZFMK Family- and class-series nomina in zoology 171 these nomina are unavailable, as published in books that are not consistently binominal. Strangely enough howe- ver, the nomen Cylinder Denys de Montfort, 1810 is cur- rently not considered valid, but its autoneonym Cylindrus Deshayes, 1824 is so, being currently treated as a subge- nus of the genus Conus Linnaeus, 1758 (e.g., Keen 1971; Pitt et al. 1986). Cylindrus Fitzinger, 1833 was established with a single valid species included, Pupa obtusa Draparnaud, 1805: 63, which is therefore its nucleospecies by original specific monophory* (monotypy). This generic nomen is current- ly (e.g., Frank 2006) considered valid for a genus of ter- restrial snails. The current nomenclatural situation concerning the use of the term Cylindrus in zoological nomenclature is not com- pliant with the Rules of the Code, for two distinct reasons: (R1) the autoneonym Cylindrus Deshayes, 1824 of Cylin- der Denys de Montfort, 1810 is considered valid instead of its archaeonym*, although the latter should be so, not being preoccupied by an available homonymous generic nomen; (R2) two homonymous genus-series nomina, Cy- lindrus Deshayes, 1824 and Cylindrus Fitzinger, 1833, are currently both considered valid in zoology, although the second one, being a junior homonym of the former, should be considered invalid (even if the former one was not so). The two nomina are listed as valid in several current on- line databases, but apparently never in the same one: Cy- lindrus Deshayes, 1824 appears as the valid nomen of a subgenus of Conus Linnaeus, 1758 in the databases Ca- talogue of recent and fossil Conus (Alan J. Kohn) [http://biology.burke.washington.edu/conus/recordview/sp ecieslist P.html], The sea shells (Nauka Bulgarie) [http://theseashells.nauka.bg/Conus_Cylindrus_ textile tex tile-html] and Hardy’s Internet Guide to marine Gastro- pods (Eddie Hardy) [http://jeh-temp.co.uk/Taxon_pages /Family CONIDAE CONINAE.shtml], whereas Cylin- drus Fitzinger, 1833 appears as the valid nomen of a ge- nus of terrestrial snails in the databases Molluscs of cen- tral Europe (Dr. Vollrath Wiese, Cismar, D-23743 Gro- mitz-Cismar) |http://www.mollbase.de/list/liste.php], Ani- malbase Goettingen [http://www.animalbase.uni-goettin- gen.de/zooweb/servlet/AnimalBase/search] and Biolib.cz [http://www.biolib.cz/en/taxon/1d18384]. The rediscovery of the nomen Cylindrus Batsch, 1789, created for a genus of marine cone shells, allows to cla- rify this nomenclatural situation. We hereby designate Conus textile Linnaeus, 1758 as its nucleospecies (type- species). The nomen Cylindrus Batsch, 1789 therefore re- places both Cylinder Denys de Montfort, 1810 and Cy- lindrus Deshayes, 1824 as the valid nomen of the subge- nus of Conus Linnaeus, 1758 including the latter species. As for Cylindrus Fitzinger, 1833, it is an invalid junior ho- Bonn zoological Bulletin 57 (2): 149-171 monym of both Cylindrus Batsch, 1789 and Cylindrus Deshayes, 1824 and it must be abandoned. The homonymy between Cylindrus Deshayes, 1824 and Cylindrus Fitzinger, 1833 was pointed out by Kennard (1942), in a work that seems to have been overlooked by most subsequent authors. This author rightly concluded that the nomen Cy/indrus Fitzinger, 1833 is invalid, and pointed to the existence of its senior objective synonym Cochlopupa Jan, 1830: 5. The nucleospecies of this no- minal genus is Pupa obtusa Draparnaud 1805 by original specific monophory. The single species currently referred to the genus Cylindrus Fitzinger, 1833 and known as Cy- lindrus obtusus, must therefore bear the nomen Cochlo- pupa obtusa (Draparnaud, 1805). Hydrocantharus Batsch, 1789: 550 The nomen Hydrocantharus Batsch, 1789, created for an aquatic beetle (dytiscid) is identical to several pre-1758 uses of the same nomen, which are nomenclaturally una- vailable. For the same genus, Linnaeus (1758: 342) used the nomen Dytiscus. In this genus, he listed (p. 411-413) 15 nominal species, among which Latreille (1810: 426) designated Dytiscus marginalis Linnaeus, 1758: 411 as nu- cleospecies. We hereby designate the same nominal spe- cies as nucleospecies (type-species) of Hydrocantharus Batsch, 1789, which therefore becomes an invalid junior objective synonym of Dytiscus Linnaeus, 1758. Turris Batsch, 1789: 691 A generic nomen Jurris was created for a gastropod ge- nus by Statius Miller (1766: 129), but this nomen is una- vailable as having been published in a book invalidated by the ICZN (Anonymous 1964) as not applying the prin- ciple of binominal nomenclature. A homonymous nomen Turris was later created by R6ding (1798: 123) also for a gastropod genus, and this nomen is currently considered valid. However, the present rediscovery of Zurris Batsch, 1789 makes Turris Roding, 1798 its invalid junior syno- nym. As reckoned by Winckworth (1945), the nucleospecies of Turris Roding, 1798 is Murex babylonius Linnaeus, 1758: 753, by subsequent designation of Bucquoy et al. (1883: 86). In order not to upset nomenclatural stability, we he- reby designate Murex babylonius Linnaeus, 1758 as nu- cleospecies (type-species) of Turris Batsch, 1789. The lat- ter must now replace its junior objective synonym Turris Roding, 1798 as the valid nomen of the genus. ©OZFMK i aang iain, rare - . - —s eae = ae = act snittiia dander et, - =r Poe lel Pa ial > oi pulp od aan ss 1) one Pipi toceide ulin | ; Hl x0 7 i | H | | { t fi | | , || ‘vai | ! pa , i . if ; = iiwen won Teaie! on « ‘xy i » Leh phim eee a 7 ‘ion | oii | aS | 4 | a iG 4 he yale ' hee ee rae et - KF ; 7 weri¢ ri) te a pili ian ety r mele Tgeltatiey MOR, ee fe " ier H4toxnaal rie hi Bonn zoological Bulletin | Volume 57 | Issue 2 pp. 173-176 Bonn, November 2010 An addition to the East African herpetofauna: the first record of Tarentola annularis relicta (Squamata: Gekkonidae) in Uganda Miloslav Jirkt!, Andrei Daniel Mihalca?, Petr Necas} & David Modry!4 | Biology Centre, Academy of Sciences of the Czech Republic, Ceské Budéjovice, Czech Republic 2 University of Agricultural Sciences and Veterinary Medicine, Cluj-Napoca, Romania 3 SvinoSsice, Czech Republic 4 University of Veterinary and Pharmaceutical Sciences, Brno Czech Republic Abstract. This is the first record of a member of the genus Zarentola from Uganda. Population of Zarentola annularis relicta was found in Ubbi village on the South-Western foot of Mt. Otzi in northern Uganda, close to the border with Su- dan. Brief comments on extraordinary biogeographical affinities of the area and characterization of the Mt. Otzi envi- ronments are provided. Key words: Zarentola, Uganda, East Africa, Mt. Otzi, new geographic record. INTRODUCTION In their account of the East African reptiles, Spawls et al. (2004) listed 55 species of geckoes in 11 genera for East Africa including Tanzania, Kenya, Uganda, Rwanda and Burundi. Although biogeographically, the East African realm includes also parts of South Sudan and Ethiopia, we follow the above mentioned political delimitation of East Africa used throughout literature for practical reasons. During a short survey of the Mt. Otzi area in the very North of Uganda on 9.X.2006, we collected specimens of geckoes, that did not match any known East African genus. Specimens were collected on buildings of the village school in Ubbi, a small settlement at the South-Western foot of Mt. Otzi. Upon collection, the animals were pho- tographed and released at the original collection site. The Ubbi village (03°35’07”N, 31°49°42”E, elevation of 690 m a.s.1.) is surrounded by a mosaic of farmland, small rocky hillocks covered by bushy vegetation and numer- ous rocky outcrops devoid of vegetation (Fig. 1a). The area is situated in the Southern Sudanian savanna zone, the nat- ural vegetation of which is mostly tree and shrub moist savanna. The dominant geomorphological feature of the area is the Mt. Otzi massif steeply rising above surround- ing landscape. The Mt. Otzi located on the Western bank of the Nile River has an undulating top plateau with Bonn zoological Bulletin 57 (2): 173-176 several emergent rocky peaks reaching a maximum ele- vation of 1565 m. Its slopes are covered by bush, while its upper parts are covered by a mosaic of highland bush, moist forest and farmland (Fig. 1b). The remaining for- est patches are mostly degraded by logging and clearing for agriculture and thus characterized by relatively open canopy (Fig. 1c). The area south of the Mt. Otzi is dom- inated by palm- and other moist-savanna types (Fig. Id). Tarentola specimens The geckoes were assigned to the genus 7arentola accord- ing to absence of claws on digits 1, 2 and 5 and presence of claws on digits 3 and 4 (as seen in Figs 2a—b). The on- ly Tarentola species occurring thus far in the South-East of the genus range is Zarentola annularis (Geoffroy, 1809), in which two subspecies are recognized (Joger 1984). The nominotypic Zarentola annularis annularis (Geoffroy, 1809) occurs throughout the Saharan region, whereas Tar- entola annularis relicta Joger, 1984 is known only from two disjunct areas — the Nile valley in the very south of Sudan and Mora in the North Cameroon south of the Lake Chad. Two confirmed Sudan localities include Juba (type locality of the subspecies) and Nimule (Joger 1984). ©ZFMK 174 Miloslav Jirkt et al. Fig. 1. | Landscape and vegetation of the Mt. Otzi and adjacent areas. Fig. la. Rocky outcrops in Ubbi village on the foot of Mt. Otzi. Photograph A. D. Mihalca. Fig. 1b. Forest patch on the Mt. Otzi top plateau with an emergent rocky peak in the background. Photograph M. Jirkt. Fig. le. Interior of the Mt. Otzi forest showing relatively open canopy and distinct Afromontane floristic el- ement, the false banana of the genus Ensete in the foreground. Photograph M. Jirku. Fig. 1d. Palm savanna south of the Mt. Otzi region in the Murchinson Falls NP, Uganda. Photograph M. Jirkd. Bonn zoological Bulletin 57 (2): 173-176 ©ZFMK First record of Zarentola from Uganda 175 [> Fig. 2. Zarentola annularis. Fig. 2a. Adult specimen of Zarentola annularis relicta from Ubbi, Uganda. Photograph D. Modry. Fig. 2b. Close-up of adult specimen of Zarentola annularis relicta from Ubbi, Uganda. Note the bright red trombiculid mites lo- calized mainly around eye. Photograph D. Modry. Fig. 2c. Adult specimen of Zarentola annularis annularis from Awash NP, Ethiopia. Note the four distinct white, dark-rimmed scapular flecks. Photograph P. Necas. Apart from details in scaling patterns, 7! a. relicta can be readily distinguished visually by an absence of four white, dark-rimmed scapular flecks typical for the nominotypic subspecies (compare Figs 2a—b with Fig. 2c). Based on Bonn zoological Bulletin 57 (2): 173-176 coloration pattern, the specimens we collected in Ugan- da can be assigned to the subspecies T. a. relicta. All ob- served specimens were pale-grey colored with orange to orange-brown blotches on dorsum of the head and body ©ZFMK 176 Miloslav Jirkt et al. with intervening irregularly distributed faint whitish flecks devoid of any dark margin, whereas the tail possessed just faint grayish transversal bands (Figs 2a—b). As far as we are aware, the Figs 2a—b are the first published color pho- tographs of live 7! a. relicta. BIOGEOGRAPHICAL CONSEQUENCES To our best knowledge, the presented record of Zarento- la in Uganda is the first record of the genus in the East African region as defined above. The genus Zarentola comprises 20 species distributed throughout the dry re- gions of the Mediterranean, Middle East, some Atlantic archipelagos (e.g. Canary and Cape Verde Islands) and African mainland north of the savanna and forest zones (see Joger 1984 for review). Three additional species are known from the West Indies (Diaz & Hedges 2008, Joger 1984). The African-mainland part of the genus geograph- ic range, 1.e. Saharan region, comprises a total of six species (including Tarentola chazaliae, still treated by some authors as the only representative of the monotyp- ic genus Geckonia — see Carranza et al. 2002) the spe- ciation of which seems to reflect the relatively recent arid- ification of the Sahara desert region, which was gradual- ly colonized from its rather mesic margins by ancestors of extant species since mid Oligocene. In general, the southern distribution limit of Zarentola spp. in African mainland is delimited by an interference zone between the southern margin of the arid Sahel belt and moist savanna and forest equatorial zones. To date, the southernmost confirmed locality of Tarento- la has been the record of 7 a. relicta from Nimule (03°35733”N, 32°04’14”E), on the Sudanian side of the Sudan-Uganda border. The southern records of T a. re- licta in Sudan and Uganda document an intrusion of Sa- haran faunistic element into the relatively humid equato- rial region which is dominated by moist savanna approx- imately from 9°30’N southwards. Presence of the reptile species associated with arid habitats here, deep in the sa- vanna zone might be facilitated by a presence of an ex- tensive network of huge rocky outcrops and ridges follow- ing Aswa fault, which extend into this region from far north-west and of which the Mt. Otzi is a magnificent southernmost outpost. It is possible, that these exposed rocky formations, largely devoid of vegetation, might serve as refuges for Saharan taxa that normally would not occur thus far south in the otherwise relatively humid re- gion. Farther to the south from the Mt. Otzi region, the landscape is dominated by gently undulating plains cov- ered by various moist-savanna types (Fig. 1d) where pres- ence of Zarentola seems unlikely due to lack of suitable habitats. Bonn zoological Bulletin 57 (2): 173-176 Interestingly, the Mt. Otzi region is the easternmost known locality of a remarkable West-African savanna element, the ball python Python regius, which was collected in the vicinity of Moyo (3°39°14"N, 31°43°22”E), just 13 km to the north-west from the Ubbi village reported here to be (together with Nimule) the southernmost locality of Tarentola. Apart from the two outstanding herpeto-faunistic ele- ments, P. regius and T! a. relicta, representatives of the West African and Saharan realm(s) respectively, the re- gion is mostly inhabited by East-African herpetofauna as reflected by distribution maps provided by Spawls et al. (2004). In addition, there is a relict population of eastern chimpanzee Pan troglodytes schweinfurthii on the Mt. Otzi (Caldecott & Miles 2005), which together with colobus monkey Colobus guereza (pers. obs.) in the Otzi forest show clear faunistic affinity to the Guineo-Congo- lian rainforest block. Finally, a presence of false bananas of the genus Ensete (pers. obs., Fig. 1c) in the Otzi for- est suggests also presence of Afromontane elements in the area. In conclusion, the Mt. Otzi region deserves further attention as a potential biogeographical match point, where East- and West-African (savanna), Saharan (desert), Cen- tral-African (forest-savanna) and possibly Afromontane biotas meet at one place. Acknowledgements. We are grateful to the local people of the Mt. Otzi region, the Ubbi village in particular. Philipp Wagner and Colin McCarthy of the Zoologisches Forschungsmuseum Alexander Koenig, Bonn (Germany) and the Natural History Museum London (UK) respectively, helped us with confirma- tion of the Zarentola locality records mentioned in the paper. REFERENCES Caldecott, J. & Miles, L. (eds.). 2005. World Atlas of Great Apes and their Conservation. University of California Press, Berkeley, 404 pp. Carranza, S., Arnold, E. N., Mateo, J. A. & Geniez, P. 2002. Relationships and evolution of the North African geckos, Geckonia and Tarentola (Reptilia: Gekkonidae), based on mitochondrial and nuclear DNA sequences. Molecular Phylogenetics and Evolution 23: 244-256 Diaz, L. M. & Hedges, S. B. 2008. A new gecko of the genus Tarentola (Squamata: Gekkonidae) from Eastern Cuba. Zootaxa 1743: 43-52 Joger, U. 1984. Taxonomische Revision der Gattung Tarentola (Reptilia: Gekkonidae). Bonner zoologische Beitrage 35: 129-174 Spawls, S., Howell, K., Drewes, R. & Ashe, J. 2004. A field guide to the reptiles of East Africa. A & C Black Publishers Ltd., London, 72 pp. Received: 30.V1.2010 Accepted: 24. VIII.2010 ©ZFMK Bonn zoological Bulletin Volume 57 Issue 2 pp. 177-188 Bonn, November 2010 The taxonomic status of Hyperolius spatzi Ahi, 1931 and HAyperolius nitidulus Peters, 1875 (Amphibia: Anura: Hyperoliidae)* Mark-Oliver Rodel!, Laura Sandberger!, Johannes Penner!, Youssouph Mané? & Annika Hillers!3 ! Museum ftir Naturkunde, Leibniz Institute for Research on Evolution and Biodiversity at the Humboldt University Berlin, Invalidenstrasse 43, D-10115 Berlin, Germany; E-mail: mo.roedel@mfn-berlin.de; Phone: +49 (0)30 20938571 2 Institut de Recherche pour le Dévelopement, Dakar, Senegal 3 Across the River Project, Royal Society for the Protection of Birds, 164 Dama Road, Kenema, Sierra Leone * this paper is dedicated to Prof. Dr. Wolfgang BoOhme, who’s 1978 paper on the herpetology of Senegal induced our investigations presented herein. Abstract. We herein re-investigate the taxonomic status of Hyperolius nitidulus Peters, 1875 and H. spatzi Ahl, 1931 by means of morphology, vocalization and genetic data. Both taxa are morphologically distinct, have different advertise- ment calls and differ genetically from each other by 5.1—5.6% sequence divergence in the investigated 16S rRNA gene. Based on these data we resurrect H. spatzi as a valid species and designate a lectotype for it. Both species occur in sa- vannas of western Africa. Hyperolius spatzi is restricted to Senegambia and thus far known from Senegal and The Gam- bia. Its occurrence in Guinea Bissau and southern Mauritania seems likely. Hyperolius nitidulus ranges from Guinea and Mali eastwards into Nigeria and Cameroon. Records from the driest savannas in north-eastern Nigerian, Cameroon and the Central African Republic are doubtful and may actually refer to H. pallidus Mertens, 1940. Key Words. Bioacoustics, biogeography, genetics, morphology, savanna, West Africa. INTRODUCTION Many species of the diverse African reedfrog genus Hy- perolius Rapp, 1842 exhibit very variable color patterns (Schigtz 1971, 1975, 1999). Some of these color variations are age and sex specific (Schiotz 1967, Veith et al. 2009). As these frogs offer comparatively few other species spe- cific morphological characters, this variability caused con- siderable taxonomic confusion in the past and resulted in the description of many taxa which are now regarded as synonyms (Frost 2010). One author in particular, Ernst Ahl, contributed to this chaos by describing many new species (e.g. Ah] 1931a, b), most of which proved to be invalid (Laurent 1961, Frost 2010). As the in-depth stud- ies of Schiotz (1967, 1971, 1975) and others have shown, color and advertisement calls are the most reliable char- acters for identification of these species. Unfortunately, al- cohol preserved Hypervolius specimens quickly loose col- or (and do not call). Therefore it is often difficult, if not impossible, to evaluate the status of older museum vouch- ers. Reliable locality data may be of help in some cases where taxa show allopatric distributions and/or different habitat requirements. Bonn zoological Bulletin 57 (2): 177-188 One group of savanna dwelling reedfrogs proved to be es- pecially variable and consequently taxonomically compli- cated: the Hyperolius marmoratus/viridiflavus complex (Laurent 1951b, c, 1981; Schiotz 1971, 1999). These amazing reedfrogs have an outstanding natural history with annual population cycles and spectacular behavioral (Grafe et al. 2002), morphological and physiological adap- tations, and altogether a unique life history strategy to sur- vive the harsh and long dry seasons (Spieler 1997; Lin- senmair 1998; Lampert & Linsenmair 2002 and literature cited therein). So far, they are the only tetrapods where sex change has been documented (Grafe & Linsenmair 1989; for literary use of this knowledge see Crichton 1991). To date, Laurent (195la, 1976, 1983) and Schietz (1971) undertook the most detailed morphological ap- proach to disentangle the nomenclatory chaos of these widespread savanna dwelling frogs, which all share a sim- ilar morphology (short snout, very large vocal sac in males, transversal gular fold in females, extensive web- bing) and call (xylophone like metallic calls; for summa- ry see Schiovtz 1971, 1999). ©ZFMK 178 Mark-Oliver Rédel et al. However, the mentioned studies of these frogs, using col- oration and acoustics, did not provide much insight into their actual taxonomic status (see review by Wieczorek et al. 1998). Only more recently Wieczorek & Channing (1997) and Wieczorek et al. (2000, 2001) started to apply molecular techniques to disentangle the taxonomic chaos. In the course of their work in particular one member of the H. viridiflavus-complex/superspecies/species-group, Fig. 1. Bonn zoological Bulletin 57 (2): 177-188 H. nitidulus Peters, 1875, was acknowledged species sta- tus, a decision previously already applied for mostly prag- matic reasons by e.g. Schietz (1967), Drewes (1984) and Rédel (1996, 2000). This widespread West African savan- na frog was described by Peters (1875) from “Yoruba (La- gos)”, Nigeria. It was treated as a synonym of H. mar- moratus by Boulenger (1882), as a synonym of H. pic- turatus by Loveridge (1955) and as synonym or subspecies Life coloration of Hyperolius spatzi and H. nitidulus; upper left: calling H. spatzi male from Sabodala, Senegal, remark uniform yellow color at night; lower left: daytime coloration of H. spatzi from Sabodala, Senegal, with numerous minute black points; upper right: calling H. nitidulus male from Pendjari National Park, northern Benin, remark dark lateral band; lower right: H. nitidulus couple from Lamto reserve, Ivory Coast, remark almost uniform yellow color of male and grey mottling on legs and on the flanks in the female. ©ZFMK Taxonomy of West African Hyperolius 179 Table 1. Morphological differences between Hyperolius nitidulus and H. spatzi based on data provided in the original descrip- tions and comments based on type specimens and additional material examined herein. Comments which are already deducible from types only, are given in italic. Characters H. nitidulus H. spatzi Comments based on types and additional material Choanae large, not hidden small, hidden below similar sized and well visible in both species beneath edge of mandible edge of mandible Tongue large, broad and unusually small tongue in head width spatzi: 3.3 times; Snout (dorsal view) Snout (lateral view) Position of narins Position of heals heart-shaped roundish pointed flattened or roundish truncate slightly closer to snout-tip than to eye cover Or Surpass rounded truncated in mid distance between eye and snout-tip in contact when hind legs arranged to — each other body at right angles Dorsal skin skin smooth, laterally thick, smooth or with small warts almost leathery, rough, beset with nitidulus: 1.7 times truncated in dorsal and lateral view in juveniles, a bit more rounded in adults of both taxa truncated in dorsal and lateral view in juveniles, a bit more rounded in adults of both taxa in both species narins closer to snout-tip than to eye surpass each other in both taxa both taxa with rough skinned juveniles in dry season and smooth skinned adults in wet season many small smooth or rough warts Male gular flap absent Dorsal color yellow often with dark indistinct chalk white or fine present in both taxa H. spatzi with white, brown or yellow back, spots on back speckled with regularly beset with small black spots; dark-brown H. nitidulus never with such uniform pattern of black spots Pattern on flanks dark canthal and lateral No pattern in H. spatzi like on back; H. nitidulus with very stripe (continuous or distinct to rather indistinct black lateral band and broken), bordered white dark spots dorsally; below the stripe flanks marbled in dark grey and white Body-length 28 mm 21mm adults of both species up to about 30 mm of Hyperolius viridiflavus by many other authors (e.g. Lau- rent 195la, c, 1961; Schigtz 1971). The latter author al- so treated frogs described as Hyperolius spatzi Ahl, 1931 from Bakel-Kidira, Senegal (Ahl 1931a, b) as either be- longing to H. nitidulus (Schietz 1967) or as a “subspecies” of H. viridiflavus (Schietz 1971). In his book, Schietz (1999) used the name “spatzi” as a vernacular name, de- scribing “H. viridiflavus” populations of uncertain taxo- nomic status from Senegambia, whereas Rédel (2000) considered H. spatzi to represent a junior synonym of H. Bonn zoological Bulletin 57 (2): 177-188 nitidulus. However, already in the late seventies, Bohme (1978) revived the name H. spatzi for reedfrogs from Senegal, thus emphasizing their distinctiveness from oth- er West African savanna populations. Recently Emms et al. (2006) adopted this view and reported H. spatzi from Gambia. Our recent studies of many Hyperolius popula- tions at various West African savanna localities are the ba- sis of a taxonomic reinvestigation of both taxa presented herein. OZFMK 180 Mark-Oliver Rodel et al. Fig. 2. olius nitidulus (ZMB 7729, holotype, adult female) and right: H. spatzi (ZMB 32602, lectotype, subadult male). Dorsal and ventral views of the types of left: Hyper- MATERIAL & METHODS Morphological measurements were taken with a dial caliper (+ 0.1 mm) and are given in millimeters. Webbing formulae follow the scheme of Rédel (2000). Museum vouchers originated from the Staatliches Museum fir Naturkunde Stuttgart (SMNS) and the Museum fiir Naturkunde Berlin (ZMB; Appendix 2). Calls were record- ed with a Sony WM-D6C tape recorder and a direction- al microphone (Sony ECM-Z157 and Sony ECU-959C9) or an EDIROL R-09 24bit digital recorder (sample rate: 44.1 kHz, record mode: wav_24bit, microphone ECM- 950). These calls were analyzed with the program Avisoft SAS Lab Pro 4.5 (R. Specht, Berlin, Germany). For se- quence comparisons, we analyzed 247 base pairs (bp) of the mitochondrial 16S ribosomal RNA gene from Hyper- olius spatzi (ZMB 74280, GenBank HQ113098; Senegal, Sabodala) and Hyperolius nitidulus (ZMB 74884, Gen- Bonn zoological Bulletin 57 (2): 177-188 Bank HQ113099, Sierra Leone, Tingi Hills; no voucher, GenBank HQ113100, Ivory Coast, Mont Sangbé Nation- al Park, tissue without voucher). Further hyperoliid gene sequences were obtained from GenBank (Tab. 1). DNA extraction, amplification and sequence alignment followed the procedures as described in Rédel et al. (2009). Uncor- rected pairwise sequence divergence was calculated us- ing PAUP*4b10 (Swofford 2002). RESULTS & DISCUSSION Morphological comparison. A major problem in using external morphological characters for determination of these frogs is their polymorphism. Schigtz’s (1963, 1967, 1971) described distinct color phases for many Hyperolius species, i.e. called F or A and J or B, respectively. The phase F/A of H. nitidulus/spatzi refers to the adult/wet sea- son pattern, whereas phase J/B is the juvenile or sub-adult dry season pattern. Young frogs in dry season condition have a rough, warty dorsal skin which is brown below 35°C and chalk white above this temperature (see figs. in Spieler 1997 and Rédel 2000). Adult frogs have smooth skin and a completely different dorsal color pattern (Fig. 1). These morphological differences are part of the amaz- ing aestivation strategy of these frogs (see Linsenmair 1998; Rédel 2000 and literature cited therein). According to the descriptions by Ahl (1931b) differences between Hyperolius nitidulus and H. spatzi would be those summarized in Tab. 1 (compare also translations of the original descriptions provided in Appendix 1). Major dif- ferences between the descriptions of H. nitidulus and H. spatzi consist in the fact that the description of the for- mer is based on an adult female, whereas the description of the latter is based on a series of subadult frogs in dry season condition (Fig. 2). Schiotz (1967, 1971, 1999) mentioned differences be- tween various West African H. nitidulus populations, in- cluding a cline in pattern from Sierra Leone (few and small spots on flanks) to Cameroon (broad lateral band; same cline in pattern on the lower legs). He also observed an hour-glass pattern and a dark vocal sac in frogs from Sier- ra Leone (likewise present in some juveniles in northern Ivory Coast, see Rédel 2000). Our specimens from Sier- ra Leone neither differed in coloration nor in genetics (see below) from e.g. H. nitidulus populations from northern Ivory Coast. Schietz (1971) further mentions that frogs from drier northern savannas are more uniform grayish colored, whereas more southern ones, 1.e. from the humid savanna types, exhibit a distinct pattern. The latter differ- ences might be related to age. R6del (2000) reported that older specimens are more distinctly colored. As adult H. nitidulus are unable to survive the dry season, all popu- ©ZFMK Taxonomy of West African Hyperolius 181 H. spaizi 10 N aE = > oO = o =) oO 2 cs 1V H._ nitidulus Frequency (kHz) 1V Fig. 3. Waveforms, spectrograms and energy plots of the advertisement calls of Hyperolius spatzi (above) and H. nitidulus (be- low; compare Tab. 2). The Hyperolius spatzi male from Sabodala, Senegal, was recorded in a terrarium. The Hyperolius nitidulus was recorded at a savanna pond in Comoé National Park, Ivory Coast. The background noise is a chorus of other H. nitidulus males. lations are annual (Linsenmair 1998). In more humid sa- vannas, the wet season lasts longer and frogs may reach older ages (and thus potentially a more colorful pattern). Almost all anatomical differences (position and size of choanae, position of narins, size, shape, length of extrem- ities, head width) deducible from Ahl’s (193 1a, b) descrip- tions (compare Tab. 1) are identical among both taxa (for specimens investigated see Appendix 2). Both species have very short, rounded snouts, females posses a typical gular fold and males have very large vocal sacs with a large but diffuse whitish yellow gular flap (gland). Juve- niles are often almost indistinguishable. Hyperolius nitidu- lus juveniles show clear dorsolateral bands or an hour- glass pattern shortly after metamorphoses (see figs. in Rédel 2000). In dry season conditions they are uniform Bonn zoological Bulletin 57 (2): 177-188 brown or white. Juvenile H. spatzi in dry season condi- tions are white with many small black dots, the latter sometimes being indistinct. In contrast, adult frogs are dis- tinctively colored. The dorsal surfaces of H. spatzi are chalk-white to yellow, densely beset with tiny black spots, whereas H. nitidulus is brownish or yellowish with black spots and has black lateral lines and spots (plate 18 in Leaché et al. 2006). H. nitidulus has white, yellow or red- dish ventral surfaces, whereas these surfaces are exclusive- ly yellow in H. spatzi (see fig. 2f in Emms et al. 2006). The hidden parts of legs are pinkish to blood red in both species (Fig. 2 and figs. 430 & 431 in Schiotz 1999, figs. in Rédel 2000). Generally, females of H. nitidulus have amore distinct lateral black pattern than males, which can be almost uniform brown (Fig. | and figs. in R6del 2000). At night, males of both taxa appear uniform yellowish. ©ZFMK 182 Mark-Oliver Rodel et al. Table 2. Characteristics of the advertisement calls of Hyperolius spatzi, recorded in Sabodala, Senegal, and H. nitidulus, record- ed in the Comoé National Park, Ivory Coast and Mount Nimba, Guinea (Fig. 3). Differences of call length, main frequency and time between calls have been tested by comparing mean values of five males of each species (Wilcoxon test). Call length [sec] Frequency [Hz] Inter-call intervals [sec] mean 0.08 sd 0.04 H. spatzi N (males) 5 N (calls) 25 mean 0.02 sd < 0.01 H. nitidulus N (males) 5 N (calls) 25 W 616 P < 0.0001 2638.0 1 139.6 0.80 5 5 25 25 2927.6 1.01 85.1 0.29 >) 4 25 20 26 218 < 0.0001 0.4756 Usually, the pattern in H. nitidulus remains vaguely vis- ible. The only morphological difference detected by us (herein confirming Ahl 1931a, b), is the size and shape of the tongue. Hyperolius spatzi usually have comparative- ly smaller and narrower tongues than H. nitidulus, whose tongue 1s broad and almost heart-shaped. This 1s also vis- ible in the type specimens of both species. Acoustics. The advertisement call of both taxa is a sin- gle, pure, metallic and very loud tone (Fig. 3). Choruses of both species resemble xylophones or bells. Although superficially similar, advertisement calls of both taxa showed significant differences. The call of H. spatzi was of comparatively longer duration and lower frequency (Tab. 2). The small sample size and the relatively slight differences in call characteristics urge for some caution in their interpretation. However, the acoustic results are not contradicting the specific distinctiveness of H. nitidu- lus and H. spatzi. Genetics. The genetic distances in the investigated frag- ment of the 16S RNA gene between Hyperolius spatzi (N= 1) and 1. nitidulus (N= 3, originating from Sierra Leone and Ivory Coast) ranged from 5.1—5.6%. The mean dis- tance between H. spatzi and various other members of the H. viridiflavus/marmoratus-complex (N= 33; including H. nitidulus) was 5.9% (+ 1.1 SD; range: 3.6-8.7%). The low- est distance was present in comparison to a H. viridiflavus angolensis, the highest to a H. viridiflavus viridiflavus sample (sequences from GenBank, compare Tab. 3). Mean Bonn zoological Bulletin 57 (2): 177-188 genetic distances between H. spatzi and nine other Hyper- olius species was 18.7% (+ 3.9 sd; range: 11.2—23.2%). The lowest distance present occurred in comparison to H. Jusciventris, the highest to a H. cinnamomeoventris sam- ple (compare Tab. 3). Based on genetic data (12S and 16S), Wieczorek et al. (2000, 2001) recognized H. nitidulus as being distinct on the species level from other members of the H. viridiflavus group. Altogether they accepted ten species within this group of which H. nitidulus was most distinct (within in- traspecific genetic variation 0.7—-4.8%; between clade vari- ation 2.4-10.0%; Wieczorek et al. 2001). Our data con- firm their results and speak in favor of likewise recogniz- ing H. spatzi as a distinct species. Distribution. Hyperolius nitidulus occurs in humid to dry savannas of West Africa (Fig. 4; Lamotte 1966; Schietz 1967, 1999; Rédel 2000). Laurent’s (1951c) doubts con- cerning the type locality of H. nitidulus were rejected by Schiotz (1963), by explaining that savanna exists at the type locality, and thus also suitable habitats for H. nitidu- lus. Records have been published for Benin (Nago et al. 2006), Ghana (Schiotz 1964a, 1967; Hoogmoed 1980; Hughes 1988; Rédel & Agyei 2003; Leaché 2005; Leaché et al. 2006), Burkina Faso (this paper), eastern and cen- tral Guinea (Laurent 1951a, c; Schiotz 1967; Rédel et al. 2004; Hillers et al. 2006, 2008; Greenbaum & Carr 2005), Ivory Coast (Laurent 1951c; Lamotte & Perret 1963; Bar- bault 1967, 1972; Lamotte 1967; Schigtz 1967; Vuattoux ©OZFMK Taxonomy of West African Hyperolius 183 Fig. 4. : 1 Kilometers Known distributions of Hyperolius spatzi (circles) and H. nitidulus (squares) based on museum and literature records (compare text and Appendix 2); stars indicate positions of type localities of H. spatzi (Senegal) and H. nitidulus (Nigeria). The north-westernmost record of H. nitidulus in Nigeria may refer to H. pallidus, southern and central Cameroonian populations are usually referred to two H. nitidulus subspecies (compare text and fig. 428 in Schiotz 1999). 1968; Euzet et al. 1969; Rédel 1996, 1998, 2000, 2003; Spieler 1997; Linsenmair 1998; Rédel & Spieler 2000; Rodel & Ernst 2003; Adeba et al. 2010), Mali (Schiotz 1967), Nigeria (Schiotz 1963, 1966, 1967; Walker 1968; Onadeko & Rédel 2009), Sierra Leone (Schiotz 1964b, 1967; Lamotte 1971), and Togo (Bourgat 1979; Segniag- beto et al. 2007). Hyperolius spatzi, as defined herein, has been recorded from Senegal (Boettger 1881, as H. cinctiventris; Loveridge 1956; Schietz 1967; Lamotte 1969; Miles et al. 1978, listed as H. nitidulus; Ahl 1931a, b; B6hme 1978), and The Gambia (Andersson 1937 as H. sp., but unam- biguous description provided; Barnett & Emms 2005 as H. nitidulus; Emms et al. 2006). A record from Guinea was actually based on 7. nitidulus (Hillers et al. 2006; see Ap- pendix 2). Schietz (1971) recognized “H. viridiflavus spatzi’’ as a taxonomic unit occurring in Senegambia and provides a map, indicating the distribution of H. spatzi and H. nitidulus, respectively (fig. 42 in Schietz 1971). Padi- al & de la Riva (2004) believed that H. nitidulus and H. viridiflavus may occur in southern Mauritania. Hyperolius viridiflavus (sensu stricto) certainly does not occur in western Africa, including Mauritania. Hyperolius nitidu- lus might reach eastern Mauritania and it seems very like- ly that H. spatzi might be a part of the Mauritanian fau- na, as is indicated by the close proximity of the type lo- cality of this species to the boarder of Mauritania (Fig. 4). Mountains and rivers can act as potential barriers between taxa (e.g. Li et al. 2009, for contrasting results see Gas- con et al. 1998). In this case, the Géba and Corubal rivers Bonn zoological Bulletin 57 (2): 177-188 along the border between Guinea-Bissau and Guinea, might fulfill such a role. It is also possible that the north- ern foothills of the Fouta Djallon serve as an altitudinal barrier. However, more data from Equatorial Guinea, west- ernmost Guinea, eastern Senegal, western Mali and Mau- ritania would be needed to clarify the exact limits of the species’ ranges. The distribution of H. nitidulus in Central Africa is more complicated. The species is listed as 1. viridiflavus (sub- species H. v. nitidulus, H. v. pallidus) for Cameroon, the Central African Republic and the Democratic Republic of Congo by Frétey & Blanc (2000). In northern Cameroon and adjacent north-eastern Nigeria, Chad and the Central African Republic (Joger 1990), H. nitidulus may be re- placed by H. pallidus which was described by Mertens (1940) from dry northern Cameroon (Poli near Garua) and which has been treated by Perret (1966) as a full species, and by Schietz (1971) and Amiet (1973) as a subspecies of H. nitidulus. From Cameroonian savannas, situated a bit further south, two H. nitidulus subspecies have been described by Perret (1966). Hyperolius n. bangwae occurs in elevated savannas, 1.e. Bamenda, Bamiléké, Adamaoua, whereas H. n. aureus is said to occur in the drier north- ern savannas and semi-deserts (Perret 1966; compare e.g. Bohme & Schneider 1987 for some records). This view was adopted by Schietz (1971) and Amiet (1973). The lat- ter provided arguments for the treatment of these taxa as subspecies of H. nitidulus, i.e. Cameroonian frogs differ from typical H. nitidulus by slightly smaller size and slightly duller coloration. The voices are “as good as iden- tical” (Amiet 1973). More recently, Amiet thought that all ©OZFMK 184 Mark-Oliver Rodel et al. Table 3. Genetic distances between Hyperolius spatzi (ZMB 74280; GenBank #: HQ113098) and other Hyperolius species. Un- corrected p-distances are based on 247 base pairs of mitochondrial 16S ribosomal RNA. Values for H. nitidulus are given in bold. Genus Species subspecies“ GenBank # p-distance Hyperolius chlorosteus FJ594076 0.214 Hyperolius cinnamomeoventris FJ594077 0.232 Hyperolius concolor FJ594078 0.203 Hyperolius fusciventris FJ594080 0.112 Hyperolius guttulatus FJ594082 0.133 Hyperolius horstocki AF282410 0.199 Hyperolius kivuensis AF282409 0.183 Hyperolius naustus AF215442 0.219 Hyperolius nitidulus HQ113099 0.051 Hyperolius nitidulus HQ113100 0.051 Hyperolius nitidulus AF282435 0.056 Hyperolius picturatus FJ594090 0.186 Hyperolius viridiflavus AF215440 0.056 Hyperolius viridiflavus AF215441 0.061 Hyperolius viridiflavus AY323920 0.077 Hyperolius viridiflavus angolensis AF282411 0.036 Hyperolius viridiflavus albofaciatus AF282433 0.065 Hyperolius viridiflavus aposematicus AF282412 0.051 Hyperolius viridiflavus argentovittis AF282431 0.046 Hyperolius viridiflavus bayoni AF282413 0.082 Hyperolius viridiflavus broadleyi AF282414 0.071 Hyperolius viridiflavus ferniquei AF282416 0.051 Hyperolius viridiflavus ferniquei AY 603987 0.051 Hyperolius viridiflavus glandicolor AF282417 0.066 Hyperolius viridiflavus goetzi AF282418 0.066 Hyperolius viridiflavus mariae AF282419 0.066 Hyperolius viridiflavus mariae AF282420 0.066 Hyperolius viridiflavus marginatus AF282430 0.051 Hyperolius viridiflavus melanoleucus AF282432 0.056 Hyperolius viridiflavus pamtherinus AF282425 0.051 Hyperolius viridiflavus pitmani AF282426 0.066 Hyperolius viridiflavus marmoratus AF282421 0.056 Hyperolius viridiflavus ngorongoro AF282423 0.066 Hyperolius viridiflavus ommatostictus AF282424 0.056 Hyperolius viridiflavus pyrrhodictyon AF282434 0.046 Hyperolius viridiflavus rhodesianus AF282427 0.038 Hyperolius viridiflavus rubripes AF282436 0.062 FHyperolius viridiflavus swymmertoni AF282415 0.071 Hyperolius viridiflavus taeniatus AF282422 0.056 Hyperolius viridiflavus verrucosus AF282428 0.062 Hyperolius viridiflavus viridiflavus AF282429 0.087 Bonn zoological Bulletin 57 (2): 177-188 ©OZFMK Taxonomy of West African Hyperolius 185 three Cameroonian taxa are subspecies of H. viridiflavus, i.e. the highlands of western Cameroon and the Adamaoua plateau being inhabited by H. v. aureus (and possibly H. v. bangwae), and populations occurring in northern Cameroon (mid-Sudanian, Sudano-Sahelian and Sahelian zones) belong to H. v. bangwae and H. v. pallidus (J.-L. Amiet pers. comm.). Conclusions. Our investigations on the type specimens, as well as on additional vouchers, revealed small but dis- tinct morphological (mostly color pattern; tongue shape and size), significant acoustic and large genetic differences (16S gene). Especially the genetic differences are clear- ly within the range that is thought to be species specific in anurans (Vences et al. 2005a, b; Rédel et al. 2009; Vieites et al. 2009). Our results thus speak in favor of rec- ognizing both taxa as distinct species. A contradicting ar- gument was seen in the very complicated situation of a large variation of color patterns between and within pop- ulations of the Hyperolius viridiflavus/marmoratus species group(s). Schietz (1999) thus questions an approach where the taxonomy for only a small part of the continent would be resolved. However, in West Africa it is possible to as- sign these frogs to particular names and we thus do not see a reason for avoiding it. We therefore herein resurrect the species status of H. spatzi, designate a lectotype from the series of syntypes and redescribe the species based on type and new material. REDESCRIPTION OF HYPEROLIUS SPATZI AHL, 1931. ZMB 32602 (lectotype; Fig. 2), 74853-74876 (paralecto- types, formerly all ZMB 32602), all from Bakel-Kidira, Senegal, coll. Spatz. Description of lectotype (all measurements in mm). Subadult frog (male, vocal sac barely developed?); short, compact body; snout-vent length 19.2; head width 7.3, head length 6.9, thus head wider than long; snout short and truncated in dorsal and lateral view; narins angular narrow slit, closer to snout-tip than to eye; tympanum hid- den; transversal gular fold; tongue small, narrow, almost parallel and notched anteriorly, tongue width 2.3, tongue length 3.2, tongue 3.3 times in head width; choanae small and round, close to edge of mandible but well visible; dor- sal skin slightly granular; belly granular (medially dissect- ed); ventral skin on thighs near vent granular, remaining ventral parts of hind limbs smooth; finger and toe tips en- larged to discs; relative lengths of fingers: 1<2<4<3; basal webbing between fingers; femur length: 8.4; tibia length: 10.4; foot incl. longest toe: 14.0; relative lengths of toes: 1<2<3<5<4; webbing formula: 1 (0), 2 (1.5-0), 3 (1.5—0.5), 4 (1-0), 5 (0); subarticular tubercles on fingers Bonn zoological Bulletin 57 (2): 177-188 and toes not very prominent. Dorsal surfaces chalk white, densely beset with minute black points; ventral skin on thighs near vent white, remaining parts of thighs and ven- tral parts of shanks, feet, inner parts of forelimbs, ventral part of hands and fingers fleshy colored. Variation. Series of paralectotypes almost indistinguish- able from lectotype, exclusively subadult frogs in dry sea- son conditions; dorsal skin partly more or less granular than in lectotype; black points on white ground sometimes more distinct or sometimes almost absent. Further mate- rial (see Appendix 2) exhibit the following variation: Male snout-vent length: 27.0—-31.3 (N= 6); female snout-vent length 30.6 (N= 1); snout shape of adult frogs in dorsal and ventral view slightly more rounded than in juveniles; adults of both sexes in ethanol with dorsal surfaces (incl. upper side of thighs) with brownish ground color (com- posed of small, very dense brownish points), many very distinct black spots; black spots sometimes a bit more abundant on flanks than on back; some specimens with black spots on throat; others with uniform clear ventral surfaces; tongue in almost all specimens small and com- paratively narrow (exception: ZMB 74279). Adult animals in life brownish to yellow with very distinct black spots. These may be not visible during night. Venter yellow. Acknowledgements. 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Videnskabelige Meddelelser fra Dansk Naturhistorisk Forening 125: 1-92 + 4 plates Schietz A (1964a) A preliminary list of amphibians collected in Ghana. Videnskabelige Meddelelser fra Dansk Naturhistorisk Forening 127: 1-17 Schigtz A (1964b) A preliminary list of amphibians collected in Sierra Leone. Videnskabelige Meddelelser fra Dansk Naturhis- torisk Forening 127: 19-33 + 1 plate Bonn zoological Bulletin 57 (2): 177-188 Schiotz A (1966) On a collection of amphibia from Nigeria. Vi- denskabelige Meddelelser fra Dansk Naturhistorisk Forening 129: 43-48 + 3 plates Schietz A (1967) The treefrogs (Rhacophoridae) of West A frica. Spolia Zoologica Musei Hauniensis 25: 1-346 Schiotz A (1971) The superspecies Hyperolius viridiflavus (Anu- ra). Videnskabelige Meddelelser fra Dansk Naturhistorisk Forening 134: 21-76 Schiotz A (1975) The treefrogs of Eastern Africa. Steenstrupia, Copenhagen Schigtz A (1999) Treefrogs of Africa. 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Philosophical Transactions of the Royal So- ciety B Biological Sciences 360: 1859-1868 Vences M, Thomas M, van der Meijden A, Chiari Y, Vieites DR (2005b) Comparative performance of the 16S rRNA gene in DNA barcoding of amphibians. Frontiers in Zoology 2: 5 Vieites DR, Wollenberg KC, Andreone F, Kohler J, Glaw F, Vences M (2009) Vast underestimation of Madagascar’s bio- diversity evidenced by an integrative amphibian inventory. Proceedings of the National Academy of Sciences of the U.S.A. 106: 8267-8272. Vuattoux R (1968) Le peuplement du palmier ronier (Borassus aethiopum) d'une savane de Cote dIvoire. Annales de Il’ Uni- versité d’ Abidjan, Sér. E, 1: 1-138 Walker RB (1968) The amphibians of Zaria, in the northern Guinea savannah, Nigeria. Copeia 1968: 164-167 Wieczorek AM, Channing A (1997) The taxonomic status of Broadley’s reed frog. African Journal of Herpetology 46: 110-116. Wieczorek AM, Channing A, Drewes RC (1998) A review of the taxonomy of the Hyperolius viridiflavus complex. Herpeto- logical Journal 8: 29-34 Wieczorek AM, Drewes RC, Channing A (2000): Biogeography and evolutionary history of Hyperolius species: application of molecular phylogeny. Journal of Biogeography 27: 1231—1243 Wieczorek AM, Drewes RC, Channing A (2001) Phylogenetic relationships within the Hyperolius viridiflavus complex (Anu- ra: Hyperoliidae), and comments on taxonomic status. Am- phibia-Reptilia 22: 155-166 ©ZFMK 188 Mark-Oliver R6édel et al. APPENDIX 1. Translations of the original descriptions of Hyperolius nitidulus by Peters (1875) and Hyperolius spatzi by Ahl (193 1a): Hyperolius nitidulus (Fig. 2): “Body shape equals that of H. marmoratus. Snout same length as eye. Tympanum hidden. Bel- ly and ventral surfaces of thighs granular. The outer two fingers and the forth toe, with the exception of the two most distal pha- langes, webbed. Dorsally purple grey, shanks likewise colored, whereas the thighs seem to be uncolored. A black band from nose through eye to belly, there band dissolving into black spots on white background; upper lip, flanks below this band, anal region, upper surfaces of forearms, outer and inner edge of shanks and external side of foot to toe tips (in ethanol) white with black spots, which plus/minus fuse. Total length 28 mm; head: 8 mm; head width: 8.5 mm; forearm: 19 mm; hand with 3" finger: 7 mm; hind leg: 44 mm; foot with fifth toe: 20 mm. From Yoru- ba (Lagos). [comment added: referring to ZMB 7729, holotype]” Plate 3 (figures 4 and 4a) in Peters (1875) figures the typical wet- season color pattern of this species. Hyperolius spatzi (Fig. 1): “stocky body shape; vomerine teeth absent; choanae very small, hidden below edge of mandible; tongue unusually small, notched posteriorly; large head, app. 1/3 of body length, wider than long; snout rounded, truncated in lat- eral view, not or only slightly surpassing mouth, as long as eye, much shorter than distance between anterior corner of eyes, slightly longer than high; canthus rostralis rounded but distinct; loreal region vertical, only slightly concave; narines in mid dis- tance between eye and snout-tip; inter-narial distance slightly narrower than inter-orbital distance, the latter twice as wide as upper eyelid; tympanum hidden beneath skin. Robust fingers, 1/3 to 1/2 webbed; well developed discs; 15* fin- ger shorter than second, second shorter than fourth, which is slightly surpassed by the 3"4 finger; 3" finger as long as snout; subarticular tubercles moderately large, not prominent. Webbing on feet complete with the exception of 4" toe where the last pha- lanx is without webbing; discs as large as those on fingers; 5‘ toe slightly longer than 3'4; external metatarsalia tightly fused, tarsal fold absent; very small inner metatarsal tubercle; outer metatarsal tubercle lacking; no tarsal tubercle; subarticular tu- bercle small, moderately distinct. Tibio-tarsal angle surpasses eye or reaches snout-tip. Femur shorter than tibia, the latter 3.5—4 times longer than wide and twice or slightly less times in body length, longer than foot; heals in contact when hind legs arranged to body at right angles. Dorsal skin thick, almost leathery, rough, beset with many small smooth or rough warts; ventrum granular; distinct postgular and postpectoral folds; no temporotemporal fold; males with subgu- lar vocal sac and a small, indistinct gular flap. Coloration in alcohol dorsally chalk white or, rarer, fine speck- led with dark-brown. Venter white. Ventral parts of thighs and inner parts of shanks flesh-colored (presumably red in life). No markings at all. Body length 21 mm. Bakel-Kidira (Upper Senegal region). 26 specimens, Bakel-Kidira, Spatz leg., types [comment added: ZMB 32602, lectotype; 74853-74876, paralectotypes; formerly all ZMB 32602]. The species is named to honor the collector, the well know researcher Spatz, whose collecting activities re- sulted in a large number of valuable reptiles and amphibians, stored in the Berlin museum.” Bonn zoological Bulletin 57 (2): 177-188 APPENDIX 2. Voucher specimens, including types, of Hyperolius spatzi and H. nitidulus in the ZMB and SMNS collections. Hyperolius nitidulus. Benin: ZMB 74896-74898, Pendjari Na- tional Park, Sudan savanna, October 2003, coll. Olaf Grell; ZMB 74890, Pendjari National Park, Tangieta, savanna, N 10°38.317’, E 01°15.746’, 1 September 2004, coll. G.A. Nago & M.-O. Rodel; Burkina Faso: ZMB 74893-74894, Dano, small river in savanna, N 11°14’16.8”, W 03°01’24.1”, 22 October 2003, coll. T. Moritz; Ivory Coast: SMNS 8995.1-2, Ananda, 1993, coll. M.-O. Rédel; SMNS 9680.1-2, Bondoukou, 1996, coll. K. Koua- dio; ZMB 74888 & SMNS 8967.1-7, Comoé National Park, sa- vanna, June 1996, coll. M.-O. Rédel; ZMB 74886, Mont Sang- bé National Park, Mare Soumarou, island forest in the savanna, pitfall trap, dry season 2001, coll. G. Gbmalin & Y. Cesar; Guinea: ZMB 74895, Mont Béro Classified Forest, savanna, N 08°08730.9”, W 08°34’09.6”, 1 December 2003, coll. M.A. Ban- goura & M.-O. Rédel; ZMB 74891-74892 Nimba Mountains, savanna Séringbara, with big ponds, close to village, N 07°36.181’, W 08°29.769’, 18 May 2006, coll. T.N.-S. Loua & A. Hillers; ZMB 74889, Pic de Fon/Simandou range, Banko, sa- vanna, 11 July 2004, coll. M.A. Bangoura & K. Kamara; ZMB 74882, Boké Préfécture/Kolaboui, swampy area in secondary forest island, N 10°45.075’, W 14°27.040’, 23 & 24 April 2005, coll. M.A. Bangoura & A. Hillers (originally listed as H. spatzi in Hillers et al. 2006); Nigeria: ZMB 7729 (holotype), Yoruba (Lagos), coll. Krause; Sierra Leone: ZMB 74884-74885, Tin- gi Hills, big pond with a few trees around and swampy area in savanna, N 08°51.047’, W 10°46.502’, 427 ma.s.l., 5 June 2007, coll. J. Johnny & A. Hillers; Togo: ZMB 39028, station Sokode, coll. Schréder. Hyperolius spatzi. Gambia: ZMB 74877, Abuko Nature Re- serve, savanna, 2005, coll. L. Barnett & C. Emms; Senegal: ZMB 32602 (lectotype), 74853-74876 (paralectotypes, former- ly all ZMB 32602), Bakel-Kidira, coll. Spatz; ZMB 74279, Sa- bodala, ponds and puddles in degraded farmbush savanna next to Oromin camp, N 13°09.368’, W 12°06.882’, 12 September 2009, coll. A. Hillers & Y. Mané; ZMB 74280-74285, Sabodala, in and around big pond in farmbush savanna/grassland, with some rocks, N 13°07.259’, W 12°07.622’, 7 September 2009, coll. A. Hillers & Y. Mané. Received: 26.VII.2010 Accepted: 24. VIII.2010 ©OZFMK Bonn zoological Bulletin Volume 57 Issue 2 pp. 189-196 Bonn, November 2010 Genetic variability in mainland and insular populations of Podarcis muralis (Reptilia: Lacertidae) Massimo Capula! & Claudia Corti? 'Museo Civico di Zoologia, Via U. Aldrovandi, 18, I-00197 Roma, Italy; E-Mail: massimo.capula@comune.roma.it 2Museo di Storia Naturale dell’ Universita di Firenze, Sezione di Zoologia “La Specola”’, Via Romana, 1-50125 Firenze, Italy, E-Mail: claudia.corti@unif1.it Abstract. Allozyme electrophoresis was used to study the distribution of genetic variation within and among mainland and insular populations of the lacertid lizard Podarcis muralis from western, southern and eastern Europe. Genetic vari- ability in the species is low and genetic subdivision is high. The highest values of percent polymorphism and heterozy- gosity were found in the samples from two Tyrrhenian islands (Elba Island, La Scola Islet). The occurrence of higher levels of genetic variability in insular populations is probably because these populations inhabit marginal environments characterized by temporal-ecological instability. In these environments high heterozygosity levels can be preserved af- ter colonization events, unless founder populations are so small that bottleneck effects occur. The genetic heterogeneity analysis demonstrates a certain amount of genetic differentiation among local populations of P. muralis, with a relative- ly high level of genetic subdivision. Allozyme data show that genetic variation in P. muralis is distributed into two ma- jor population groups: the first includes the closely related samples from Spain and SW France, the second the geneti- cally recognizable samples from Germany, Italy, and Greece. The average genetic distance between the two groups is relatively high (Ne1’s D = 0.059), with D ranging from 0.043 to 0.100. Key words. Podarcis muralis, Lacertidae, allozyme electrophoresis, population heterogeneity, Tyrrhenian islands, Eu- rope. INTRODUCTION There have been numerous surveys of the genetic struc- ture of insular populations of vertebrates, especially rep- tiles (e.g. Soulé & Yang 1974; Gorman et al. 1975; Pat- ton et al. 1975). From these studies it became evident that many demographic, historical, and geographic factors in- fluence the pattern of genetic variation in the insular pop- ulations (e.g. Soulé et al. 1973; Soulé 1976). The Mediterranean lacertid lizards of the genus Podarcis seem to be particularly useful for this type of investiga- tion because they are widespread on several Mediterranean islands and are normally characterized by high inter- and intra-population morphological and genetic variability (e.g. Harris & Arnold 1999; Arnold & Ovenden 2002; Cor- ti & Lo Cascio 2002; Salvi et al. 2009). Although the evo- lutionary significance of the pattern of variation observed in these lacertid lizards has been unstudied for most taxa, in some cases at least it was pointed out that species which are characterized by a high degree of phenotypic plastic- ity in the pattern of the upper parts may have levels of ge- netic variability higher than those found in the morpho- Bonn zoological Bulletin 57 (2): 189-196 logically low variable species (see e.g. Selander 1976; Ca- pula 1994a, 1996, 1997; Losos et al. 1997; Capula & Cec- carelli 2003; Caputo et al. 2008). In this paper, based primarily on allozyme data, the dis- tribution of genetic variation within and among mainland and insular populations of the lacertid lizard Podarcis mu- ralis from western, southern and eastern parts of its Eu- ropean range was estimated. Podarcis muralis was cho- sen as it is a morphologically and ecologically variable species occurring in a wide variety of habitats over its dis- tribution range, which extends from the northern border of the Iberian Peninsula to north-western Turkey, and throughout central and southern Europe (Arnold & Ovenden 2002; Corti & Lo Cascio 2002). In the northern part of its range this lizard is typically a thermophilous and lowland species, with a reduced variability in the pat- tern of the upper parts, while in the southern part it is more often a mountain species, occurs especially in wet and shady habitats, and is characterized here by high pheno- typic variability (see Capula et al. 1993, 2009; Corti et al. ©ZFMK 190 Massimo Capula & Claudia Corti Table 1. Geographic and collecting data for the Podarcis muralis samples used in this study. Population Locality Sample size Guadarrama (Spain) Anso (Spain) Ordesa (Spain) Deba (Spain) Albaran (Spain) Bidache (SW France) Le Chiroulet (SW France) St. Gaudens (SW France) Bonn (Germany) Cavalese (Italy) Cesena (Italy) Resceto (Italy) Chiusdino (Italy) Populonia (Italy) Uccellina Mountains (Italy) Ostia (Italy) NS SY J 70%) and different from nearly all the other samples in ANOSIM. Guadarrama is a recently differentiated but well diagnos- able (morpho)subspecies (with lower values of dorsalia, ventralia and greater values of circumanalia). Despite the mtD- NA differences of the Béjar specimens, their morphology is largely equivalent to that of 1. cyreni castiliana (Gredos), but clearly differ in their female body elongation (near 1 cm) with shorter limbs, a possible strategy to increase clutch size. Populations from the Sierras de Avila (Villafranca, Serrota and Paramera) are very similar among them. Villafran- ca (in males) together with Beéjar (in females) are the most connected samples in MST, and the root of the species dif- ferentiation from a morphological point of view, once discarded geographical and climatic influence on morphology. All populations except Guadarrama shall be considered as J. c. castiliana by their morphological identity with Gredos. These morphological similarities probably are the reflect of extensive gene flow among them, responsible of maintaining their morphology largely equivalent. Key words. Lacertidae, /berolacerta cyreni, Intraspecific variability, subspecies, Geographical variation, Iberian Penin- sula. Resumen. Se ha calculado un Analisis Discriminante Canonico, ANOVA y ANOSIM con toda el area de distribucion de Iberolacerta cyreni, incluyendo varias poblaciones periféricas no estudiadas hasta la fecha. La unica muestra diferencia- da es Guadarrama (la subespecie nominal), con muy poco solapamiento en el CDA (clasificacion correcta > 70%) que difiere de practicamente todas las demas muestras en el ANOSIM. Guadarrama es una poblacion recientemente diferen- ciada, pero bien diagnosticable como (morfo)subespecie (valores bajos de dorsalia y ventralia, y altos de circumanalia). A pesar de las diferencias mitocondriales de Béjar, su morfologia es ampliamente asimilable a J. c. castiliana (Gredos), siendo destacable el relativo elongamiento corporal de las hembras (casi | cm) con miembros proporcionalmente cortos, una posible estrategia para incrementar el tamafio de puesta. Las poblaciones de las Sierras de Avila (Villafranca, Serro- ta y Paramera) son muy similares entre si. Villafranca junto con Béjar (en machos y hembras respectivamente) estan mor- fologicamente en la raiz de la diferenciacion de la especie (MST), una vez descartada cualquier influencia climatica o de distancia geografica. Excepto Guadarrama, todas deben considerarse como J. c. castiliana por su identidad morfologica con Gredos, lo que refleja la probable presencia de un flujo genético extensivo y reciente entre ellas. INTRODUCTION The Spanish Sistema Central consists of a series of Sier- ras, more or less aligned in a ENE-WSW direction, which separate the Duero (to the North) and Tajo (to the South) river drainages, or what is the same, the Old and New Castile plateauxes. It runs from the Portuguese Serra da Estrela (inhabited by a relict population of /berolacerta monticola |Boulenger, 1905]), across the Spanish Sierra de Gata (apparently too low and dry for /berolacerta), the Sierra de Francia with /berolacerta martinezricai (Arribas, 1996) and the main part of the Spanish portion of its range Bonn zoological Bulletin 57 (2): 197-210 (over 240 km in length), which is inhabited in several points by populations of the Carpetane Lizard (/berolac- erta cyreni [Miller & Hellmich, 1937]). The Carpetane Lizard is widespread through the main parts of the Sistema Central and is mainly known from Sierras de Béjar, Gredos and Guadarrama (Fig. 1). It was raised to species level (Arribas 1996) based on allozymes (Mayer & Arribas 1996), karyology (heterochromatiniza- tion of sex-chromosome and localization of the NORs; OZFMK 198 Oscar J. Arribas Odierna et al. 1996) and adult and hatchlings pattern and coloration. Two subspecies were defined, the nominal /. c. cyreni from Guadarrama (type locality: Puerto de Navacerrada), and /. cyreni castiliana from Gredos (type locality: Circo de Gredos, Avila) to which frequently are assimilated Béjar specimens. This latter subspecies differs from the nominate one by a reduced dark pattern, more dorsalia, ventralia, slightly larger diameter of the masse- teric and hindlimb length, and lower circumanalia (Arribas 1996). The degree of genetic differentiation between /. c. cyreni (Guadarrama) and /. c. castiliana (Gredos) was analyzed by Mayer & Arribas (2003), who found a mtDNA se- quence divergence of 0.6 % in the 12s rRNA (12s) and 16s rRNA (16s) mitochondrial genes, which corresponds to approximately 0.6 MY BP. Carranza et al. (2004) sug- gested that both subspecies diverged approximately 0.8+0.2 MY BP, an estimation mainly based on the Cy- tochrome b (Cyt b) mitochondrial coding gene (the 12s and the nuclear gene C-mos were uninformative at this lev- el), a divergence time almost identical to the one calcu- lated by Crochet et al. (2004) using also the Cytb gene [1.6 % genetic divergence, which roughly corresponds to 0.6 to 1, with a mean of 0.8 MY BP). These two values were very similar to the above-mentioned ciphers. The inferred divergence time increased up to 1.2 (Cyt b) or 1.6 MY BP (Cyt b+12s) when different terminal taxa evolutionary models and phylogenetic methods were used (Arribas et al. 2006; Arnold et al. 2007; respectively). On the other hand, specimens from Sierra de Béjar branched at the base of the 7. cyreni clade in some mtD- NA analyses (Carranza et al. 2004; Arribas & Carranza 2004). It was suggested that the split of this populations occurred approximately 1.7+0.3 MY BP (Carranza et al. 2004). However, in analyses using the same mtDNA re- gions but different taxa and other evolutionary models and phylogenetic methods than above (Arribas et al. 2006; Arnold et al. 2007), the specimens from Béjar formed a trichotomy with /. c. castiliana from Sierra de Gredos and I. c. cyreni from Navacerrada. Apart from the uninformative C-mos nuclear gene frag- ment analyzed by Carranza et al. (2004) there is only one other information about differences at the nuclear level, the analysis of allozyme data by Almeida et al. (2002), which showed a Nei’s distance of 0.002 between speci- mens from Gredos and Guadarrama. From West to East, the distribution of J. cyreni can be di- vided into two axes connected by low mountain valleys (see appendix II), but not clearly interrupted by clear cut barriers as river valleys. One axis runs across Sierra de Béjar (summit in La Ceya, 2.425 m) and Gredos (Alman- Bonn zoological Bulletin 57 (2): 197-210 zor, 2.592 m), whereas the other axis is constituted by the Sierras de Villafranca (Moros, 2.065 m), La Serrota (Ser- rota, 2294 m), La Paramera (Zapatero, 2.160 m) and slightly separated by lower areas, Guadarrama (Penalara, 2.430 m). The two axes greatly overlap longitudinally leaving the Villafranca, Serrota and Parameras just to the North of the Sierra de Gredos, but at their orographic shad- ow for rains, and climatically more continentalised. This explains the botanical similarities between the Paramera- Serrota-Villafranca axis and the Sierra de Guadarrama (Lucefio and Vargas 1991). In Guadarrama (where Podarcis muralis also exists), I. cyreni occurs only at the highest areas, from 1.760 m (Puerto de Cotos, Puerto de Navacerrada) up to the peaks (2.340 m in Penalara). In Gredos it lives almost from 1.700 to 2.500 m. It was seen in 17. VII. 1986 (own data) in Puerto del Pico (at 1.352 m close to one of the fountains of the pass) but recent research in this area has been to- tally unfruitful. It is possible that these lower stations favoured by accelerated cold winds in the mountain pass- es (Venturi effect) had disappeared by climatic or best, by habitat degradation due to human over-frequentation dur- ing the last 20 years. In Béjar it has been found between 1.837 m (own data) and 2.443 m (see Lizana et al. 1988, 1992, 1993; and Martin 2005 for general data; own data corrections for the confirmed lower limits). Apart from the better known Sierras, the presence of the Carpetane Lizard in the small parallel mountain ranges called “Sierras de Avila” or ““Parameras” (composed by three Sierras: Villafranca, La Serrota, and La Paramera) was first discovered by the mountaineering group “Valle de Ambles” (Lizana et al. 1993), but no specimens have been studied so far. All aspects of morphology, status and relationships of these small and isolated populations from the Sierras de Avila are totally unknown. In these Sierras the species is extremely localized, especially in La Ser- rota and Paramera. In Sierra de Villafranca, the area with a relatively more extended suitable area, I have found it from 1.850 m probably up to the highest areas (Pico Mo- ros, 2.065 m). In La Serrota it is extremely rare and lo- calized, also cornered in the highest parts, from 2.284 m (perhaps 1.935 m where excrements, possibly of this species, were seen; pers. obs.) to the very summit (Pico Serrota, 2.294 m):; and in La Paramera from 1.700 m in the northern slopes to the summit (Pico Zapatero, 2.160 m) (own data). After a three-year prospection of these parallel ranges, I gathered data from these localized and barely known pop- ulations in order to check the relationships of all the Car- petane Lizards throughout its range. My aim is: a) to re- assess differences between J. c. cyreni and I. c. castiliana in the light of the existence of other small and isolated pop- ©OZFMK Intraspecific variability in Jberolacerta cyreni 199 ulations; b) to ascertain the taxonomic status of the Bejar populations and to check if these represent a further sub- species; c) to study both the relationships among the sam- ples from the Sierras de Avila (=Parameras) massifs, as well as their similitude and differences with their neigh- bouring and well known populations from Gredos, or the more distant populations from Guadarrama and Bejar; and d) as the type series of /. cyreni was destroyed during the Second World War (SWW), to choose a Neotype for the species (see appendix I) in order to fix unequivocally the type locality (although apparently all lost, there were al- sO specimens from Gredos in the original type series). MATERIAL & METHODS Morphology A total of 106/92 male specimens, and 136/135 female specimens of /. cyreni with a complete measurements dataset and snout-vent length greater than 45 mm, were included in the univariate (ANOVA) and multivariate (dis- criminant) analyses, respectively. Given that these popu- lations present sexual dimorphism (Arribas 1996, 1999a; Arribas et al. 2006), analyses were carried out for males and females separately. All material is from Oscar Arribas (OA) database. OTUs names, localities and specimens included in the morphological multivariate analysis were as follows (Fig. Ie GUADARRAMA: Sierra de Guadarrama (Madrid and Segovia provinces, Spain). 25 males and 36 females [/. cyreni cyreni]. MIJARES: Puerto de Miyares (Sierra del Fig. 1. Schematic representation of the distribution of /bero- lacerta cyreni in the Spanish Sistema Central. The different lo- calities (OTUs) cited in the text are represented. 1: Béjar; 2: Gre- dos; 3: Villafranca; 4: Serrota; 5: Paramera; 6: Mijares; 7: Gua- darrama. Bonn zoological Bulletin 57 (2): 197-210 Cabezo, Gredos Oriental Massif, Avila province, Spain). 6 males and 7 females. GREDOS: Circo de Gredos (Gre- dos Central Massif, Avila province, Spain). 23 males and 46 females [/. cyreni castiliana]. BEJAR: Sierra de Bé- jar (Gredos Occidental Massif, also known as Sierra de Candelario, Salamanca province, Spain). 11 males and 28 females. VILLAFRANCA: Sierra de Villafranca (Avila province, Spain). 20 males and 14 females. SERROTA: La Serrota (Avila province, Spain). 3 males and 2 females. PARAMERA: Sierra de La Paramera (Avila province, Spain). 4 males and | female. These populations are discontinuous among them (Appen- dix IT) and constitute discrete geographical OTUs (Fig. 1). Due to lower sample, it was necessary to cluster the Ser- rota and Paramera specimens in the male discriminant analysis, and these two plus Villafranca (all of them “Sier- ras de Avila’”’) in the female one. However, reciprocal dis- tances between each one of these poorly represented sam- ples to the best represented ones were carefully checked and commented in the results section. As the three popu- lations from Sierras de Avila seemed to be largely equiv- alent in the multivariate analyses, a posteriori, all of them were treated as a single OTU (S: AVILA) in ANOVA. Characters studied Biometric characters. Snout-vent length (SVL); Forelimb length (FLL); Hindlimb length (HLL); Pileus length (PL); Pileus width (PW); Parietal length (PaL); Masseteric scale diameter (DM); Tympanic scale diameter (DT); Anal width (AW) and Anal length (AL). All linear measure- ments were made with a digital calliper to the nearest 0.01 mm. These measurements were transformed to the follow- ing more informative and not dimensional-depending ra- tios: FLL/SVL (relative forelimb length; “FLL index’); HLL/SVL (relative hindlinb length, “HLL index”); PL/PW (pileus shape, ““Pileus index”); DM/PaL (relative masseteric plate size, ““Masseteric index”); DT/PaL (rel- ative tympanic size, “Tympanic index”); AL/AW (anal plate surface, “Anal form index”) and AS/SVL (V(AL*AW)*100/SVL, relative anal plate size with re- spect to the total length, “Anal size index”’) (Arribas 1996, 2001). The results of the linear measurements and index- es yielded largely similar results. All ratios were given multiplied by 100 to avoid excessive decimal scores. Scalation characters. Supraciliar Granula (GrS) for the right and left sides; Gularia (GUL); Collaria (COLL); Dor- salia (DORS); Ventralia (VENT); Femoralia rigth (FEMr) and left (FEMI); 4". digit Lamellae (LAM); and Circum- analia (CIRCA). ©ZFMK 200 Oscar J. Arribas Statistical Procedures Statistical analyses used in the morphological study in- cluded both Univariate (ANOVA for SVL, scalation char- acters and indexes, with post-hoc Tukey-Kramer tests at P< 0.05 and P < 0.01 to detect differences among sam- ples) as well as Multivariate techniques (Canonical Dis- criminant Analysis, CDA). In this later analysis, each pop- ulation is represented by a centroid (a hypothetical mid- dle individual). Minimum-length spanning tree (MST) was computed from the Mahalanobis’ distance matrix to de- tect the nearest neighbours based on their position in the multidimensional space. MST representation also avoids distortion of UPGMA trees by the reciprocal pairwise dis- tances recalculation in every step during their construc- tion. UPGMA frequently clusters samples reflecting sam- ple sizes than their true relationships. Distances of small samples or isolated specimens appear greatly exaggerat- ed with respect to the well represented ones. As a result of that, the small-sized samples appear ever as the most external or differentiated in UPGMA derived trees (Kunkel et al. 1980; Cherry et al. 1982; Arribas 1997). Moreover, the UPGMA trees based in very unevenly sized samples also gave very poor Cophenetic Correlation In- dexes between the tree-derived ultrametric distances ma- trix and the original Mahalanobis distance matrix and therefore we have not used them (Arribas et al. 2006). To test the significance of the differences among pre-es- tablished groups for the Discriminant Analysis (based in a geographical origin), we carried out an Analysis of Sim- ilarity (ANOSIM) (Clark 1988, 1993) that tests if the as- signed groups are meaningful, this is, more similar with- in groups than with samples from different groups. The method uses the Bray-Curtis measure of similarity to con- struct clusters of specimens. The null hypothesis 1s there- fore that there are no differences between the members of the compared groups (they are randomly blended). R-sta- tistic scales from +1 to -1. Values closer +1 correspond to a perfect case in which all groups were completely dif- ferent (all specimens of the same group are more similar among them than to any specimens of the other groups). R = 0 occurs if the high and low similarities are perfect- ly mixed and bear no relationship to the group, a common situation if some of the groups are largely equivalent. A value of -1 indicates that the most similar samples are all outside of the groups (all groups largely equivalent and randomly formed). To check for significance, pseudorepli- cation tests (1000 randomizations) were run to test if the given results can occur by chance. If the value of R is sig- nificant, there is evidence that the samples within groups are more similar than would be expected by random chance. Even more important, pairwise tests among Bonn zoological Bulletin 57 (2): 197-210 populations permit to test significance of the differences among the concerned groups and to detect which ones are really different from the others. Mahalanobis’ (squared) distance matrices were compared by means of Mantel Test (with 1000 permutations) with matrices composed by Euclidean (squared) distances for the climatic characteristics of localities: a) Precipitation (mm) during the incubation months (July and August, as scalation is invariant during lizard’s life); b) Annual pre- cipitation (mm), c) Temperature (°C) (July and August); d) Annual average temperature e) Sun radiation (n 10 kJ/(m2*day*micrometer)(July and August), and f) Annu- al Sun radiation. Data were extracted from Ninyerola et al. (2005). Also, these Mahalanobis’ distances were com- pared with (d) the aerial (straight) geographical distances among the sampling localities. Multivariate (Discriminant and ANOSIM) analyses were performed with Community Analysis Package 4.0 (Hen- derson & Seaby 2007). MST trees and Mantel tests were calculated with NTSYS 2.1° (Rohlf 2000). Univariate sta- tistics were processed with NCSS 2001° package (Hintze 2001). RESULTS Males Canonical Discriminant Analysis. The CDA conducted with 106 male specimens shows three significant axes that explain an 85 % of the total intersample variation. The two first axes together explain the main part (70.6 %) and dis- criminate fairly well the samples, especially the first one, the unique with an eigenvalue >1. The first discriminant axis has an Eigenvalue of 1.54 (51.2 % of variance ex- plained; Chi-Sq. with 85 df= 200.71, P< 0.0001) and dis- tributes the samples with fairly overlap among them, ex- cept Guadarrama, that has only a small coincidence with the other ones (Fig. 2 A). Guadarrama appears in the neg- ative part of the axis, characterised by the lower values for DORS (0.441553) and VENT (0.560765) and greater values of CIRCA (-0.572683). Second and third axes (eigenvalues < 1) present a considerable overlap among the samples and do not discriminate populations. This discriminant analysis applied to the samples reached a 72.6% of correct classification among the specimens. The Guadarrama sample (/. cyreni cyreni) reaches a 71.9 % of correct classification in respect to all the other sam- ples UZ. c. castiliana). ©ZFMK Intraspecific variability in Jberolacerta cyreni 201 Discriminant Plot - Ib. cyreni MALES Function 2 Function 1 Discriminant Plot - Ib. cyreni FEMALES Function 2 Fig. 2. Function 1 Canonical Discriminant Analysis (CDA) plots for a) males (above) and b) females (below). Specimens, sample centroids and group perimeters are represented. Guadarrama (inverted triangles), Gredos (triangles), Béjar (irregular circles), Mijares (cross), Villafranca (asterisk) and Serrota-Paramera (sail). In females, the three last samples are grouped as Sierras de Avila (sail). Sample centroids are represented by a square. See text for axis characteristics and results. Minimum-length spanning tree (not represented) connect- ing the centroid (hypothetical middle specimens) of each sample is fairly congruent with their geographical posi- tion, connecting in general neighbouring samples. The most “central” (most connected) population is Villafran- ca that connects with Gredos (at Mahalanobis Distance of 3.1870), Béjar (3.2199), Paramera (4.8042) and finally, to the most isolated one, Guadarrama (6.8412). Two popu- lations show overestimated distances due to their small sample sizes: Mijares (East Gredos) that connects with Bé- jar (7.0951), and Serrota with their neighbouring Param- era (9.1822). Bonn zoological Bulletin 57 (2): 197-210 Analysis of Similarity (ANOSIM) (Table 3) shows that there is a considerable overlap among samples (R-statis- tic = 0.122088, P < 0.005; 1000 randomizations) as our value (that can range from | to -1), although positive, is very small. Very significant differences among the (geo- graphically) assigned groups, appear only among Guadar- rama and Bejar, Gredos and Villafranca (P < 0.01) but do not reach significance with Mijares and Serrota + Para- mera (both with small samples). The other populations are not differentiated among them (P > 0.01). ©ZFMK NO S NO Fig. 3. [berolacerta cyreni castiliana. a) La Covatilla Sky re- sort (Sierra de Beéyar), July 2007, Male ; b) El Travieso (S# de Beéjar), July 2004, Female ; c) El Calvitero (Sierra de Béjar), Ju- ly 2004, Female (atypical pattern, with diffumination and coa- lescence in a unique vertebral line); d) Puerto de Miyares (Gre- dos Oriental Massif), July 2006, Female. Bonn zoological Bulletin 57 (2): 197-210 Oscar J. Arribas The Analysis of Variance (ANOVA) (Appendix III, Table 1) indicate that Guadarrama differs from all or nearly all the other populations in VENT and CIRCA (with the smaller and greater values for these parameters, respec- tively, in the former population), but also appeared dif- ferences between Guadarrama and Beyar in Dors (small- er in the former), and with Gredos in PV (greater in the former). An interesting and significant difference appears in DORS among Gredos and Béjar samples (clearly greater in the later). There is no significant correlation among Mahalanobis’ distances and any of the geographic and climatic param- eters analyzed (all Mantel Tests P > 0.05). Females Canonical Discriminant Analysis: The CDA conducted with 136 female specimens shows three significant axes that explain a 93.8 % of the total intersample variation. The two first axes together explain a large part of the vari- ance (85.7 %), and especially along the first one, that ac- counts itself for 62.7 % of the total variation and is the unique with an eigenvalue >1 (1.77), discriminating Guadarrama specimens from the other neighbour samples only with a small overlap (Fig. 2B). The other samples show a considerable overlap among them. Guadarrama discriminates towards the negative part of the axis, char- acterised by the lower values of DORS (0.62) and VENT (0.66) and greater ones of CIRCA (-0.38). Second and third axes (eigenvalues < 1) present a considerable over- lap among the samples and do not discriminate popula- tions. The discriminant analysis applied to the samples reached a 74.26% of correct classification among the specimens. Guadarrama sample (J. c. cyreni) reaches an 87.2 % of cor- rect classification with respect all the other samples (J. c. castiliana). Minimum-length spanning tree (not represented) connect- ing centroids is very similar to the male one. The most con- nected sample is Béjar, which clusters with Villafranca (at 2.9), Gredos (3.5) and Mijares (7.38, but here exaggerat- ed by the scarce sample of the later). Guadarrama connects with the scarcely represented (and geographically inter- mediate) Mijares (East Gredos) (at 6.09), and all the Sier- ras de Avila samples cluster together (Villafranca with Ser- rota + Paramera at 8.66). Analysis of Similarity (ANOSIM) (Appendix III, Table 3) shows that there is a considerable overlap among sam- ples (R-statistic = 0.162588, P < 0.001; 1000 randomiza- tions), but the results are slightly best than for the male ©OZFMK Intraspecific variability in /berolacerta cyreni 203 Fig. 4. [berolacerta cyreni castiliana. a) Pico Zapatero (Si- erra de la Paramera), July 2005, Male; b) Puerto de Pena Negra (Sierra de Villafranca), July 2006, Male; c) Pico Serrota (La Ser- rota Massif), July 2005, Female; d) Pico Serrota (La Serrota Mas- sif), July 2006, Male. Bonn zoological Bulletin 57 (2): 197-210 analysis. Very significant differences (P < 0.01) appear among Guadarrama and all the other samples except with Miares. The other populations are not differentiated among them (P > 0.01). The Analysis of Variance (ANOVA) (Appendix II, Table 2), as in the male analysis, it shows that Guadarrama is the most different one, especially in DORS, VENT and CIRCA (the first two characters smaller, and the third one greater in the former population). Guadarrama also dif- fers from Gredos by its lower GUL, from Béjar by its greater FLL, HLL, a lower SVL; and from Sierras de Avi- la by its greater relative anal scale surface. Also, significant differences appear in SVL between Bé- jar (clearly the great sized female population) and Gre- dos, and among this latter (with relative greater FLL and HLL) with Béjar and Villafranca. As in male analysis, there is no significant correlation among Mahalanobis’ distances and the geographic and cli- matic parameters analyzed (all Mantel Tests P > 0.05). DISCUSSION From the Discriminant and ANOVA analyses it appears that the only differentiated sample is Guadarrama. It ap- pears with very limited overlap with the other samples in the CDA graphs (Figs 2 A and B). Diagnostic characters for this population (nominate subspecies: /. c. cyreni) are the lower values of DORS (difference more marked in fe- males), lower values of VENT and greater CIRCA. More- over, ANOSIM analyses show that Guadarrama is the unique OTU that is significantly different from nearly all the other samples, except from the close population of Mi- jares (in both sexes) and Serrota + Paramera (but these ex- ceptions occur only in the males and probably due to their scarce sample size). Mijares sample (very small) seems in some aspects ap- proaching to Guadarrama (specially in DORS and VENT values) but globally are clearly closer to /. c. castiliana (specially to Béjar in male and female MST). Populations West from Guadarrama show a great overlap in CDA and lack differences in ANOSIM, being morpho- logically fairly equivalent and all of them assimilable to I. c. castiliana. There are only a few scattered very sig- nificant differences among them (P < 0.01) in ANOVA, as for instance among Gredos and Béjar (this latter has greater DORS and a strikingly greater SVL and propor- tionally shorter limbs that Gredos, but only in female spec- imens). The reason of the longer SVL in Béjar females (from 8 mm to | cm greater than in the other populations) ©ZFMK 204 Oscar J. Arribas which leaves proportionately shorter limbs, can be a con- sequence of body elongation that in lacertids appears linked to a greater clutch size (Brana 1996). This is an in- teresting question for future study: if Béjar specimens ef- fectively have greater clutch size that other 1. cyreni pop- ulations. Both the MST results (in which Villafranca and Beéjar are the most connected samples) as well as the presence of related species further West (/. martinezricai and I. mon- ticola), suggest an origin of the species towards the west- ern extreme of their current distribution area. From these westernmost parts, where it also occurs the higher haplo- type diversity (see below), /. cyreni spread towards the East. Despite that the Sierras de Avila (Villafranca, Ser- rota and Paramera) are slightly more aligned with the Guadarrama axis than with Gredos one, we cannot be sure from which of these two mountain ranges the former was colonized, as MST results in males and females are con- tradictory. According to the male analysis Guadarrama is more related to Villafranca, whereas in the female analy- ses, it is Mijares (Eastern Gredos) the most related one. The results of the mitochondrial analyses of these sam- ples (Cyt B and 12s) (unpublished, Carranza, pers. com.) indicates that the interruption of gene flow 1s fairly recent, as a common haplotype appears in all populations except Villafranca and Bejar. All Gredos, Guadarrama and La Serrota specimens are identical for these two mitochon- drial fragments. Independent changes in one nucleotide with respect to the common haplotype appear in Villafran- ca (the unique change is different in two specimens), Para- mera, Miares and Béjar specimens, and two changes ac- cumulate in one Béjar and one Mijares specimen (others have only one). The current morphological differences of Guadarrama specimens seem to be relatively recent, and are possibly the result of bottleneck effects during the colonization process, or alternatively of strong selective pressures (or a combination of both causes). Conversely, the absence of marked differences among the other populations (more or less with a similar age) could be due to the main- tenance of a more continuous gene flow among them, re- sponsible of maintaining their morphology largely equiv- alent (nuclear genes remain unstudied). The current larg- er geographical gap in the distribution of /. cyreni occurs precisely between Guadarrama and the remaining popu- lations to the West. Despite the presumably short isolation time, as comment- ed above, a considerable selection pressure or a genetic bottleneck in the expanding populations might have pro- moted and fixed the morphological differences now seen in Guadarrama specimens. These factors do not seem to Bonn zoological Bulletin 57 (2): 197-210 be due to isolation-by-distance processes but by histori- cal vicariant events (cf. Irwin 2002) as there is no rela- tionship between morphological differentiation and geo- graphic distances. Also there is no relationship among the more obvious climatic parameters (precipitation, temper- ature and sun radiation) and these differences. Concerning the position of Béjar populations, only spec- imens from the west-facing slopes of the massif have been studied, and therefore it is possible that in other parts of the massif other haplotypes (the common one with Gre- dos) could be present. The species was cited from “Puer- to de Tornavacas, SA” (for Salamanca, sic.!; a mistake as this locality is in Avila) (Lizana et al. 1992). This is the natural pass between Béjar and Gredos, but I have been unable to find it there. This place 1s a sub-Mediterranean environment with Pyrenean Oak open forest inhabiting populations of Zimon lepidus, Psammodromus algirus and even Buthus occitanus, all thermophylous species typical from dry conditions. The lower height of Puerto de Tor- navacas (1010m) makes me to suspect that /. cyreni is not there and the record is possibly a mistake. One possibil- ity is that it was from the higher neighboring mountains. An account about the pattern and coloration of [. cyreni is in Arribas (1996) and it is especially detailed for the main Sistema Central massifs in Perez-Mellado et al. (1993). Both colour as well as the dark pattern, seem to be selected in accordance to the substrate characteristics. Overall, the background colour is brown in juveniles and subadult specimens, changing in different percentages to green in adult specimens (more frequently in males and becoming more vividly linked to reproductive processes). In populations inhabiting rocks (plenty of Rhizocarpon gtr. geographicum lichens) as in Gredos and the upper parts of Guadarrama (Penalara) green adults are more frequent (both males and big females). When living in rocky talus with sands and bare ground (as in Navacerrada area) brownish adult specimens are more frequent (Arribas 1999b: Figs 9 and 10). Concerning the reticulate pattern, it also varies in a dif- ferent degree among the different populations depending on the substrates inhabited. Juveniles and subadults have temporal uniform or reticulate bands that coalesce during growth with dorsal spots (more frequently in males) giv- ing reticulated-like patterns. Usually, there is a relation- ship between the size of the granite phanerocrystals (the granite-rock spotting) and the habitus of the lizards liv- ing on it. Lizards living on rocks that present large crys- tals (as for instance the Béjar ones) are more reticulated than specimens living in places in which the rocks pres- ent smaller crystals (and thus finely spotted). Coloration accounts described in Perez Mellado et al. (1993) are fair- ly precise, especially for Gredos specimens. Concerning ©ZFMK Intraspecific variability in Jberolacerta cyreni 205 Bejar (=Candelario in Perez Mellado et al. [1993]) the de- scription should be corrected as, although it is true that very old specimens are fairly reticulated, especially males, females more frequently have two paravertebral rows of distinctive spots (photo 2), as in females of other popu- lations and in mid-grow specimens of both sexes in all lo- calities. The statement that “the most common background colour of the back and flanks is greenish or bluish” prob- ably is true for fully adult specimens during the breeding period, but in July and August only some big males con- serve greenish tones, appearing even fully adult females more or less brownish. Concluding: a) /. c. cyreni is a recently differentiated but well diagnosable (morpho)subspecies, a case paralleling the relationship of 1. monticola monticola from Serra da Estrela (Portugal) with repect to 1. m. cantabrica (from Galicia and Cantabrian Mts.), in this later case, with no genetic differences, but with singular morphological traits that distinguishes it from other /. monticola in a multivari- ate analysis (Arribas & Carranza 2004; Arribas et al. 2006). Highly variable nuclear markers as for instance in- trons or microsatellites may help to clarify definitively the status of these well diagnosable (morpho)subspecies. b) Despite the mtDNA differences of the Béjar specimens (one or two nucleotides), their morphology is largely equivalent to /. c. castiliana. Lacking data from other Beé- jar populations and genetic nuclear markers, I assume that these morphological similarities reflect the presence of a very recent gene flow with other neighbouring popula- tions. The Béjar populations are however outstanding by their female body elongation (up to near | cm larger), con- serving proportionately shorter limbs, which can be a strat- egy to increase clutch size. c) The Sierras de Avila populations are very similar; the closer among all the populations compared. One of them, the Sierra de Villafranca, is together with Béjar, the most connected sample, and it is, from a morphological point of view, at the root of the species expansion. Both also present the unique slightly variant haplotypes. All they should be considered as /. c. castiliana by their closer iden- tity with Gredos. Acknowledgements. Dr. Sergi Pla (Barcelona, Spain), Jesus Garcia (Huesca, Spain) and Dr. Vicente M. Ortuno (Madrid, Spain) helped in some prospections. Dr. Salvador Carranza (Barcelona, Spain) corrected and improved parts of the manu- script. This study was financed completely by the author and therefore it did not cost any Euro to the public arks. Prospec- tions of the small Sierras de Avila and Béjar populations were carried out under permissions n° 20051630007003 (2005), 20061630024599(2006), 2007167004130 (2007), 20081630020386 (2008), 20092390004760 (2009), issued by the competent organisms in charge of the wildlife protection (Jun- ta de Castilla y Leon, Spain) Bonn zoological Bulletin 57 (2): 197-210 REFERENCES Almeida AP, Rosa HD, Paulo OS, Crespo EG (2002) Genetic differenciation of populations of Iberian rock-lizards [bero- lacerta (Iberolacerta) sensu Arribas (1999). Journal of Zoo- logical Systematics and Evolutionary Research 40: 57-64 Arnold EN, Arribas O, Carranza S (2007) Systematics of the Palaearctic and Oriental lizard tribe Lacertini (Squamata: Lac- ertidae: Lacertinae), with descriptions of eight new genera. 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Spixiana 30, 2: 201-274 Henderson PA & Seaby RMH (2007) Community Analysis Pack- age 4.0 Pisces Conservation Ltd, Lymington, UK. 164 pp. (www.pisces-conservation.com) Irwin DE (2002) Phylogeographic breaks without geographic barriers to gene flow. Evolution 56: 2383-2394 Kunkel JG, Cherry LM, Case SM, Wilson AC (1980) Reply to W.R. Atchley’s “M-Statistic and Morphometric divergence. Science 208: 1060-1061 ©ZFMK 206 Oscar J. Arribas PSE Sa dee ow Custer eer a os Fig. 5. Lizana M, Ciudad MJ, Pérez-MelladoV (1988) Distribucion al- titudinal de la herpetofauna en el Macizo Central de la Sierra de Gredos. Rev. Esp. Herpetol. 3: 55—67 Lizana M, Ciudad MJ, Gil M, Guerrero F, Pérez-Mellado V, Martin-Sanchez R (1992) Nuevos datos sobre la distribucion de los anfibios y reptiles en el macizo central de la Sierra de Gredos. Rev. Esp.Herpetol. 6: 61-80 Lizana M, Martin-Sanchez R, Morales JJ, Lopez-Gonzalez J, Gutierrez J (1993) Nuevas poblaciones de la lagartija serrana (Lacerta monticola cyreni) en las sierras de Avila. Bol. Asoc. Herp. Esp. 4: 5—6 Martin J (2005) Lagartija carpetana - [berolacerta cyreni. En: Enciclopedia Virtual de los Vertebrados Espanoles. Carrascal LM, Salvador A (Eds.) Museo Nacional de Ciencias Natura- les, Madrid. http://www.vertebradosibericos.org/ Mayer W & Arribas O (1996) Allozyme differentiation and re- lationship among the Iberian-Pyrenean Mountain Lizards (Squamata: Sauria: Lacertidae). Herpetozoa 9 (1/2): 57-61 Mayer W & Arribas OJ (2003) Phylogenetic relationships of the European lacertid genera Archaeolacerta and Iberolacerta and their relationships to some other ‘Archaeolacertae’ (sensu la- to) from Near East, derived from mitochondrial DNA se- quences. Journal of Zoological Systematics & Evolutionary Research 41: 157-161 Bonn zoological Bulletin 57 (2): 197-210 reset) Cee eees 5o4 pee eeomet sis aan ee Iberolacerta cyreni NEOTYPE (here designed). MNCN n. 39934. Miller L & Hellmich W (1937) Mitteilungen tiber die Herpeto- fauna der Iberischen Halbinsel. II. zur Kenntnis der Lacerta monticola. Zool. Anz. 117: 65—73 Ninyerola M, Pons X, Roure JM (2005) Atlas Climatico Digi- tal de la Peninsula Ibérica. Metodologia y aplicaciones en bio- climatologia y geobotanica. Universidad Autonoma de Bar- celona, Bellaterra. Odierna C, Aprea G, Arribas OJ, Capriglione T, Caputo V, Ol- mo E (1996) The Karyology of the Iberian Rock Lizards. Her- petologica 52 (4): 542-550 Pérez-Mellado V, Barbadillo LJ, Barahona F, Brown RP, Corti C, Guerrero F, and Lanza, B (1993) A systematic survey of of the Iberian Rock lizard Lacerta (Archaeolacerta) montico- la. In: Valakos E, Boehme W, Pérez-Mellado V and Maragou P (eds.) Lacertids of the Mediterranean region. Athens: Hel- lenic Zoological Society 85—105 Received: 29.VI.2010 Accepted: 15.X.2010 ©ZFMK Intraspecific variability in Jberolacerta cyreni 207 APPENDIX I Neotype designation for /berolacerta cyreni (Miller & Hellmich, 1937) The type series of /berolacerta cyreni (Lacerta monticola cyreni Miiller & Hellmich, 1937) included 66 specimens (not only from Guadarrama, the species’ type locality, but also some paratypes from Gredos) formerly deposited in the Zool. Staatssamlung (Herpet. Samml.) Miinchen. In fact, two syntype specimens, male and female, respectively numbered “ZSM (SLM) 2329 a” and “ZSM (SLM) 2329 b”, were originally considered as types labelled “Guadarrama, Puerto de Navacerrada. W. Hellmich”. (Miller & Hellmich, 1937). Although it seems that all the original type series was destroyed during the Second World War (Franzen & Glaw, 2007), and due to the fact that in this type series there were included some Gre- dos specimens (today part of another subspecies; /. c. castiliana), and also that there was early confusion about the /. monticola type locality which lead to the description of a new taxon as Lac- erta estrellensis Cyren, 1928 (Arribas 2008), I design a new type specimen (neotype) to fix unequivocally the type locality against any contingence (as could be the highly improbable apparition of any “surviving” original Gredos paratype). I designate here as NEOTYPE for the species a specimen from the Museo Nacional de Ciencias Naturales (Madrid) (MNCN n. 39934) (Fig. 5). A male labelled as follows: Left hindleg: (white label, Typewriter letter) MNCN (anverse), 39934 (reverse). Right hindleg: (white label, pencil handwritten) “Pto. de Cotos—Pto de Navacerrada. Srra. de Guadarrama (_ )[blank in- side parenthesis], 21-IV—84, 18,15 h. Sol. Pedriza en pinar con nieve. Ps=8.5 gr.” (no collector’s data). Left foreleg: (White label, ink handwritten) Neotypus. O. Ar- ribas designatio (anverse), “Lacerta monticola cyreni Miller & Hellmich, 1937” (=/berolacerta cyreni) (reverse). Right foreleg: (Red plastic label, Dymo® lettering) NEOTYPUS. Neotype description (Fig. 5): Biometry: Adult male with snout—vent length of 66.85 mm. Tail 126 mm (intact). Forelimb length 23.34 mm. Hindlimb length 34.52 mm. Pileus length 16.4 mm. Pileus width 8.2 mm. Pari- etal legth 5.5 mm. Masseteric widest diameter 2.71 mm. Tym- panic widest diameter 1.94 mm. Anal plate width 5.04 mm. Anal length 3.19 mm. FLL/SVL (relative forelimb length): 0.349. HLL/SVL (relative hindlimb length): 0.5163. PL/PW (pileus Bonn zoological Bulletin 57 (2): 197-210 shape): 2.003. DM/PaL (relative masseteric plate size): 0.491. DT/PaL (relative tympanic size): 0.352. AL/AW (anal plate sur- face): 0.6329. AS/SVL (relative anal plate size in respect to to- tal legth): 5.998. Scalation: Number of supraciliary granules: 9 (rigth) and 11 (left). Supralabials: 5 (both sides). Sublabials: 6 (right side) and 7 (left side). Submaxillars: 6 (both sides). Gularia: 25. Collar- ia: 9. Dorsalia: 53. Ventralia: 26. Femoral pores 19 (right) and 18 (left). Lamellae: 25. Circumanal Plates: 8. Rostral in full con- tact with frontonasal. Supranasal separated from first loreal. One postnasal (in both sides). First Postocular separated from Pari- etal plate. Alternate wide and narrow scale rings in the tail. Twen- ty—six scales across one of these rings. Coloration: (in alcohol). Dorsal tract and pileus brown (proba- bly also in life), densely spotted of medium-sized black spots that in the middle of the dorsum nearly form transverse bands and connect the two temporal bands. Temporal bands reticulat- ed fairly dark (black—brown) with traces of clear occelli (bare- ly visible) inside, also connecting with more light reticulated with infratemporal band (barely discernible). No blue axillar occel- li. Traces of blue points in the outermost ventral scales. Only the outermost ventral scale ranges are clearly black spotted. Bel- ly light bluish or white—bluish. APPENDIX II Barriers and high mountain passes (among parentheses) between the different /. cyreni populations. All these intermediate areas are at present apparently devoid of /. cvreni, thus constituting these OTUs discrete populations: BEJAR-GREDOS: no barrier (Puerto de Tornavacas, 1275m). BEJAR-VILLAFRANCA: Tormes River Valley (no pass). GREDOS-VILLAFRANCA: Tormes River Valley (Collado de Cepegato, 1550m). GREDOS-S? AVILA[Serrotat+Paramera]: Alberche River Valley (no passes). SERROTA—PARAMERA: no barrier (Puerto de Menga, 1566m). VILLAFRANCA-SERROTA: no barrier (Puerto de Chia, 1663m). S* AVILA (as a whole) -GUADARRAMA: no barrier (Puerto del Boqueron, 1315m). GREDOS—GUADARRAMA: Alberche River Valley (no pass) ©ZFMK Oscar J. 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(91=u) (s) (gz=u) (p) — (9b=) (¢) (L=u) (2) (9€=u) (1) = Sr S-£ P= SZ at Sant Ge] OU co] So) a d Our hy V'TIAV ¢S UVrAA soadauDd SAUVETA waAVdVAD saTVNAd S f=) 5 faa} “So[eUUloy Mad J OF yNQ “| BqQui uUlsy °Z 9IqUL 210 Oscar J. Arribas Table 3. Analysis of Similarity (ANOSIM) results (with 1000 randomizations). Males above diagonal and females below. The number is the test probability results among each two concerned populations (significant results in bold). FEM \ MAL BEJAR GREDOS GUAD. MIJARES SERR-PAR VILLAFR. BEJAR — 0.058 0.001 0.644 0.138 0.064 GREDOS 0.021 — 0.001 0.691 0.867 0.198 GUAD. 0.001 0.001 = 0.097 0.128 0.001 MIJARES 0.53 0.039 0.327 — 0.224 0.513 SERR-PAR OM27, 0.025 0.001 0.217 — 0.86 Bonn zoological Bulletin 57 (2): 197-210 ©ZFMK [ Bonn zoological Bulletin Volume 57 | Issue 2 pp. 211-229 Bonn, November 2010 Insights into chameleons of the genus Trioceros (Squamata: Chamaeleonidae) in Cameroon, with the resurrection of Chamaeleon serratus Mertens, 1922 Michael F. Barej!;*, Ivan Ineich?, Vaclav Gvozdik34, Nathaly Lhermitte-Vallarino>, Nono Legrand Gonwouo®, Matthew LeBreton’, Ursula Bott’, & Andreas Schmitz? 1 Museum fiir Naturkunde, Leibniz Institute for Research on Evolution and Biodiversity at the Humboldt University Berlin, Invalidenstrasse 43, D-10115 Berlin, Germany 2 Muséum National d’Histoire Naturelle, Département Systématique et Evolution (Reptiles et Amphibiens), UMR CNRS 7205, CP 30, 25 rue Cuvier, F-75005 Paris, France 3 Institute of Animal Physiology and Genetics, Academy of Sciences of the Czech Republic, Department of Vertebrate Evolutionary Biology and Genetics, Rumburska 89, CZ-277 21 Libéchov, Czech Republic 4 National Museum, Department of Zoology, Cirkusova 1740, CZ-19300 Prague, Czech Republic 5 Muséum National d’Histoire Naturelle, USM 30705 Département Ecologie et Gestion de la Biodiversité & CNRS IFR 101, Parasitologie comparée et Modéles expérimentaux, 61 rue Buffon, 7 CP52, F-75231 Paris cedex 05, France 6 Université of Yaoundé I, Faculty of Science, Laboratory of Zoology, P.O. Box 812, Yaoundé, Cameroon 7 Global Viral Forecasting Initiative, BP 7039, Yaoundé, Cameroon 8 Zoologisches Forschungsmuseum Alexander Koenig, Adenauerallee 160, D-53113 Bonn, Germany ° Muséum d’histoire naturelle, Department of Herpetology and Ichthyology, C.P. 6434, CH-1211 Geneva 6, Switzerland * Corresponding author: E-mail: michael@pbarej.de Abstract. Relationships among chameleons of the genus Zrioceros in Cameroon are reviewed on a molecular basis us- ing mitochondrial genes and by morphology. Trioceros oweni is placed basal to two distinct clades (lowland-submon- tane species vs. submontane-montane species) and its position is discussed due to high genetic differences to the remain- ing taxa. Within the lowland-submontane species group, distinct subclades with low genetic differences exist within 7° montium and T. cristatus. Differing relationships to previously published results are observed within the submontane- montane species group, resulting in taxonomic changes: 7rioceros eisentrauti 1s grouped with the two 7) quadricornis subspecies, showing only low genetic differences, which also correlates with the similar overall morphology. The taxon is thus assigned to a subspecific rank: T. quadricornis eisentrauti. Within the wiedersheimi-group, the former southern subspecies is elevated to species rank, 7rioceros perreti, and two additional species have been distinguished by molec- ular and morphological methods in the former nominate taxon. Trioceros wiedersheimi is restricted to northernmost lo- calities, while remaining populations have been assigned to the revalidated taxon Trioceros serratus (Mertens, 1922). Differentiating morphological characters for the three species are provided and a neotype of Chamaeleon serratus Mertens, 1922 is designated and described to ensure clarification of its taxonomic status and type locality. Key words. Reptilia, Chamaeleonidae, Trioceros, Trioceros serratus, Africa, Cameroon, phylogeny, taxonomy. INTRODUCTION Only recently, Tilbury & Tolley (2009) provided molec- ular evidence that the two former subgenera (Chamaeleo Laurenti, 1768 sensu stricto and Trioceros Swainson, 1839) of the chamaeleonid genus Chamaeleo as recog- nized by Klaver & BOhme (1986) represent two distinct and valid genera. For a diagnosis of the two genera see Klaver & Bohme (1986, 1992) and Tilbury & Tolley (2009). Klaver & Bohme (1992) additionally provided a detailed overview of formerly published knowledge on the Bonn zoological Bulletin 57 (2): 211-229 cristatus-subgroup inside the Trioceros-group from Cameroon. The Republic of Cameroon exhibits a very high diversi- ty of chameleon species compared to adjacent countries, especially in montane areas (BOhme & Klaver 1981; Gonwouo et al. 2006; Herrmann et al. 2005, 2006). At present 14 species of chameleons are known to occur in Cameroon. They belong to the genera Chamaeleo [five ©ZFMK 212 Michael F. Barej et al. species: C. africanus Laurenti, 1768; C. dilepis Leach, 1819; C. gracilis Hallowell, 1842; C. quilensis Bocage, 1886; C. senegalensis Daudin, 1802], Rhampholeon [one species: R. spectrum (Buchholz, 1874)] and Trioceros [eight species: 7. camerunensis (Miller, 1909); T. crista- tus (Stutchbury, 1837); 7! eisentrauti (Mertens, 1968); T. montium (Buchholz, 1874); 7. oweni (Gray, 1831); T. pf- efferi (Tornier, 1900); 7. quadricornis (Tornier, 1899); T. wiedersheimi (Nieden, 1910)]. Trioceros quadricornis and T. wiedersheimi are polytypic, with one more subspecies [TZ g. gracilior (BOhme & Klaver, 1981), TZ. w. perreti (Klaver & Bohme, 1992)], resp. (BOhme & Klaver 1981; Chirio & LeBreton 2007; Gonwouo et al. 2006; Klaver & Bohme 1986; Tilbury & Tolley 2009). According to Klaver & Bohme (1997) and Uetz & Hallermann (2010) one additional species (Chamaeleo laevigatus Gray, 1863) is present in Cameroon, but this species has not been listed by other recent authors (Chirio & LeBreton 2007; Gonwouo et al. 2006; Tilbury 2010). While some species such as Chamaeleo gracilis or Trioceros cristatus show a large distribution ranging at least from Nigeria to Gabon and the Congo (Necas 2004), five species are regarded as montane endemics occupying restricted high elevation ar- eas along the Cameroon mountain chain, with 7) eisen- trauti the most restricted, being endemic to the Rumpi Hills in western Cameroon (Chirio & LeBreton 2007; Gonwouo et al. 2006; Klaver & BOhme 1992). Pook & Wild (1997) published a preliminary phylogeny of Trioceros from Cameroon, and we herein provide ad- ditional and new insights into this species group based on additional material. MATERIAL AND METHODS In all, 49 combined, mitochondrial 16S and 12S rRNA gene fragments, sequences (Tab. 1, Appendix II; museum acronyms see below) comprising 964 bp (lengths refer- ring to the aligned sequences including gaps) were ob- tained. One short section (4 bp from the 12S gene) was too variable to be reliably aligned, and was excluded from the analyses, resulting in a total of 960 bp which were used in the analyses. Kinyongia tavetana (AM422414/ AM422433; Mariaux et al. 2008) was used as outgroup. Its position outside of 7rioceros was demonstrated by Tilbury & Tolley (2009). DNA was extracted using Qi- Amp tissue extraction kits (Qiagen) and the peqGold Tis- sue DNA Mini Kit (PEQLAB Biotechnologie GmbH) (see Wagner et al. 2009a). The primers l|6sar-L (light chain; 5’— CGC CTG TTT ATC AAA AAC AT - 3’) and 1 6sbr- 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. Additionally, a section of the mitochondrial 12S ri- Bonn zoological Bulletin 57 (2): 211-229 bosomal RNA gene was amplified 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). PCR cycling procedures were as described in Schmitz et al. (2005). PCR products were purified using Qiaquick purification kits (Qiagen). 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). PAUP* 4.0b10 (Swofford 2002) was used to com- pute the uncorrected pairwise distances for all sequences (Tab. 2, Appendix II). We performed neighbour-joining (NJ), maximum parsimony (MP), maximum likelihood (ML) and Bayesian reconstructions. For ML and Bayesian analysis parameters of the model were estimated from the data set using Modeltest 3.7 (Posada & Crandall 1998) and MrModeltest 2.3 (Nylander 2002), respectively. For the MP analysis, we used the “heuristic search” with the “ran- dom addition” option of PAUP* (Swofford 2002) with 10 replicates, using the TBR (tree bisection-reconnection) branch swapping option. For the ML tree we used the PhyML 3.0 computer cluster of the Montpellier bioinfor- matics platform (http://www.atgc-montpellier.fr/phyml/) (Guindon & Gascuel 2003). All Bayesian analyses were performed with MrBayes, version 3.12 (Huelsenbeck & Ronquist 2001). The exact parameters used for the Bayesian analyses followed those described in detail by Reeder (2003). For the Bayesian reconstruction clades with posterior probabilities (PP) = 95% were considered strongly (significantly) supported. Additionally, we used bootstrap analyses with 1000 (for ML), 2000 (for MP) and 20000 (for NJ) pseudoreplicates to evaluate the relative branch support in the phylogenetic analysis. In the morphological analysis measurements follow stan- dard procedures (e.g.Werner 1902; Mariaux et al. 2008) and were taken on preserved material with an electronic dial calliper (+ 0.1 mm). All measurements are given in mm (Tab. 3, Appendix II). Analysis of morphological da- ta has been performed using PAST software (Version 1.82b; Hammer et al. 2001). If measurements (e.g. femur length) differed between body sides, mean values were used. Photos of living specimen have been used to analyse colouration patterns. Investigated specimens are deposited in Muséum d’his- toire naturelle, Geneva (MHNG); Muséum national d’Histoire naturelle, Paris (MNHN); National Museum, Museum of Natural History, Prague (NMP6V); Zoologi- sches Forschungsmuseum Alexander Koenig, Bonn (ZFMK); Museum fiir Naturkunde, Leibniz-Institut fiir Evolutions- und Biodiversitatsforschung an der Humboldt- Universitat zu Berlin (ZMB); Zoologische Staatssamm- lung Miinchen (ZSM). OZFMK Chameleons of the genus Trioceros from Cameroon 21 RESULTS To date ten taxa belonging to the genus Trioceros have been recognized in Cameroon (eight species + two sub- species), but the present phylogenetic analysis is incon- sistent with this arrangement (Fig. 1). All four used phy- logenetic methodologies strongly agree in the overall topology and in all cases support the same terminal clades. The phylogenetic analyses reveal only a single difference (discussed below) for the individual analysis of the two applied gene fragments (not shown), therefore, we only discuss the results of the combined analysis. Distances between ingroup and outgroup species averaged 11.48% (10.58%-—12.30%); Tab. 2, Appendix II). Interspe- cific distances within the ingroup ranged from 3.21%-6.90% excluding 7: oweni. Trioceros oweni 1s the most basal taxon in respect to all ingroup taxa, which are grouped within one clade fully supported in NJ, MP and ML, while still strongly, but not fully significantly supported in the Bayesian reconstruction (PP: 0.91). The main clade is divided into two major subclades with strong statistical support. The first subclade includes 7. camerunensis, T. cristatus, and T; montium, but their mutual relationships remain unresolved. However, partly well supported substructure can be recognized within the two species, 7. montium and T. cristatus. Trioceros camerunensis stands in a basal position to 7’ montium, but this is only significantly supported by the MP reconstruction. Within 7. montium, we find a subdivision into three only slightly differentiated subclades. Overall the uncorrected p-distances of the included 7 montium vouchers range between 0.00%- 0.75%. Similarly, within 7. cristatus a similar subdivision into three more distinct subclades is apparent. Here, the genetic distances between the included 7) cristatus vouchers ranges between 0.00%—1.28%. Contrary to the first major ingroup subclade, relationships of the species of the second major subclade remain unre- solved and form a basal polytomy. Nonetheless, all ter- minal clades in this second major subclade are strongly supported and are well distinct regarding the individual branch lengths and bootstrap support for each terminal clade, mostly corresponding to currently accepted species within Zrioceros. The morphologically very distinct tax- on 7: eisentrauti is grouped together with the two de- scribed 7. quadricornis subspecies with uncorrected p-dis- tance values of between 0.51%-—1.08% between these three taxa. We found only one haplotype in each of the two sub- specific taxa, 7. g. quadricornis and T. q. gracilior, while in T. eisentrauti we uncovered a difference of two nu- cleotide substitutions in our newly gained sequences in comparison to the published 12S sequence of Pook & Wild Bonn zoological Bulletin 57 (2): 211-229 Ww (1997). The distances of 7. eisentrauti to the nominate form T. q. quadricornis (0.51%-—0.64%) are about equal in size to the distances of the latter to 7. qguadricornis gra- cilior (0.63% 0.64%). The distance of 7. eisentrauti to T. quadricornis gracilior is only moderately higher (1.02%-—1.08%). These values are clearly within the in- traspecific distance range of all included Trioceros species. Contrarily, the remaining taxa of this subclade show a much higher genetic differentiation between each other, ranging from 3.18%-—5.00%. These other terminal clades correspond to the taxa 7! pfefferi and T: wiedershei- mi. The latter hornless taxon is represented by three ge- netically well differentiated clades. Two of them corre- spond to the two so far described subspecies, but we find a further significant split within the populations current- ly assigned to the nominate form. DISCUSSION Following our molecular and morphological results sev- eral changes are necessary among Cameroonian chameleons of the genus Trioceros. The overall number of Trioceros taxa in Cameroon is raised to eleven and two already known taxa are revised in their taxonomic rank. Within the Cameroonian Jrioceros, T. oweni is the most basal taxon, while the other taxa form two subclades, in which 7. camerunensis, T. cristatus, T. montium form a lowland to submontane group while remaining taxa of the second subclade inhabit submontane to montane habitats (Pook & Wild 1997). Results and required changes will be discussed below in separate sections referring to the relevant species groups. Trioceros oweni (Gray, 1831) (Fig. 2A) Trioceros oweni, the type species of the genus Trioceros, is the most basal in respect to all remaining Cameroon- ian taxa (Fig. 1). The value of uncorrected p-distance be- tween 7. oweni and the outgroup taxon Kinyongia tave- tana (12.21%) is within the genetic distance range of all included Trioceros taxa to the outgroup (10.58—12.30%, Tab. 2, Appendix II). However, values of uncorrected p- distances between 7) oweni and remaining Cameroonian Trioceros taxa (8.57—10.22%) are significantly higher than values in-between the remaining ingroup taxa (see Tab. 2, Appendix II), and the maximum distance value is on- ly marginally lower than the minimum distance of all 7ri- oceros to the outgroup taxon. Based on molecular data, Pook & Wild (1997) suggested that 77 oweni might belong to a distinct species group, being closer related to 7. john- stoni, an East African species, than to other western 7ri- oceros. In the past, Werner (1902) grouped 7: oweni to- gether with 7. johnstoni, T: melleri and T: werneri, while T. cristatus, T: montium, T. pfefferi and T: quadricornis be- ©OZFMK 214 Michael F. Barej et al. Kinyongia tavetana AM422414/AM422433 Ea fad 2 */* Trioceros ap oweni Campo region/Nkoelon E146.15 camerunensis Mt. Cameroon/Njonji E130.1 montium Rumpi Hills/Mofako Balue E179.18 montium Bakossi Mts./Edib Hills E188.19 montium Bakossi Mts./Edib Hills E188.18 montium Mt. Kupe/Nyasoso E180.15 montium Bakossi Mts./Edib Hills E188.20 montium Mt. Kupe E130.5 montium Mt. Kupe E131.3 3| montium Mt. Cameroon E130.4 montium Mt. Cameroon E131.2 cristatus Rumpi Hills/Mofako Balue E180.2 © cristatus Rumpi Hills/Big Massaka E180.7 El ie Lx 5 fe a/* 3 cnistatus Mt. Cameroon/Njonji E131.1 cristatus Mt. Cameroon/Njonji E130.2 cnstatus Mt. Cameroon/Njonji E130.3 cristatus Campo region/Nkoelon E150.8 cristatus Mamfe region/Amebishu E146.13 cristatus Mamfe region/Amebishu E150.7 quadricomis eisentrauti gquadricornis eisentrauti Rumpi Hills/Mt. Rata E178.10 7| © quadncornis eisentrauti Rumpi Hills/Mt. Rata E178.11 Lx bl fd + /% */x ai fs */e quadricornis quadricomis Mts. Manengouba E131.5 8| quadricornis quadricornis Mts. Manengouba E130.9 quadricornis quadricomis Mts. Manengouba E130.10 quadricornis quadricornis Mts. Manengouba E131.8 H guadncornis gracilior Mt. Oku E130.8 guadnicomis gracilior Mt. Oku E130.7 guadncornis gracilior Oku village E131.4 pfeffen perreti Mts. Manengouba E131.6 perreti Mts. Manengouba E130.11 perreti Mts. Manengouba E130.12 serratus Mt.Mbam E178.3 serratus Mt.Mbam E178.5 eof fs tlhe ake 0.1 serratus Mt.Mbam E178.2 serratus Mt.Mbam E178.4 serratus Big Babanki E188.16 serratus Belo, Mt. Oku [NEOTYPE] E130.17 serratus Big Babanki E189.8 serratus Oku village E131.16 serratus Oku village E131.17 serratus Lake Oku E130.16 serratus Mt. Oku E130.15 serratus Oku village E131.7 wiedersheimi Tchabal Mbabo E91.6 wiedersheimi Tchabal Mbabo E178.1 wiedersheimi Tchabal Gangdaba E188.13 Fig. 1. | Phylogram of the combined analysis of the 16S and 12S rRNA sequence fragments (49 sequences / 960 bp in total). The star symbol “*” denotes significantly supported nodes. [The values for the internal nodes are as follows (NJ/MP/PP/ML, respec- tively): 1:(78/61/0.78/73); 2:(95/98/1.00/96); 3:(99/87/0.99/99); 4:(100/98/1.00/100); 5:(100/93/1.00/100); 6:(86/96/1.00/92); 7:(81/89/0.82/86); 8:(97/67/0.56/94); 9:(100/89/0.97/99); 10:(69/83/0.61/63)]. Bonn zoological Bulletin 57 (2): 211-229 OZFMK Chameleons of the genus 7rioceros from Cameroon 215 longed to a different morphological group. A simple BLAST search in GenBank, performing a similarity check of sequences, of the applied 7’ oweni-sequence identified T. melleri (16S) or T: sternfeldi (12S) to show the high- est similarity values; both again East African species. Ac- cording to Townsend & Larson (2002) and Tilbury & Tol- ley (2009), 7. melleri is related to T. johnstoni. While the only western 7rioceros (T: feae from Bioko Island, Equa- torial Guinea) in the study of Tilbury & Tolley (2009) is placed basal to all other Zrioceros. Similarly, Townsend & Larson (2002) found that all western 7rioceros (includ- ing 7: feae) studied by them stand as a sister clade to the other members of the genus. Hence, concerning 7: oweni our results support the view of Pook & Wild (1997) that Trioceros taxa in western Cen- tral Africa are more closely related to each other than to T. oweni. The exact position of 7: oweni remains to be as- sessed in future studies with a wider sampling from the whole distribution area of this genus. Lowland-submontane clade Trioceros camerunensis (Miller, 1909) (Fig. 2B) In the past Mertens (1964) classified 7) camerunensis as a subspecies of 7’ montium based on morphological sim- ilarities and zoogeographical affinity, but Klaver & Bohme (1992) reclassified the taxon as a valid species. Our mo- lecular results do support close relationships between 7° camerunensis and T: montium but also confirmed its full species status. According to Pook & Wild (1997), 7: camerunensis 1s more closely related to T. feae (not in- cluded in our study) than to 7’ montium. Lowland-submontane clade Trioceros montium (Buchholz, 1874) (Fig. 2C) Within the well supported monophyletic 7’ montium-clade, distinct subclades appear (Fig. 1; Tab. 2, Appendix II). Buchholz (1874) described T. montium from Bonjongo, Mt. Cameroon. Later Mertens (1938) described a sub- species T. montium grafi from Mongonge, on the oppo- site side of Mt. Cameroon. Klaver & Bohme (1992) re- garded it only as an aberrant form and moved it in syn- onymy with the nominate form. Based on dorsal crest shape, Perret & Mertens (1957) indicated a possible sub- species from the Manengouba Mts. but, as in 7. m. grafi, Klaver & Bohme (1992) proved the occurrence of this character to be more widespread. However, Pook & Wild (1997) mentioned differences in the courtship livery of 7: montium between populations. Differences in colouration are of importance in species recognition and may play a role in character displacement (Pook & Wild 1997; Rand Bonn zoological Bulletin 57 (2): 211-229 1961; Wagner et al. 2009b) but further studies on this as- pect are required. 7rioceros montium inhabits the submon- tane zone of Mt. Cameroon, Rumpi Hills, Manengouba Mts. area and parts of the south-western edge of the Ba- menda Highlands (Gonwouo et al. 2006). At first glance no morphological characters indicate a separation of pop- ulations. Hence, we refrain to draw any premature con- clusions at this point. Lowland-submontane clade Trioceros cristatus (Stutchbury, 1837) (Fig. 2D) A similar situation appears in 7? cristatus and distinct sub- clades are detectable within this taxon and as in the pre- ceding case, uncorrected p-distances show only compar- atively low differences between the clades (Fig. 1; Tab. 2, Appendix II). Stutchbury (1837) described T. cristatus from Gabon and since then no further subspecies have been described or taxa synonymised with 7. cristatus. Tri- oceros cristatus is widespread in the lowland to submon- tane zone from Nigeria to the Central African Republic, Gabon and the Republic of the Congo (Klaver & Bohme 1992; Pauwels & Vande weghe 2008). Furthermore, the species has been reported from Ghana and Togo (see ref- erences in Klaver & Bohme 1992) but, these localities must be regarded with caution, as they have not been con- firmed recently. In contrast to 77 montium the species 1s more widespread. A more detailed analysis of the overall distribution must be applied before any conclusions can be drawn. Submontane-montane clade Trioceros quadricornis (Tornier, 1899)-group (Figs 2E-—G), including Trioceros quadricornis eisen- trauti (Mertens, 1968) NEW RANK Molecular results require changes in the former quadri- cornis-group. Morphological distinctness (body size, shape of dorsal crest, number and size of rostral horns, lung morphology) between populations from southern parts of the Cameroon mountain chain (Mt. Kupe, Manen- gouba Mts.) and northern parts (Bamenda Highlands to Obudu Plateau in eastern Nigeria) have already been rec- ognized by BGhme & Klaver (1981). Uncorrected p-dis- tance values between the taxa quadricornis and gracilior (Tab. 2, Appendix II), indicate a very recent split and these taxa correspond to subspecies. 7. g. gracilior is known from the Bamboutos Mts, Mbulu Hills, Mt. Lefo, Mt. Oku and the Obudu Plateau, while 7) g. quadricornis is pres- ent on Manengouba Mts. and Mt. Kupe (Bohme 1975; Bohme & Klaver 1981; Gonwouo et al. 2006; Joger 1982; Klaver & Bohme 1986, 1992). ©ZFMK 216 Michael F. Bare} et al. —, Fig. 2. Cameroonian chameleons (in life): A = Trioceros oweni male (Campo region; photo by J.A.M. Wurstner). B = 7. camer- unensis (Njonji, Mt. Cameroon). C = T. montium male; specimen with an aberrant horn shown (Big Massaka, Rumpi Hills). D = T. cristatus male (Nkoelon, Campo region). E = 7. g. quadricornis male (Mt. Kupe). F = T. q. gracilior male (Mt. Lefo; photo by W. Bohme). G = T. q. eisentrauti female (Mt. Rata, Rumpi Hills). H = 7: pfefferi male (Kodmin, Bakossi Mts.). Bonn zoological Bulletin 57 (2): 211—229 ©ZFMK Chameleons of the genus Trioceros from Cameroon DG. We group the morphologically highly distinct taxon 7. eisentrauti as a distinct subspecies of 7) guadricornis (Fig. 1), a position already indicated by Pook & Wild (1997). But, while in the latter publication and in our 12S-only analysis (not shown) eisentrauti is placed as the basal sis- ter taxon to the two 7! quadricornis subspecies, accord- ing to our combined results (16S, 12S) this is not the case. Despite its morphological uniqueness (gular crest formed of flaps in eisentrauti and a gular crest formed of conical scales in other 7rioceros taxa), molecular results reveal close relationships between these three taxa with values of uncorrected p-distances within intraspecific ranges. Val- ues of uncorrected p-distances between eisentrauti and T. q. quadricornis are comparable to values between the 7. quadricornis subspecies and values are only marginally higher between eisentrauti and the subspecies T. q. gra- cilior (Tab. 2, Appendix II). However, the taxa show a dis- junct distribution with 7 g. quadricornis occurring in the Manengouba area (see above) and 7! eisentrauti being en- demic to the Rumpi Hills in western Cameroon (Gonwouo et al. 2006; Klaver & BOhme 1997). All taxa inhabit pris- tine montane habitats, 7? g. quadricornis occurring at al- titudes between 1.800—2.250 maz.s.l., 7. g. gracilior at al- titudes between 1.800—2.400 ma.s.l. and 7. eisentrauti in altitudes above 1.150 m a.s.1., respectively (Gonwouo et al. 2006; Pook & Wild 1997). We have located T. eisen- trauti on Mt. Rata in the Rumpi Hills only above 1.600 m a.s.l., hence, it is probably even more restricted in its altitudinal and overall distribution range than previously indicated. In the original description of 7. eisentrauti Mertens (1968) had already indicated relatedness to 7. quadricornis taxa according to body size and shape of the dorsal and tail crests. Bohme & Klaver (1981) emphasized the similarities of 7’ g. quadricornis and eisentrauti in comparison to 7: q. gracilior and remarked that rostral tu- bercles in eisentrauti might represent reduced rostral horns, which are present in 7: g. quadricornis (up to two pairs of rostral horns) and 7. qg. gracilior (up to three pairs of rostral horns). However, reduction of rostral horns is also known in T. gq. quadricornis and T. q. gracilior (Bohme & Klaver 1981; Mertens 1968) and BOhme & Klaver (1981) assumed that reduced horns represent a more derived character state. From the genetic point of view, we are aware that the low genetic differentiation in mitochondrial DNA might be in some cases caused by in- trogressive hybridization in the evolutionary history of two species. However, we believe that the similar overall mor- phology (body shape and size, shape of the crests) of T. quadricornis and eisentrauti also further supports our hy- pothesis of two closely related, but conspecific taxa. Due to the constant morphological differences between them and their allopatric distributions we regard the taxa quadri- cornis and eisentrauti as subspecies of a single species. Trioceros quadricornis quadricornis (Tornier, 1899) from the Manengouba area represents the nominate form Bonn zoological Bulletin 57 (2): 211-229 while the taxon eisentrauti from the Rumpi Hills is giv- en a new systematic status Trioceros quadricornis eisen- trauti (Mertens, 1968) NEW RANK. In contrast to the afore discussed species (7! montium and T. cristatus), any contact zone between these two allopatric taxa can be ex- cluded due to their highly restricted altitudinal distribu- tion. As above, low genetic differences suggest a very re- cent split presumably connected to the altitudinal range shifts of the lower-temperature forests up to the mountains after the end of the Pleistocene Ice Ages (when montane forests in the tropics expanded to the lower elevations; He- witt 2004). All three subspecific taxa of 7. guadricornis could now represent species in statu nascendi. Submontane-montane clade Trioceros wiedersheimi (Nieden, 1910)-group (Figs 3A—-G), including Trioceros perreti (Klaver & Bohme, 1992) NEW RANK Further changes are necessary within the former wieder- sheimi-group. Klaver & Bohme (1992) described the sub- species 7. w. perreti from Manengouba Mts. Molecular re- sults however reveal full species status for this taxon, as the uncorrected p-distances between 7. wiedersheimi pop- ulations from Manengouba Mts. and populations further north (Bamenda area, Tchabal Mbabo) are clearly within the interspecific range of other western Trioceros species (Tab. 2, Appendix IT). We thus herein elevate the taxon to the full species rank: Trioceros perreti (Klaver & Bohme, 1992) NEW RANK. The present distribution of T: perreti covers the Manengouba Mts. and the Bakossi Mts. (Euskirchen et al. 2000; Gonwouo et al. 2006). Regarding the former nominate 7’ w. wiedersheimi, mo- lecular and morphological results lead to recognition of two distinct clades with uncorrected p-values within in- terspecific range of this genus (Tab. 2, Appendix II). For- merly, 7’ w. wiedersheimi has been thought to occur in Cameroon north of the Manengouba Mts. (inhabited by T. perreti (Klaver & Bohme, 1992), see above). It has been found along the Cameroon mountain chain (Bamboutos Mts., Mbulu Hills, Mt. Lefo, Mt. Mbam, Mt. Oku and Mt. Tchabal Mbabo) and in eastern Nigeria (Gotel Mts., Mam- billa Plateau and Obudu Plateau), where it inhabits mon- tane savannas and grasslands between 1400 and 2450 m a.s.l. (BOhme & Nikolaus 1989; Chirio & LeBreton 2007; Dunger 1967; Gonwouo et al. 2006; Herrmann et al. 2006; Klaver & Béhme 1992). Nieden’s (1910) description of T. wiedersheimi is based on two specimens, a female from Genderogebirge (=Tchabal Mbabo) and a subadult male from the village Tsch’a (Bekom), Bamenda area. In the course of describing 7. w. perreti, Klaver & Bohme (1992) designated the female specimen as lectotype and conse- quently restricted the type locality to the Genderogebirge ©ZFMK 218 Michael F. Bare et al. A re Fig. 3. Cameroonian chameleons (in life): A = Trioceros wiedersheimi male (Mt. Tchabal Gangdaba). B = T. perreti male (Kod- min, Bakossi Mts.). C = T. serratus male (Kedjom Keku = Big Babanki, Bamenda Highlands). D = T. serratus female (Kedjom Keku = Big Babanki, Bamenda Highlands). E = Male neotype of 7. serratus (in alcohol; MNHN 2007.1494; Belo, Mt. Oku). F = Illustration of 7? serratus after Mertens (1922; “Sitidkamerun”’). G = Male neotype of 7. serratus (in life; MNHN 2007.1494, Be- lo, Mt. Oku) in situ. Bonn zoological Bulletin 57 (2): 211-229 OZFMK Chameleons of the genus Trioceros from Cameroon 219 (=Tchabal Mbabo). As one of the uncovered molecular clades contains specimens from Tchabal Mbabo, topotyp- ic material, thus this clade should correspond to 7. wieder- sheimi. The occurrence of ZT. wiedersheimi on Tchabal Gangdaba has already been assumed in the past (Chirio & LeBreton 2007; Klaver & B6hme 1992) and we can confirm its occurrence on this mountain range. Thereafter, the species is known from the northernmost parts of its former assumed distribution, while populations of the southern molecular clade from the Bamenda Highlands, Mt. Mbam, and Mt. Oku represent a distinct taxon. It is also very likely that this clade covers populations from the Mbulu Hills, Mt. Lefo and the Obudu Plateau in south- ern Nigeria as this would correspond to a bordering range from other studies (Gonwouo et al. 2006). Solely one lo- cality in direct proximity north of the Manengouba Mts. (see map in Gonwouo et al. 2006) appears uncertain, as T: perreti has been regarded as restricted to the mountain range, but the specimen was not available to us for this study. Mertens (1922) described Chamaeleon serratus from “Stidkamerun” (= South Cameroon, Fig. 3F), being most similar to T. wiedersheimi, but differing by size, promi- nence of the temporal cristae and course of the lateral cristae (Fig. 3E). The species has been later synonymized with 7: wiedersheimi by Mertens himself (1940; see be- low). Klaver & Bohme (1992) argued that 7. serratus is a synonym of 7: wiedersheimi, as the original description, especially the low number of scales on the scalloped dor- sal ridge, is not consistent with 7. perreti from Manengou- ba Mts. A comparison with the type specimen of 7 ser- ratus was not possible, as the type specimen was proba- bly destroyed during the Second World War (H. Wermuth 16.4.1979 in litt., in Klaver & Bohme 1992). Mertens (1922) rightly suggested that 7’ wiedersheimi 1s morphologically the most similar species to 7. serratus, but obviously he only compared his material with Nieden’s (1910) original description and not with the type specimen, as he only cited the original sections for com- parison. Beside the characters cited above, Mertens (1922) mentioned that no additional distinct characters like the shape of the dorsal crest, which is at the origin of the spe- cific name (Mertens 1968), are given in Nieden’s (1910) description. [Remark: Nieden (1910) stated that a dorsal crest is lacking but, a dorsal midline is formed of two rows of tubercle scales which are separated in groups of 3-4 scales in the male specimen from the Bamenda region]. Later, Mertens (1940) reported on a collection delivered by M. Kohler including chameleons from the Bamenda Highlands (four males + two females) and concluded that T. serratus is in fact a junior synonym of T. wiedershei- mi. His conclusion was based on the fact that males of the Bonn zoological Bulletin 57 (2): 211-229 new material corresponded to the “paratypoid” (= paratype) of 7 serratus, while females are consistent with the female cotype (=syntype) of 7. wiedersheimi and fi- nally he recognized that the prominence of the lateral and temporal cristae is subject to individual variation. In a sub- sequent publication on material collected by Eisentraut at Lake Oku and Lake Manengouba, Mertens (1968) con- firmed his former statement and remarked that males in T. wiedersheimi also do possess a serrated dorsal crest, while the dorsal crest is straight and simple in females. With the exception of Lake Manengouba (recognized as distinct by Klaver & Bohme 1992), all localities of the ma- terial examined by Mertens belong to the newly discov- ered southern clade. Our morphological analysis of material throughout the dis- tribution range of the former taxon T. w. wiedersheimi re- vealed that distinguishing characters chosen by Mertens (1922) are hard to assign to members of one clade, as many characters are present in members of both clades (lack of heel spur, lack of occipital lobes, etc.) separating them from other taxa. Of the three main characters given by Mertens (1922), two of them seem to be inapplicable. According to Mertens (1922): (a) T. serratus grows larg- er than 7) wiedersheimi, but four of ten males from Tch- abal Mbabo and the Gotel Mts. (= 7? wiedersheimi) pos- sess a larger body length than the largest member of the southern clade, and the largest female also belongs to T. wiedersheimi; (b) temporal cristae are distinct in 7. wieder- sheimi and indistinct in 7 serratus, but this character varies within both clades (Fig. 4), which was already men- tioned for Bamenda populations by Mertens (1940); and (c) Mertens (1922) differentiated the course of the later- al cristae (in front of the eye first running along the eye then in a weaker slope to the tip of the snout in 7’ wieder- sheimi, in contrast to an even slope in direction to the tip of the snout in 7! serratus) and this character is clearly more applicable to specimens belonging to the southern, previously unrecognized clade (Fig. 4). Nonetheless, with just a few specimens of each clade a determination on this character alone is difficult. At last, the name-giving char- acter, a serrated dorsal crest is also present in males of both clades. The number of scale rows forming the crenulation is consistent in both clades (being formed of up to three rows of scales) and the extent of crenulation along the dor- sum and base of tail also varies in both clades. Accord- ing to Mertens (1922) each cusp of the crenulation is 3 mm high and 4.5 mm long, but only in one very large spec- imen of 7. wiedersheimi a comparable size has been reached, while specimens of similar size to Mertens’ (1922) specimens possess smaller cusps in both clades. The given type locality “Stidkamerun” does not allow any direct localization of 7: serratus. Moreover, the subadult male paralectotype of 7. wiedersheimi originates from the Bamenda area (part of the southern clade) and might have OZFMK 220 Michael F. Barej et al. Fig. 4. | Heads in lateral view. Row 1-3: Trioceros wiedersheimi (from left to right): ZMB 21873 female (lectotype), ZFMK 75744 female, ZFMK 68943 male, ZFMK 75740 male, ZMB 21857 male, ZMB 74805 female, ZFMK 78714 male, MNHN 2005.2753 male, NMP6V 74112 male, ZFMK 75745 female, MHNG 1544.001 male, MHNG 1544.002 male. Row 4—6: T. serratus (from left to right): MNHN 2007.1494 male (neotype), MNHN 2007.1465 female, NMP6V 74104 male, MHNG 1365.023 female, ZSM 13.1925 subadult male, MNHN 2007.1464 male, ZFMK 5801 male, MNHN 2007.1463 male, MNHN 2007.1461 female, MHNG 1365.024 female, MHNG 1010.049 male, ZFMK 5800 female. Bonn zoological Bulletin 57 (2): 211—229 ©ZFMK Chameleons of the genus Trioceros from Cameroon 221 understandably mistaken as part of the distribution of 7. wiedersheimi. The distribution of the southern clade cov- ers the main part of former distribution of 7: wiedershei- mi and makes it more plausible to be termed “Siid- kamerun” (South Cameroon) in comparison to the Tcha- bal Mbabo area. Of Mertens (1922), most informative characters, the only reasonably useful for the recognition of T. serratus is the course of the lateral cristae. A course corresponding to Mertens’ (1922) information is present in members of the southern clade. According to Article 75 of the International Code of Zo- ological Nomenclature (ICZN 1999) a neotype is required when no name-bearing types are believed to be extant and it is necessary to define a taxon objectively. In our case, a designation of a neotype is necessary because the holo- type is lost (H. Wermuth 16.IV.1979 in litt., in Klaver & Bohme 1992; A. Schliter, herpetological curator of the SMNS, in litt. 8.I11.2010) and the taxon requires unam- biguous clarification of its taxonomic status. The type lo- cality of the taxon is now set as the collecting locality of the neotype (Article 76 in ICZN 1999). Hence, we here revalidate 7. serratus and provide a description of the neo- type at the end of this section. Submontane-montane clade Trioceros pfefferi (Tornier, 1900) (Fig. 2H) According to Townsend & Larson (2002) T: pfefferi is related to (what these authors termed) 7’ quadricornis and T: wiedersheimi, but more closely to the latter species. While the 12S-only analysis of Pook & Wild (1997) also suggested a grouping of 7: pfefferi and T: wiedersheimi, an unambiguous position of this rare taxon was not possible in our analysis with the combined 16S+12S-gene fragments (see Fig. 1). Further, because of the lack of a working 12S sequence for the sample of 7: pfefferi that we used, we used a chimera-sequence consisting of the original 12S data as published by Pook & Wild (1997; only available in the original publication, not in GenBank) and our new 16S data of another specimen. Therefore, all that can be said for now is that 7: pfefferi is more closely related to the taxa of the hornless 7. wiedersheimi-complex than to 7’ montium or T. quadricornis, which share morphological characters like horns with 7: pfefferi. It is also interesting to underline that the horned species, 7. montium, T. quadricornis and T. pfefferi, do not form a monophyletic clade, and thus, horns evolved several times in the evolutionary history of the western Jrioceros species group. Regarding the overall distribution of 7. pfefferi, this species shows a similar distribution pattern to two other species groups (a) 7. perreti—T. serratus — T: wiedershei- Bonn zoological Bulletin 57 (2): 211-229 mi and (b) T. qg. quadricornis —T. q. gracilior. Both groups show a rough distribution with one taxon in the Manen- gouba area and a second one in the Bamenda Highlands (additionally a third in the northernmost parts in the case of T. wiedersheimi). As in both these groups former al- lopatric populations have been recognized as valid taxa, the recently discovered populations of 7. pfefferi from Mbulu Hills and Ediango (see Gonwouo et al. 2006) should be compared to southern populations in future stud- les, especially as the species also inhabits submontane/montane altitudes between 1100-1800 m a.s.l. and might show a disjunct distribution, as well. Noteworthy is, that present distribution data reveal a dis- junct partitioning of montane areas in the Cameroon mountain chain with related taxa (with the exception of T. pfefferi, but see above) but apparently the highest peak (Mt. Cameroon) does not posses an endemic montane tax- on. Only the submontane 7: montium is present on Mt. Cameroon and elevations further north, but this taxon re- quires further studies to understand a potential distribu- tional separation (see above). Designation of neotype and _ redescription of Chamaeleon serratus Mertens, 1922 (now considered as a member of the genus T7rioceros sensu Tilbury & Tolley 2009) Holotype (lost). Chamaeleon serratus Mertens (1922), Zool. Anz., 54: 191. Mus. Stuttgart, Nr. 4640 (male), pro- bably destroyed during the Second World War (H. Wer- muth 16.4.1979 in litt., in Klaver & Bohme 1992; A. Schliiter, herpetological curator of the SMNS, in litt. 8.3.2010), type locality: ,,Stidkamerun™. Neotype. MNHN 2007.1494, adult male with everted hemipenes. Collected by Ivan Ineich on 9 May 2007 on a palm tree near road border of the road from Anyajua to Belo, close to Belo, Mt. Oku, Cameroon. Coordinates: N 06°10°32” E 10°21°09” (Lat.: 6.17547°, Lon.: 10.35244), 1339 m (4394 feet) a.s.]. Type locality. Belo, Mt. Oku, Cameroon Distribution. Cameroon, Nigeria Additional material examined (Appendix I) Diagnosis. 7rioceros serratus differs from all other 77i- oceros except T. wiedersheimi and T. perreti by a crest formed by the canthi rostrales merging above the snout, forming a depression between the tip of the snout and the merged crest. Moreover, it can be distinguished from 7: q. eisentrauti by the absence of gular flaps (Fig. 2D), from T. montium, T. oweni, T. q. quadricornis, T: q. gracilior ©ZFMK NO i) tO and 7: pfefferi by the absence of rostral appendages in male specimens (Figs 2A, 2C, 2E-F, 2H) and from 7: camerunensis and T. cristatus by the presence of a gular crest (Figs 2B, 2D). Trioceros serratus can be differentiated from 7. wieder- sheimi by a combination of the following characters: 7. serratus tends to stay smaller than 7? wiedersheimi and the tail length / body length ratio is lower in T. serratus (Tab. 3, Appendix II): total body length, 77 wiedersheimi (max. total length in males: 208 mm; in females: 172 mm) grow- ing larger than 7. serratus (max. total length in males 179 mm; in females 158 mm), but this observation is not sta- tistically significant (in males: p > 0.05 N,;,,=10, N;,=27; in females: p > 0.05 Ny,,=8 N;,=15); mean tail length / body length ratio is significantly higher in 7: wiedersheimi (in males: p < 0.05 Ny, =10, Nr, =27; in fe- males: p < 0.05 Ny, =8 N;,=15); the mean numbers of flank scales at midbody, although values overlap, the num- ber of flank scales at midbody is significantly higher (p< 0.01) in 7. serratus (N= 44; range: 56—76, mean: 66) than in T: wiedersheimi (N= 19; range: 57-68, mean: 62); num- ber of scales between the eye and the end of the head is significantly higher (p < 0.001) in 7) serratus (N= 44; range: 6-11, mean: 8) than in 7) wiedersheimi (N= 19; range: 5—9, mean: 7) and single scales tend to be distinct- ly larger in 7. wiedersheimi (Fig. 4); the course of the lat- eral cristae in front of the eye, decreases almost steadily from the eye to the tip of the snout in 7! serratus, while it first runs along the eye and then, from a point approx- imately at the middle of the eye, in a lower slope to the tip of the snout in 7. wiedersheimi. Trioceros serratus can be differentiated from 7: perreti by a combination of the following characters: total body length of similar size but, although largest specimens be- long to T. serratus, the mean total length is slightly high- er in 7. perreti (in males: p > 0.05 Ny, =25, Ny, =27; in females: p > 0.05 Ny, =10, N;,=15); mean tail length / body length ratio is significantly higher in 7. serratus in males (p < 0.05 N;,,=25, Ny, =27) lower, but not signif- icantly, in females (p > 0.05 N7,=10 N;,=15); mean numbers of flank scales at midbody, although values clear- ly overlap, the number of flank scales at midbody is sig- nificantly lower (p < 0.001) in 7! serratus (N= 44; range: 56-76, mean: 66) than in 7. perreti (N= 36; range: 65-86, mean: 74; in one single specimen even 93); number of scales between the eye and the end of the head is signif- icantly lower (p < 0.001) in 7. serratus (N= 44; range: 6-11, mean: 8) than in 7! perreti (N= 37; range: 9-15, mean: 11), scales behind the eyes are of similar size to flank scales in T. perreti and slightly enlarged in 7. ser- ratus; maximum length of gular crest is significantly high- er in T. serratus than in T: perreti (in males: p < 0.001 Nz) =25, Ny,=27; i females: p < 0.001 N;,=12, Bonn zoological Bulletin 57 (2): 211—229 Michael F. Barej et al. N;,=!7); length of mouth gap / distance mouth gap to tip of helmet ratio is significantly higher in males of 7: ser- ratus (p < 0.05; Np, =25, Ny, =27), while the value is not significant in females (p > 0.05; Ny, =12, Ny. =17); dor- sal part of the casque flat in 7 serratus (and T: wieder- sheimi) and convex in T. perreti (BOhme & Klaver 1992). For morphometrics see Tab. 3, Appendix II. Description of the neotype. Adult male in good condi- tion; body shape slender, laterally compressed; body length (measured from snout tip to cloaca) 83.0 mm; tail length 76.0 mm; tail base swollen and both hemipenes everted (Fig. 3F); vertical eye diameter 7.0 mm; canthus parietalis formed of 7 scales, measuring 6.8 mm (few ad- ditional slightly rough and ridged scales cranially of the parietal crest); distance snout tip to tip of helmet 24.3 mm; rostral crest merging above snout tip; rostral appendages absent; lateral and temporal crest distinct, both crests fus- ing posteriorly; occipital lobes absent; length of mouth gap 12.3 mm, mouth slightly opened; throat with fine longi- tudinal grooves between scale rows; gular crest present, formed of 24 scales, max. length of gular crest scale 1.7 mm, gular crest passing into ventral crest; lateral head scales behind eye enlarged in comparison to flank scales; lateral body scalation subhomogeneous but with inter- posed enlarged scales; dorsal crest present, serrated, formed out of 2-4 scales, diminishing in height caudally, scales of dorsal crest larger than flank scales; scales on extremities and tail (laterally and ventrally) of similar size as on body; fingers terminate in a single claw; hind feet without tarsal spurs. Colouration in live (Fig. 3G): general body colouration greenish, enlarged scales on flanks pale brown, lateral and upper side of head pale bluish, upper eye border bright yel- low, gular region green, upper most part of flanks and low- er flank pale blue. Colouration in alcohol (Fig. 3E): head and body dark grey- ish, gular crest of the same colour as body, ventral crest pale grey; sole of foot and palm of hand whitish; everted hemipenes whitish. Variation. The number of interposed enlarged scales on flanks vary in number per line and also in number of lines. Length of gular crest scales and their number is quite vari- able (Tab. 3, Appendix I). Comparing the sexes, males show a higher body length — tail ratio than females and longer scales forming the gular crest (Tab. 3, Appendix II). Females generally possess a dorsal ridge, the dorsal midline is slightly serrated in one specimen (MNHN 1998.0415) forming an indistinctly serrated dorsal crest. Colouration. The general ground coloration tends to be greenish or brownish. Enlarged scales on the flanks tend ©ZFMK Chameleons of the genus Jrioceros from Cameroon 223 toward being brown or blue and more conspicious (Figs 3C—D). A bright stripe of differing colouration runs from the anterior part of the eye (or even starts in front of the eye) and splits into rays of colour on the eyelid; one of them running backwards along the temporal crest or be- tween the temporal and lateral crest (Figs 3C, G). As in T. wiedersheimi the venter is brighter than the flanks and is pale blue or a lighter green / brown in living specimens (Fig. 3C). Blue colouration may be scattered around the mouth gap, on the top of the head, and on the flanks. The ventral line is whitish. In alcohol colourations fade away and specimens turn either pale grey-bluish (leaving some of the brighter rays on the eye visible) or turn dark and colours disappear almost completely. Distribution. The species is known to occur in Cameroon and Nigeria. Within Cameroon it inhabits montane savan- nahs in the Bamenda Highlands and Mt. Mbam, and in Nigeria it has been reported from the Obudu Plateau (Bohme 1975; Akani et al. 2001). In contrast, reports from the Gotel Mts in Nigeria (BOhme & Nikolaus 1989) can be referred to T. wiedersheimi. Genetics. The genetic comparison for the uncorrected p- distances of the combined mitochondrial 16S+12S rRNA fragments (Tab. 2, Appendix II) with the two morpholog- ically most similar species 7: wiedersheimi and T: perreti gave the following results: interspecific comparison be- tween 7) serratus and T. wiedersheimi ranged between 3.22% 4.08%, while the intraspecific variation within 7° serratus was much lower at 0.00%-—0.42% (N= 12). The interspecific difference between 7. serratus and T. perreti ranges from 4.33%—4.86%, while those between the taxa T: perreti and T. wiedersheimi reach 3.74%-4.13%. Natural history remark: Six gravid females contained the following numbers of eggs: 7, 8 (x2), 9 (x2) and 11. Size of measured eggs (N=10) ranges between 13.0—15.0 x 7.7—8.3 mm. In contrast, Angel (1940) reported ten eggs in a gravid female from the Mt. Bamboutos of 8 x 7 mm in size, which have been most probably not fully devel- oped. Acknowledgements. The study was accomplished under the conditions of the research, collecting and export permits issued by the Cameroonian Ministry of Forestry and Wildlife and Min- istry of Scientific Research and Innovation in 2007 and 2009 (Nos. 049/MINRESI/B00/C00/C10/C12; 0836/PRBS/MIN- FOF/SG/DFAPSDVEF/SC; 0613/PRBS/MINFOF/SG/DFAP/ SDVEF/SC; 0132/MINRESI/B00/C00/C10/C13; 1010/PRBS /MINFOF/SG/DFAP/SDVEF/SC). The senior author would al- so like to thank the Federal Agency for Nature Conservation (BEN) for the import permits (No. E-0997/08, No. E-03454/09). Thanks are also given to Laurent Chirio, Marcel Talla Kouete, Ivo Melle Ngwese, Samuel Wanji and Julia A. M. Wurstner for assistance in fieldwork, and/or logistics, and/or additional tis- Bonn zoological Bulletin 57 (2): 211-229 sue samples or specimens. Jean Mariaux provided the 16S se- quence for 7: pfefferi. We thank Philipp Wagner (ZFMK) for his help regarding the correct interpretation of some specific rules of the ICZN code. Thanks to Andreas Schliiter and Axel Kwet (both SMNS) for the effort to get information about A. Diehl and his collection and to Frank Tillack (ZMB) for fruitful dis- cussions. II and NLV work was supported by the Agence Na- tionale de la Recherche Biodiversity Project, Iles Forestic¢res Africaines, IFORA, 2006-2009. Field work of MFB was sup- ported by the Alexander Koenig Stiftung. VG is thankful to Ernest Vunan (SATEC, Big Babank1) for his kind assistance and hospitality. 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Bonner zoologi- sche Beitrage 56: 285-297 Werner F (1902) Prodromus einer Monographie der Chamale- onten. Zoologische Jahrbticher (Abteilung fur Systematik, Okologie und Geographie der Tiere) 15: 295-460 + Plates XV-XXVII Received: 30. VII.2010 Accepted: 18.X.2010 ©OZFMK Chameleons of the genus Trioceros from Cameroon 225 Appendix I List of examined specimens Trioceros serratus (Mertens, 1922) MNHN 2007.1494 (male neotype), Cameroon, Mt. Oku, border of the road from Anyajua to Belo (near Belo, at low altitude), date 9.V.2007, coll. I. Ineich; MHNG 964.037 (male), Cameroon, Bamenda, Kishong, 1.II.1939, coll. J.- L. Perret & R. Mertens; MHNG 1010.049-50 (2 males), Cameroon, Bafoussam, Bangwa, 1959, coll. J.-L. Perret; MHNG 1365.010 (male), Cameroon, Foumban, Mt. Nkogam, III.1969, coll. J.L. Amiet; MHNG 1365.019 (male), Cameroon, Bamiléké, Foto, X1I.1972, coll. J.L. Amiet; MHNG 1365.023-24 (2 females), Cameroon, Dschang, Foto, XI.1972, coll. J.L. Amiet; MNHN 1997.3642 (male), Cameroon, Oku village, V.1997, coll. L. Chirio; MNHN 1998.0415, Cameroon, Lake Oku, al- titude 2200 m, 7.VI.1998, coll. L. Chirio; MNHN 1998.0416-19, Cameroon, Mt. Oku, altitude 2000-2500 m, 25.V1.1998, coll. L. Chirio; MNHN 1998.0425, MNHN 1998.0429, Cameroon, Mt. Oku, altitude 2000-2500 m, 25.V1.1998, coll. L. Chirio; MNHN 2005.2781-2787, MNHN 2005.2900 (5 males + 3 females), Cameroon, Mt. Oku, Simonkuh, 10.572°E/6.234°N, altitude 2109 m, 8.VII.2002, coll. Programme CamHerp; MNHN 2005.2788 (male), Cameroon, Oku village, 19.1V.2000, al- titude 2000 m, 10.505°E/6.202°N, coll. Programme CamHerp; MNHN 2007.1461-64 (2 males + 2 females), Cameroon, Mt. Oku area, around village of Elak Oku, 6.2441°N/10.5076°E, altitude 6474 ft, 6.V.2007, coll. I. Ineich & N. Lhermitte-Vallarino; MNHN 2007.1465 (male), Cameroon, Mt. Oku_ area,’ Lake, 6.2019°N/10.4609°E, altitude 7456 ft, 8.V.2007, coll. I. Ineich & N. Lhermitte-Vallarino; NMUP6V 74104 (male), Cameroon, Kedjom Keku (= Big Babanki), Bamenda Highlands, 6°06.968’N 10°15.760’E, 1290 m a.s.l., 9.X1.2009, coll. V. Gvozdik; ZFMK 5798-5801 (2 males +2 females), Cameroon, Lake Oku, 20-30.1.1967, coll. M. Eisentraut; ZFMK 15283 (male), Cameroon, Mt. Lefo, 5.- 11.X.1974, coll. W. Bohme & W. Hartwig; ZFMK 18105- 6, ZFMK 18108 (male + 2 females), Cameroon, Mezam, Bafout, 1975-76, coll. P. Shu Mfosono; ZMB 21860 (fe- male), Cameroon, Bamenda, no date; ZMB 24909 (male), Cameroon, Bamenda, coll. Adametz; ZSM 13/1925 (male), Cameroon, Tsch’a Bekom, Bamenda District, no date, coll. Glauming. Trioceros perreti (Klaver & Bohme, 1992) MHNG 920.068-9 (male paratype + female paratype), MHNG 964.038 (female paratype), MHNG 965.054 (male paratype), MHNG 1010.052 (male holotype), MHNG 1010.053 (male paratype) Cameroon, Manengouba Mts., 1956, coll. J.-L. Perret; MHNG 1365.011 (male paratype), Cameroon, Manengouba Mts., Mwandong, 26.11.1972, Bonn zoological Bulletin 57 (2): 211-229 coll. J.L. Amiet; MHNG 1365.012-18 (5 male paratypes + 2 female paratypes), Cameroon, Manengouba Mts., 111.1973, coll. J.L. Amiet; MNHN 2007.1455-57 (3 fe- males), Cameroon, Manengouba Mts., around Mouame- na village, 9.796°E/4.984°N, altitude 4450 ft, 28.1'V.2007, coll. I. Ineich & N. Lhermitte-Vallarino; MNHN 2007.1458-60 (2 males), Cameroon, Manengouba Mts., around Mouabi village, 5.0613°N/9.8155°E, altitude 5283 ft, 29.1V.2007, coll. I. Ineich & N. Lhermitte-Val- larino; MNHN 2007.1460 (male), Cameroon, Manengou- ba Mts., border of the road going to the lakes, 5.0512°N/9.8069°E, 28.IV.2007, coll. I. Ineich & N. Lher- mitte-Vallarino; ZFMK 5802-3 (2 male paratypes), Cameroon, Manengouba Mts., Lake Manengouba, 20.XII1.1966-5.1.1967, coll. M. Eisentraut; ZFMK 55599 (male), Cameroon, Manengouba, river east of Lake Ma- nengouba, no date, F. Le Berre; ZFMK 58896-7 (2 males), Cameroon, Manengouba Mts., 1993, coll. E. Wallikewitz; ZFMK 59030 (male), Cameroon, Manengouba Mts., 1994, from pet trade; ZFMK 61836 (male), Cameroon, Manengouba area, no date, F. Le Berre; ZFMK 62571 (male), Cameroon, Manengouba area, no data, coll. un- known; ZFMK 66579 (female), Cameroon, Manengouba area, no date, from pet trade; ZFMK 66738, ZFMK 66740 (2 males), Cameroon, Manengouba area, 1998, from pet trade; ZFMK 69828, ZFMK 69830 (male + female), Cameroon, Bakossi Mts., 5.1.1998, coll. O. Euskirchen; ZFMK 69848-51 (male + 3 females), Cameroon, Bakos- si Mts., 5.1.1998, coll. A. Schmitz. Trioceros wiedersheimi (Nieden, 1910) MHNG 1544.1-2 (2 females), Cameroon, Mayo Darlé, 1941, coll. R. de Kalbermatten; MNHN 2005.2753 (fe- male), Cameroon, Fongoi Village, Tchabal Mbabo, 12.053°E/7.230°N, altitude 1900 m, 12.1.2002, coll. Pro- gramme CamHerp; NMP6V 74112 (male), Cameroon, Tchabal Gangdaba, 7°44.678’N 12°42.741’E, 1560 m a.s.l., 26.X.2009, coll. V. Gvozdik; ZFMK 47941 (male), Nigeria, Gotel Mts. Gangirwal, app. 2400 m a.s.l., 15.11.1988, coll. G. Nikolaus; ZFMK 68943 (male), Cameroon, Tchabal Mbabo, southern slopes, XI.- 5.X1I.1998, coll. George & Johnson; ZFMK 75740-3 (3 males + female), Cameroon, Mayo Kelele, app. 1600 m a.s.l., 7.1.2000, coll. H.-W. Herrmann & A. Schmitz; ZFMK 75744-6 (male + 2 females), Cameroon, Tchabal Mbabo, 25.1.2001, coll. H.-W. Herrmann & A. Schmitz; ZFMK 78714 (male), Cameroon, Tchabal Mbabo, I.2001, coll. H.-W. Herrmann & A. Schmitz; ZMB 21857, ZMB 74806 (formerly part of ZMB 21857) (male + female), Cameroon, Banjo Gebirge, no date, coll. Riggenbach; ZMB 21861, ZMB 74805 (formerly part of ZMB 21861) (male + female), Cameroon, Banjo Gebirge, no date, Riggenbach; ZMB 21873 (female lectotype), Cameroon, Genderogebirge, 1500 maz.s.1., no date, coll. Riggenbach. ©ZFMK 226 Michael F. Barej et al. Appendix II Table 1. List of voucher specimens for each species included in the present study, with their respective localities, collection num- bers and GenBank accession numbers (16S, 12S) [*sequence from Pook & Wild 1997; not in GenBank; same species but different vouchers used for 16S and 12S] [**sequence from Pook & Wild 1997, not in GenBank; same voucher used for both sequences]. Species Locality Collection number Accession number Kinyongia tavetana Trioceros oweni [E146.15] Trioceros camerunensis [E130.1] Trioceros cristatus [E130.2] Trioceros cristatus [E130.3] Trioceros cristatus [E131.1] Trioceros cristatus [E146.13] Trioceros cristatus [E150.7] Trioceros cristatus [E150.8] Trioceros cristatus [E180.2] Trioceros cristatus [E180.7] Trioceros montium [E130.4] Trioceros montium [E130.5] Trioceros montium [E131.2] Trioceros montium [E131.3] Trioceros montium [E179.18 Trioceros montium [E180.15 E188.19 E188.20 Trioceros perreti [E130.11] Trioceros montium [ ] [ | Trioceros montium [E188.18] [ ] [ ] Trioceros montium South Pare, Kilimanjaro, Tanzania Nkoelon, Campo region Njonji, Mt. Cameroon Njonji, Mt. Cameroon Njonji, Mt. Cameroon Njonji, Mt. Cameroon Amebishu, Mamfe region Amebishu, Mamfe region Nkoelon, Campo region Mofako Balue, Rump Hills Big Massaka, Rumpi Hills Mt. Cameroon Mt. Kupe Mt. Cameroon Mt. Kupe Mofako Balue, Rumpi Hills Nyasoso, Mt. Kupe Edib Hills, Bakossi Mts. Edib Hills, Bakossi Mts. Edib Hills, Bakossi Mts. Mts. Manengouba MHNG 2612.58 ZFMK 87642 MNHN 2007.0037 MNHN 2007.1447 MNHN 2007.1448 MNHN 2007.1449 ZFMK 87646 ZFMK 87647 ZFMK 87649 ZFMK 89455 MNHG 2716.39 MNHN 2007.1429 MNHN 2007.1445 MNHN 2007.1430 MNHN 2007.1446 MNHG 2716.47 MNHG 2716.41 NMPO6V 74130/2 NMPO6V 74130/1 NMP6V 74130/3 MNHN 2007.1458 AM422414 / AM422433 HQ337816 / HQ337864 HQ337798 / HQ337846 HQ337799 / HQ337847 HQ337800 / HQ337848 HQ337801 / HQ337849 HQ337802 / HQ337850 HQ337803 / HQ337851 HQ337804 / HQ337852 HQ337805 / HQ337853 HQ337806 / HQ337854 HQ337807 / HQ337855 HQ337808 / HQ337856 HQ337809 / HQ337857 HQ337810 / HQ337858 HQ337811 / HQ337859 HQ337812 / HQ337860 HQ337813 / HQ337861 HQ337814 / HQ337862 HQ337815 / HQ337863 HQ337828 / HQ337876 Trioceros perreti [E130.12] Mts. Manengouba MNHN 2007.1459 HQ337829 / HQ337877 Trioceros perreti [E131.6] Mts. Manengouba MNHN 2007.1460 HQ337830 / HQ337875 Trioceros pfefferi Afua jim Forest, 10,4°E/6,15°N MNHN 2007.1499 HQ337817 / —* Trioceros quadricornis eisentrauti Rumpi Hills voucher not collected HQ337820 / She Trioceros quadricornis eisentrauti [E178.10] Mt. Rata, Rumpi Hills ZFMK 89466 HQ337818 / HQ337866 Trioceros quadricornis eisentrauti [E178.11] Mt. Rata, Rumpi Hills MNHG 2716.40 HQ337819 / HQ337867 Trioceros quadricornis gracilior [E130.7] Oku village, Mt. Oku, Bamenda Highlands MNHN 2007.1426 HQ337821 / HQ337868 Trioceros quadricornis gracilior [E130.8] Oku village, Mt. Oku, Bamenda Highlands MNHN 2007.1423 HQ337822 / HQ337869 Trioceros quadricornis gracilior [E131.4] Oku village, Mt. Oku, Bamenda Highlands MNHN 2007.1424 HQ337823 / HQ337870 Trioceros quadricornis quadricornis [E130.9] | Mts. Manengouba MNHN 2007.1470 HQ337824 / HQ337871 Trioceros quadricornis quadricornis [E130.10] Mts. Manengouba MNHN 2007.1466 HQ337825 / HQ337872 Trioceros quadricornis quadricornis [E131.5] | Mts. Manengouba MNHN 2007.1469 HQ337826 / HQ337873 Trioceros quadricornis quadricornis [E131.8] | Mts. Manengouba MNHN 2007.1468 HQ337827 / HQ337874 Trioceros serratus [E130.15] Oku village, Mt. Oku, Bamenda Highlands MNHN 2007.1463 HQ337831 / HQ337878 Trioceros serratus [E130.16] Lake Oku, Mt. Oku, Bamenda Highlands MNHN 2007.1465 HQ337832 / HQ337879 Trioceros serratus [E130.17] (NEOTYPE) Belo, Mt. Oku MNHN 2007.1494 HQ337833 / HQ337880 Trioceros serratus [E131.7] Oku village, Mt. Oku, Bamenda Highlands MNHN 2007.1464 HQ337834 / HQ337881 Trioceros serratus [E131.16] Oku village, Mt. Oku, Bamenda Highlands MNHN 2007.1461 HQ337835 / HQ337882 Trioceros serratus [E131.17] Oku village, Mt. Oku, Bamenda Highlands MNHN 2007.1462 HQ337836 / HQ337883 Trioceros serratus [E178.2] Mt. Mbam voucher not collected HQ337837 / HQ337884 Trioceros serratus [E178.3] Mt. Mbam voucher not collected HQ337838 / HQ337885 Trioceros serratus [E178.4] Mt. Mbam voucher not collected HQ337839 / HQ337886 Trioceros serratus [E178.5] Mt. Mbam voucher not collected HQ337840 / HQ337887 Trioceros serratus [E188.16] Trioceros serratus [E189.8] Trioceros wiedersheimi [E91.6] Big Babanki, Bamenda Highlands Big Babanki, Bamenda Highlands Tchabal Mbabo Tchabal Mbabo Tchabal Gangdaba NMP6V 74104 HQ337841 / HQ337888 voucher not collected HQ337842 / HQ337889 ZFMK 75744 HQ337843 / HQ337890 voucher not collected HQ337844 / HQ337891 NMP6V 74112 HQ337845 / HQ337892 Trioceros wiedersheimi [E178.1] Trioceros wiedersheimi [E188.13] Bonn zoological Bulletin 57 (2): 211-229 ©OZFMK Chameleons of the genus Trioceros from Cameroon oy) Table 2. Uncorrected p-distances between Cameroonian Trioceros taxa based on 960 bp of the 16S + 12S rRNA gene fragments. 1 2 3 4 5 6 7 8 9 10 11 12 1 = Kinyongia tavetana AM422414/AM422433 - 2 oweni Nkoelon, Campo region [E146.15] 0.1221 - 3. camerunensis Mt. Cameroon [E130.1] 0.1230 0.0986 — 4 cristatus Big Massaka, Rumpi Hills [E180.7] 0.1138 0.0868 0.0381 — 5 cristatus Nkoelon, Campo region [E150.8] 0.1109 0.0868 0.0386 0.0086 — 6 — cristatus Amebishu, Mamfe region [E146.13] 0.1083 0.0857 0.0402 0.0095 0.0075 — 7 cristatus Amebishu, Mamfe region [E150.7] 0.1095 0.0868 0.0412 0.0106 0.0086 0.0011 — 8 cristatus Njonji, Mt. Cameroon [E130.2] O.11S1 0.0890 0.0396 0.0011 0.0096 0.0107 0.0118 = — 9 cristatus Njonji, Mt. Cameroon [E130.3] 0.1162 0.0901 0.0407 0.0021 0.0107 0.0118 0.0128 0.0011 — 10 cristatus Njonji, Mt. Cameroon [E131.1] 0.1150 0.0879 0.0391 0.0011 0.0096 0.0105 0.0116 0.0000 0.0011 — 11 cristatus Mofako Balue, Rumpi Hills [E180.2] 0.1138 0.0868 0.0381 0.0000 0.0086 0.0095 0.0106 0.0011 0.0021 0.0011 — 12. montium Edib Hills, Bakossi Mts. [E188.20] 0.1159 0.0958 0.0343 0.0365 0.0326 0.0365 0.0376 0.0380 0.0391 0.0375 0.0365 — 13 montium Edib Hills, Bakossi Mts. [E188.18] 0.1152 0.0943 0.0338 0.0360 0.0321 0.0360 0.0370 0.0375 0.0385 0.0370 0.0360 0.0000 14 montium Edib Hills, Bakossi Mts. [E188.19] 0.1152 0.0943 0.0338 0.0360 0.0321 0.0360 0.0370 0.0375 0.0385 0.0370 0.0360 0.0000 15 montium Mt. Kupe [E130.5] 0.1153 0.0944 0.0374 0.0386 0.0332 0.0375 0.0385 0.0396 0.0407 0.0396 0.0386 0.0032 16 montium Mt. Kupe [E131.3] 0.1153 0.0944 0.0374 0.0386 0.0332 0.0375 0.0385 0.0396 0.0407 0.0396 0.0386 0.0032 17 montium Nyasoso, Mt. Kupe [E180.15] 0.1152 0.0943 0.0338 0.0360 0.0321 0.0360 0.0370 0.0375 0.0385 0.0370 0.0360 0.0000 18 montium Mt. Cameroon [E130.4] 0.1153 0.0964 0.0381 0.0392 0.0364 0.0402 0.0413 0.0407 0.0418 0.0402 0.0392 0.0064 19 montium Mt. Cameroon [E131.2] 0.1153 0.0964 0.0381 0.0392 0.0364 0.0402 0.0413 0.0407 0.0418 0.0402 0.0392 0.0064 20 montium Mofako Balue, Rumpi Hills [E179.18] 0.1141 0.0923 0.0349 0.0360 0.0311 0.0349 0.0360 0.0375 0.0385 0.0370 0.0360 0.0011 21 perreti Mts. Manengouba [E131.6] 0.1194 0.0943 0.0623 0.0528 0.0525 0.0560 0.0571 0.0545 0.0556 0.0539 0.0528 0.0591 22. perreti Mts. Manengouba [E130.11] 0.1227 0.0953 0.0612 0.0517 0.0514 0.0549 0.0560 0.0534 0.0545 0.0528 0.0517 0.0580 23 perreti Mts. Manengouba [E130.12] 0.1227 0.0953 0.0612 0.0517 0.0514 0.0549 0.0560 0.0534 0.0545 0.0528 0.0517 0.0580 24 pfefferi 0.1099 0.0934 0.0627 0.0603 0.0563 0.0588 0.0575 0.0616 0.0629 0.0616 0.0603 0.0538 25 quadricornis eisentrauti Rumpi Hills 0.1058 0.0966 0.0690 0.0639 0.0614 0.0612 0.0626 0.0652 0.0665 0.0652 0.0639 0.0611 26 quadricornis eisentrauti Mt. Rata, Rumpi Hills 0.1118 0.0948 0.0638 0.0605 0.0591 0.0606 0.0617 0.0621 0.0632 0.0616 0.0605 0.0551 {E178.10] 27. quadricornis eisentrauti Mt. Rata, Rumpi Hills 0.1124 0.0957 0.0644 0.0611 0.0598 0.0611 0.0623 0.0629 0.0640 0.0622 0.0611 0.0556 (E178.11] 28 quadricornis gracilior Mt. Oku [E130.7] 0.1117 0.0911 0.0603 0.0539 0.0525 0.0539 0.0550 0.0556 0.0567 0.0550 0.0539 0.0526 29 quadricornis gracilior Mt. Oku [E130.8] 0.1117 0.0911 0.0603 0.0539 0.0525 0.0539 0.0550 0.0556 0.0567 0.0550 0.0539 0.0526 30 © quadricornis gracilior Oku village [E131.4] 0.1118 0.0922 0.0610 0.0545 0.0525 0.0545 0.0557 0.0556 0.0567 0.0556 0.0545 0.0532 31 quadricornis quadricornis Mts. Manengouba 0.1105 0.0933 0.0624 0.0560 0.0547 0.0560 0.0571 0.0577 0.0588 0.0571 0.0560 0.0548 [E130.10] 32 quadricornis quadricornis Mts. Manengouba 0.1105 0.0933 0.0624 0.0560 0.0547 0.0560 0.0571 0.0577 0.0588 0.0571 0.0560 0.0548 [E130.9] 33 quadricornis quadricornis Mts. Manengouba 0.1105 0.0933 0.0624 0.0560 0.0547 0.0560 0.0571 0.0577 0.0588 0.0571 0.0560 0.0548 {E131.5] 34 quadricornis quadricornis Mts. Manengouba 0.1108 0.0944 0.0632 0.0567 0.0546 0.0567 0.0579 0.0578 0.0589 0.0578 0.0567 0.0554 [E131.8] 35 serratus Belo, Mt. Oku [E130.17] 0.1174 0.1008 0.0651 0.0609 0.0601 0.0619 0.0620 0.0621 0.0632 0.0620 0.0609 0.0641 36 serratus Big Babanki, Bamenda Highlands 0.1174 0.0998 0.0646 0.0603 0.0601 0.0614 0.0615 0.0621 0.0632 0.0614 0.0603 0.0635 [E188.16] 37 serratus Big Babanki, Bamenda Highlands 0.1180 0.1014 0.0654 0.0611 0.0609 0.0622 0.0623 0.0629 0.0640 0.0622 0.0611 0.0634 [E189.8] 38 serratus Mt. Mbam [E178.2] 0.1163 0.1000 0.0664 0.0621 0.0612 0.0632 0.0633 0.0632 0.0643 0.0632 0.0621 0.0653 39 serratus Mt. Mbam [E178.3] 0.1165 0.0990 0.0654 0.0611 0.0602 0.0622 0.0622 0.0622 0.0633 0.0622 0.0611 0.0643 40 serratus Mt. Mbam [E178.4] 0.1163 0.0999 0.0664 0.0621 0.0612 0.0632 0.0632 0.0632 0.0643 0.0632 0.0621 0.0653 41 serratus Mt. Mbam [E178.5] 0.1151 0.0977 0.0646 0.0603 0.0601 0.0614 0.0615 0.0621 0.0632 0.0614 0.0603 0.0635 42 serratus Mt. Oku [E130.15] 0.1186 0.1022 0.0664 0.0621 0.0613 0.0632 0.0633 0.0632 0.0643 0.0632 0.0621 0.0654 43 serratus Lake Oku [E130.16] 0.1186 0.1021 0.0663 0.0621 0.0612 0.0632 0.0632 0.0632 0.0643 0.0632 0.0621 0.0653 44 serratus Oku village [E131.16] 0.1175 0.0998 0.0646 0.0604 0.0601 0.0614 0.0626 0.0622 0.0633 0.0615 0.0604 0.0636 45 serratus Oku village [E131.17] 0.1185 0.1009 0.0656 0.0614 0.0612 0.0624 0.0625 0.0632 0.0643 0.0625 0.0614 0.0646 46 serratus Oku village [E131.7] 0.1090 0.0988 0.0680 0.0645 0.0636 0.0633 0.0634 0.0656 0.0667 0.0656 0.0645 0.0670 47 wiedersheimi Tchabal Mbabo [E91.6] 0.1147 0.0923 0.0635 0.0520 0.0529 0.0533 0.0534 0.0538 0.0550 0.0531 0.0520 0.0590 48 wiedersheimi Tchabal Mbabo [E178.1] 0.1163 0.0900 0.0620 0.0508 0.0494 0.0498 0.0498 0.0526 0.0537 0.0519 0.0508 0.0565 49 wiedersheimi Tchabal Gangdaba [E188.13] 0.1146 0.0903 0.0637 0.0519 0.0527 0.0530 0.0541 0.0536 0.0547 0.0530 0.0519 0.0592 Bonn zoological Bulletin 57 (2): 211-229 ©ZFMK 228 Michael F. Bare] et al. Table 2. Continued. 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 13 = 14 0.0000 — 15 0.0032 0.0032 - 16 0.0032 0.0032 0.0000 — 17 0.0000 0.0000 0.0032 0.0032 —- 18 0.0063 0.0063 0.0075 0.0075 0.0063 — 19 0.0063 0.0063 0.0075 0.0075 0.0063 0.0000 — 20 0.0011 0.0011 0.0021 0.0021 0.0011 0.0053 0.0053 — 21 0.0582 0.0582 0.0578 0.0578 0.0582 0.0603 0.0603 0.0571 — 22 0.0571 0.0571 0.0567 0.0567 0.0571 0.0592 0.0592 0.0560 0.0032 — 23 0.0571 0.0571 0.0567 0.0567 0.0571 0.0592 0.0592 0.0560 0.0032 0.0000 — 24 0.0535 0.0535 0.0521 0.0521 0.0535 0.0574 0.0574 0.0510 0.0365 0.0378 0.0378 — 25 0.0599 0.0599 0.0587 0.0587 0.0599 0.0613 0.0613 0.0575 0.0370 0.0382 0.0382 0.0392 — 26 0.0542 0.0542 0.0536 0.0536 0.0542 0.0563 0.0563 0.0522 0.0403 0.0435 0.0435 0.0368 0.0026 — 27 0.0547 0.0547 0.0542 0.0542 0.0547 0.0568 0.0568 0.0526 0.0407 0.0439 0.0439 0.0369 0.0027 0.0000 — 28 0.0518 0.0518 0.0513 0.0513 0.0518 0.0539 0.0539 0.0497 0.0433 0.0443 0.0443 0.0366 0.0102 0.0106 0.0107 — 29 0.0518 0.0518 0.0513 0.0513 0.0518 0.0539 0.0539 0.0497 0.0433 0.0443 0.0443 0.0366 0.0102 0.0106 0.0107 0.0000 — 30 0.0524 0.0524 0.0513 0.0513 0.0524 0.0545 0.0545 0.0503 0.0438 0.0448 0.0448 0.0366 0.0103 0.0107 0.0108 0.0000 0.0000 — 31 0.0539 0.0539 0.0535 0.0535 0.0539 0.0560 0.0560 0.0518 0.0422 0.0432 0.0432 0.0367 0.0051 0.0064 0.0064 0.0063 0.0063 0.0064 — 32 0.0539 0.0539 0.0535 0.0535 0.0539 0.0560 0.0560 0.0518 0.0422 0.0432 0.0432 0.0367 0.0051 0.0064 0.0064 0.0063 0.0063 0.0064 0.0000 33 0.0539 0.0539 0.0535 0.0535 0.0539 0.0560 0.0560 0.0518 0.0422 0.0432 0.0432 0.0367 0.0051 0.0064 0.0064 0.0063 0.0063 0.0064 0.0000 34 0.0546 0.0546 0.0535 0.0535 0.0546 0.0567 0.0567 0.0525 0.0427 0.0438 0.0438 0.0367 0.0051 0.0064 0.0065 0.0064 0.0064 0.0064 0.0000 35 0.0630 0.0630 0.0622 0.0622 0.0630 0.0652 0.0652 0.0620 0.0458 0.0447 0.0447 0.0434 0.0435 0.0480 0.0487 0.0426 0.0426 0.0427 0.0437 36 0.0625 0.0625 0.0622 0.0622 0.0625 0.0646 0.0646 0.0614 0.0454 0.0444 0.0444 0.0434 0.0435 0.0478 0.0483 0.0423 0.0423 0.0428 0.0433 37 0.0634 0.0634 0.0631 0.0631 0.0634 0.0655 0.0655 0.0623 0.0461 0.0451 0.0451 0.0437 0.0442 0.0484 0.0489 0.0428 0.0428 0.0433 0.0439 38 0.0643 0.0643 0.0632 0.0632 0.0643 0.0664 0.0664 0.0632 0.0459 0.0448 0.0448 0.0434 0.0435 0.0471 0.0477 0.0416 0.0416 0.0417 0.0427 39 0.0633 0.0633 0.0622 0.0622 0.0633 0.0654 0.0654 0.0622 0.0449 0.0438 0.0438 0.0434 0.0435 0.0460 0.0467 0.0406 0.0406 0.0407 0.0417 40 0.0643 0.0643 0.0632 0.0632 0.0643 0.0664 0.0664 0.0632 0.0459 0.0448 0.0448 0.0434 0.0435 0.0471 0.0477 0.0416 0.0416 0.0417 0.0427 41 0.0625 0.0625 0.0622 0.0622 0.0625 0.0646 0.0646 0.0614 0.0444 0.0433 0.0433 0.0447 0.0448 0.0478 0.0482 0.0423 0.0423 0.0428 0.0433 42 0.0643 0.0643 0.0633 0.0633 0.0643 0.0665 0.0665 0.0633 0.0470 0.0460 0.0460 0.0447 0.0447 0.0493 0.0500 0.0438 0.0438 0.0439 0.0449 43 0.0643 0.0643 0.0632 0.0632 0.0643 0.0664 0.0664 0.0632 0.0470 0.0459 0.0459 0.0447 0.0447 0.0492 0.0499 0.0438 0.0438 0.0438 0.0449 44 0.0626 0.0626 0.0623 0.0623. 0.0626 0.0647 0.0647 0.0615 0.0455 0.0444 0.0444 0.0446 0.0434 0.0479 0.0483 0.0423 0.0423 0.0428 0.0434 45 0.0636 0.0636 0.0633 0.0633 0.0636 0.0657 0.0657 0.0625 0.0465 0.0454 0.0454 0.0447 0.0447 0.0488 0.0493 0.0433 0.0433 0.0439 0.0444 46 0.0658 0.0658 0.0648 0.0648 0.0658 0.0669 0.0669 0.0647 0.0486 0.0475 0.0475 0.0447 0.0447 0.0475 0.0483 0.0419 0.0419 0.0419 0.0429 47 0.0591 0.0591 0.0587 0.0587 0.0591 0.0612 0.0612 0.0568 0.0374 0.0363 0.0363 0.0337 0.0318 0.0342 0.0345 0.0329 0.0329 0.0333 0.0329 48 0.0565 0.0565 0.0562 0.0562 0.0565 0.0609 0.0609 0.0543 0.0388 0.0377 0.0377 0.0340 0.0321 0.0345 0.0348 0.0333 0.0333 0.0337 0.0332 49 0.0583 0.0583 0.0580 0.0580 0.0583 0.0604 0.0604 0.0562 0.0413 0.0403 0.0403 0.0389 0.0341 0.0352 0.0354 0.0350 0.0350 0.0354 0.0360 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 32 - 33 0.0000 — 34 0.0000 0.0000 — 35 0.0437 0.0437 0.0438 = — 36 0.0433 0.0433 0.0439 0.0000 — 37 0.0439 0.0439 0.0445 0.0000 0.0000 — 38 0.0427 0.0427 0.0428 0.0011 0.0011 0.0011 — 39 0.0417 0.0417 0.0417 0.0011 0.0011 0.0011 0.0000 — 40 0.0427 0.0427 0.0428 0.0011 0.0011 0.0011 0.0000 0.0000 — 41 0.0433 0.0433 0.0439 0.0021 0.0021 0.0021 0.0011 0.0011 0.0011 — 42 0.0449 0.0449 0.0450 0.0011 0.0011 0.0011 0.0021 0.0021 0.0021 0.0032 — 43 0.0449 0.0449 0.0449 0.0011 0.0011 0.0011 0.0021 0.0021 0.0021 0.0032 0.0000 — 44 0.0434 0.0434 0.0439 0.0021 0.0021 0.0022 0.0032 0.0032 0.0032 0.0042 0.0011 0.0011 — 45 0.0444 = 0.0444 0.0449 0.0011 0.0011 0.0011 0.0021 0.0022 0.0021 0.0032 0.0000 0.0000 0.0011 — 46 0.0429 0.0429 0.0430 0.0011 0.0011 0.0012 0.0023 0.0023 0.0023 0.0034 0.0000 0.0000 0.0011 0.0000 — 47 0.0329 0.0329 0.0334 0.0344 0.0341 0.0340 0.0333 0.0322 0.0333 0.0341 0.0357 0.0356 0.0352 0.0352 0.0378 — 48 0.0332 0.0332 0.0337 0.0369 0.0365 0.0365 0.0359 0.0348 0.0359 0.0365 0.0382 0.0381 0.0377 0.0376 0.0403 0.0045 — 49 0.0360 0.0360 0.0365 0.0375 0.0371 0.0377 0.0365 0.0355 0.0365 0.0371 0.0387 0.0386 0.0372 0.0382 0.0408 0.0068 0.0066 — Bonn zoological Bulletin 57 (2): 211-229 ©ZFMK Chameleons of the genus Zrioceros from Cameroon £00 £0°0 800 £0°0 £0°0 £00 10°0 10°0 900 VV Gil vO 81 Lal vl cl Tl Ve L0 Gla AE € ‘Aap’s 4 6L0 LS'0 Clit LS'0 veo cs'0 910 910 LL0 L9 8 Caesecls Gull vrt vil GL FENG SOE WLS OVE GEG uvoul = L3'0 £9°0 Lol £9°0 8e0 8S°0 810 810 93°0 OL I] 6T 6¢I SHI Sol Vel €6 8V~ SL “0i0L 30°C6 Te xeul S 0L0 cs'0 10'T cS'0 670 Lv'0 vam) STO £9°0 09 oF 9i0) 2016 o'8 OCI 88 Ss «LT 8I go OSP Oss 61 urur LI LI LI LI LI LI LI LI I LI LI LI LI LI LI LI LI LI LI sa LI LI N (So[VUlaJ) SMJDsAAS SOADIOIT, £00 c0°0 80°0 c0'0 c0'0 c0'0 10:0 100 80°0 ve Le! so. 6 UTT 80 60 OT GO) LA 50 86 L9 € “ASp's 820 890 60'1 850 €€0 cs'0 L10 910 98°0 £9 L CAE Sieles 265 GS -8l= Vile “6K? 69 OL9 68L VE6I uvoul 80 19°0 Ic 19°0 9¢°0 €S'0 810 L10 00'T 89 6 Sc OST 6F€I CO Sl Svs “Se EL 10:98 “ONES €C xXeUl IL'0 cs'0 L60 cs'0 0c0 870 910 910 9L'0 LS Si OW Orl= 9301 ES viOl= 9:9. 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Se MUG TOSS OES SR OKED Il ura SC SC ST SC Sc SC Sc SC SC SC SC SG SG SC SC SC Sc Sc Sc Sc SC SC N (sajeul) Yassad Sosad01N], JHOW/D OW/daA MA/GH OW/da 3HIS/MH 3HIS/DOW T4/T9d TH/TEL TH/1L WSH AHS TID TH TH JHOW OW MH IHS) Ga IL Wa OO ‘ones yowyoy Jo dy 0} des yynow / des ynow = iOW/OW ‘ones ypIM peoy / Jojowerp sA0 = MH/Ga_ ‘ones des yynour/ 1ajowerp sA0 = OW/C_ ‘ones yourpay Jo dy oj dy nous / (PIM peoy = IHIS/MH “ones jourjoy Jo dy o} dy ynous / des yynour = IHIS/OW ‘ones ySua] Apog / yISue] wWueoIO} Tq/ToJ ‘ones ySuo] Apog / yBuo] eign = Tq/THL “ones yISuI] Apog / yjsus] [1e} = Tq/TL ‘Apogprur ye sayeos yuey = WSJ ‘peoy Jo puoa pure oX9 usamyaq a]kOs JO JoquINU = FHS ‘}Saso Jens Jo YSU] = TH ‘yISuo] wue o10f = TOI ‘ySug] viqn = TIL owyoy Jo dy 0} de8 yynow sourjsip = IO ‘des now = yyy ‘yIpIM peoy = MH “owyoy Jo dy 0} dy ynous sourysip = HIS “Jo}ourerp dA0 [eoI}IOA = QA ‘[!e} JO dy 0} vovojo WOT, YISUD] [Ie] = TL “VOVO]D 0} nous Jo dy Woy YSU] Apog = Tg ‘jSo1O Ie[NS BUIULIO} sapeos Jo JoquINU = DN (‘Aop's) UOeIASp plepuejs pue (URoIT) SonjeA UROW “(xeU) WINUITX -PUL “(UTUT) WINUITUTU “(N{) OZIS o[dues oI UDAID “SNJDAIAS ‘J, Pu IUMaYssapaIM “J ‘Yadssad SOAJIOLAT, B[CUIDJ PUL [LUI W]Npe JO (UIW) SoINseoU [eoIso;oYydiow jo Arewung *¢ sq Bonn zoological Bulletin 57 (2): 211-229 a i | a “ _ ang ae rod ci nation Bonn zoological Bulletin Volume 57 Bonn, November 2010 Another new Cophixalus species (Amphibia: Anura: Microhylidae) from western New Guinea Rainer Gtinther Museum ftir Naturkunde Berlin, Leibniz Institute for Research on Evolution and Biodiversity at the Humboldt-University, Invalidenstr. 43, D-10115 Berlin, Germany Abstract. Based on external morphological, anatomical, bioacoustic, and molecular traits, a new species in the micro- hylid genus Cophixalus is described. The new species was discovered in the Fakfak Mountains, northwestern corner of the Bomberai Peninsula, Papua Province, Indonesia. The new taxon is most closely related to the sympatric Cophixalus tetzlaffi. It differs from that species in several morphological traits, but primarily by its advertisement call: the new species utters a single peeping note with a mean duration of less than 200 milliseconds, whereas the advertisement call of C. tetz- laffi consists of three to four notes, with a mean note duration of more than 400 milliseconds. Molecular data (mitochon- drial 16S rRNA) support the determination of the specific distinctness of the new species. Key words. Anura, Asterophryinae, Cophixalus, new species, New Guinea. INTRODUCTION Fifty-one species in the microhylid frog genus Cophixalus are known at present (Frost 2010). Of these, 14 occur in north-eastern Australia, 30 in Papua New Guinea, three are known only from the Papua Province of Indonesia, three are recorded from both Papua New Guinea and the Papua Province of Indonesia, and one species seems to be endemic to the Island of Halmahera, located about 300 km west of the western tip of New Guinea. Although many new Cophixalus species are expected to be described al- so from western New Guinea, the distribution centre of the genus seems to be clearly in eastern New Guinea and north-eastern Australia. Cophixalus montanus has been known since 1895 from Halmahera, and the detection of three new species in the western part of New Guinea (on Yapen Island, on the Wandammen Peninsula, and on the Bomberai Peninsula) came as a surprise (Gunther 2003, 2006). Here I describe another new species from the Fakfak Mountains on the Bomberai Peninsula (located on the “throat” of the Vogelkop) found during an expedition in September 2008. Moreover, a population of Cophixalus tridactylus and one specimen of a second undescribed species was observed there. Consequently, at least four Cophixalus species occur syntopically in the middle and higher elevations (400-1000 m above sea level, a.s.l.) of the Fakfak Mountains. Bonn zoological Bulletin 57 (2): 231-240 MATERIAL AND METHODS Most frogs were collected at night after locating them by their advertisement calls. Some specimens were pho- tographed in life the next day and all specimens were anaesthetized with chlorobutanol and subsequently fixed in 2 % formalin. Tissue probes from thigh muscle were taken from some frogs and stored in about 96 % ethanol to enable DNA sequencing, before fixing the animals in formalin. All specimens were transferred to 75 % ethanol later in the Berlin museum. One specimen was cleared and stained as an osteological preparation according to a method modified from Dingerkus & Uhler (1977). The following measurements were taken with a digital cal- liper (> 10 mm) or with a binocular dissecting microscope fitted with an ocular micrometer (< 10 mm) to the near- est 0.1 mm: SUL — snout-urostyle length: from tip of snout to distal tip of urostyle-bone; SUL is about one to two mm shorter than the snout-vent length (SVL). As the measurement error is higher in the latter, I prefer to use the former. In general, both measurements are more or less identical and are used interchangeably in this paper. TL -—tibia length: external distance between knee and an- kle; TaL — length of tarsus: external distance, tarsal and ankle joints held at a right angle; ©ZFMK 232 Rainer Gtinther 10° I i 3] Fig. 1. cality (1). Map of the western part of New Guinea with type lo- T4L — length of fourth toe: from tip of toe to proximal end of inner metatarsal tubercle; T4D — transverse diameter of disc of fourth toe; F3L — length of third toe; F3D — transverse diameter of disc of third finger; F1D — transverse diameter of disc of first finger; T1D — transverse diameter of disc of first toe; HL — head length: from tip of snout to posterior margin of tympanum; HW — head width, taken in the region of the tympana; SL — snout length: from an imaginary line that connects the centres of eyes to tip of snout; END — distance from anterior corner of orbital opening to centre of naris; IND — internarial distance between centres of nares; ED — eye diameter: from anterior to posterior corner of orbital opening; TyD — horizontal diameter of tympanum. Advertisement calls were recorded under natural condi- tions with a Sony Digital Audio Tape (DAT) Walkman TCD-D 100 and a Sennheiser microphone MKE 300 and analysed with Avisoft-SAS Lab Pro software. All speci- mens are currently stored in the Museum fiir Naturkunde Berlin (ZMB) and bear registration numbers of that insti- tution. Part of the type series will later be transferred to the Museum Zoologicum Bogoriense (MZB). RESULTS AND DISCUSSION Cophixalus monosyllabus sp. n. Holotype. ZMB 74993 (field number, FN: RG 7888) ; adult male collected by R. Giinther and A. Piahar 6 km Bonn zoological Bulletin 57 (2): 231-240 Fig. 2. Holotype of Cophixalus monosyllabus sp. n. head in lateral view (above); head in dorsal view (below). direct line NNE of Fakfak town, near the Fakfak-Kokas road, Bomberai Peninsula (neck of Vogelkop), Papua Province, Indonesia, 2°53’S and 132°18'E, elevation 500 ma.s.l., 9 September 2008 (Fig. 1). Paratypes. ZMB 74994 (FN: RG 7889), ZMB 74995 (FN: RG 7890), ZMB 74996 (FN: RG 7912), ZMB 74997 (FN: RG 7915), ZMB 74998 (FN: RG 7916), ZMB 74999 (FN: RG 7926), ZMB 75000 (FN: RG 7927), ZMB 75001 (FN: RG 7951), ZMB 75002 (FN: RG 7952). ZMB 74997 is now an osteological preparation. All nine paratypes are males. They were collected from 9 to 14 September 2008 along the Fakfak-Kokas road in the southern part of the Fakfak Mountains, at elevations of from 400 to 700 m a.s.l. Collectors were R. Giinther, M. Kapisa, and A. and F. Piahar. ©ZFMK New Cophixalus from western New Guinea 233 Table 1. Body measurements and body ratios of the type series of Cophixalus monosyllabus sp. n. ZMB-No are the inventory numbers of the Museum ftir Naturkunde Berlin, FN are the field numbers of the author, SD indicates the standard deviation. ZMB 74993 is the holotype; ZMB 74997 is now an osteological preparation. All specimens are adult males. All measurements are in mm; abbreviations are explained in “Material and methods”. ZMB-No 74993 74994 74995 74996 74997 74998 74999 75000 75001 75002 mean SD FN 7888 7889 7890 7912 PONS OMG, 7926 7927 7951 7952 22.9 1.04 SUL 22.8 DBolk 24.1 22.5 22.4 23.6 20.6 22.8 24.3 23.0 WE 10.9 ey) IEA ele Wes} ES: 10.1 INS 7/ 12.3 USF TaL 7.4 U2 Yell 6.8 6.9 eZ UP 7.6 7.3 7) T4L 11.5 11.6 i EG 11.2 11.1 10.5 LE 12.5 10.9 T4D 1.3 1E2'5 1.4 1.0 163 3 1.0 LED: 13'S 1:25 TID 0.7 0.65 OTS “OS 0.7 0.6 0.45 0.7 0.6 0.6 SIE 6.0 6.4 6.8 6.4 6.1 6.5 5)5) 59 6.8 6.0 F3D 1.4 1.4 Ned 1.45 es) 1.3 1.25 1S) 1) 1.4 FID 0.6 0.45 0.50 0.50 0.5 0.4 0.45 0.5 0.5 0.5 HL 7.5 V2) 8.2 Toll es) 8.0 6.8 Wess ES ie HW 8.5 9.0 9.1 9.0 95 9.8 8.5 9.0 9.6 9.6 SL 3.2 3.3 3.6 33 3.5 3.4 32 33 Shep) 3.6 END 2.2 2.1 2.5 2.0 2.1 De 2.0 2.1 DD 2.0 IND D3} DES) 7) D3 DED 2.4 22 2S 2.5 DED ED 2.8 2.9 3.1 DRS 2.9 2.8 Dall 2.8 3.0 3.0 TyD 1.2 1.0 12 1.0 1.0 1.1 1.0 0.9 1.0 1.0 TL/SUL 0.48 0.51 0.47 0.51 0.50 0.49 0.49 0.51 0.51 0.51 0.50 0.051 TaL/SUL 0.32 0.31 0.29 0.30 0.31 0.31 0.35 0.33 0.30 0.32 0.31 0.017 T4L/SUL 0.50 0.50 0.49 0.52 0.50 0.47 0.51 0.49 0.51 0.47 0.50 0.016 F3L/SUL 0.26 0.28 0.28 0.28 0.27 0.28 0.27 0.26 0.28 0.26 0.27 0.009 F3D/SUL 0.061 0.060 0.071 0.064 0.067 0.055 0.061 0.066 0.062 0.061 0.063 0.004 FID/SUL 0.026 0.019 0.021 0.022 0.031 0.017 0.022 0.022 0.021 0.022 0.021 0.002 T4D/SUL 0.057 0.054 0.058 0.044 0.058 0.055 0.049 0.053 0.056 0.054 0.054 0.004 TID/SUL 0.031 0.028 0.031 0.022 0.031 0.025 0.022 0.031 0.025 0.026 0.027 0.004 HL/SUL 0.33 0.31 0.34 0.32 0.33 0.34 = 0.33 0.32 0.31 0.33 0.33 0.011 HW/SUL 0.37 0.39 0.38 0.40 0.42 0.42 041 0.39 0.40 0.42 0.40 0.018 HL/HW 0.88 0.80 0.90 0.79 0.77 0.82 0.80 0.81 0.78 0.78 0.81 0.043 END/IND 0.96 0.84 0.84 0.87 0.95 0:92) 0:91 0.84 0.88 0.89 0.89 0.044 ED/SUL 0.123 0.125 0.129 0.111 O29 OMS On3it 0.123 0.120 0.130 0.124 0.006 TyD/ED 0.43 0.34 0.39 0.40 0.34 039.037 0.32 0.33 0.33 0.36 0.037 SL/SUL 0.140 0.143 0.149 0.147 0.156 0.144 0.155 0.145 0.141 0.156 0.148 0.006 Bonn zoological Bulletin 57 (2): 231-240 ©ZFMK 234 Rainer Giinther Fig. 3. Holotype of Cophixalus monosyllabus sp. n. ventral view of right hand (left); ventral view of right foot (right). Diagnosis. With a snout-urostyle length of from 20.6 to 24.3 mm in ten adult males, the new species belongs to the middle-sized species of the genus. It is obviously a sis- ter species of the sympatric Cophixalus tetzlaffi and dif- fers from all other species in the same characters as the latter. The new species differs from C. fefzlaffi, among others, by its larger body size, its wider finger and toe discs, and its advertisement call which consists of only one peeping syllable (note) that lasts, on average, 196 millisec- onds (ms). In contrast, the advertisement call of C. tetzlaf- fi consists of three to four peeping notes, with a mean note length of more than 400 ms. Description of the holotype. For measurements see Table 1. Head broader than long (HL/HW ratio 0.88), canthus rostralis roundish; loreal region straight; snout slightly pro- truding in profile (Fig. 2, above) and rounded in dorsal view (Fig. 2, below); horizontal eye diameter greater than eye-naris distance; borders of tympanum scarcely visible, its size less than half of the eye diameter (TyD/ED 0.43), no supratympanic fold; internarial distance slightly greater than distance between eye and naris (END/IND 0.96); tongue large, posteriorly broadened and without posterior notch, its posterior and lateral margins free; a strongly serrated fold present in front of the pharynx; long Fig. 4. Bonn zoological Bulletin 57 (2): 231—240 Dorsolateral view of a more brownish coloured paratype of Cophixalus monosyllabus sp. n. (ZMB 74995). ©ZFMK New Cophixalus from western New Guinea NO Ww Nn Fig. 5. Dorsolateral view of a more greyish coloured paratype of Cophixalus monosyllabus sp. n. (ZMB 74999). Fig. 6. Ventral view of a paratype of Cophixalus monosylla- bus sp. n. (ZMB 74995). Bonn zoological Bulletin 57 (2): 231-240 slits on both sides of the tongue are entrances to a subgu- lar vocal sac. Legs moderately long, no webs between fin- gers or toes (Fig. 3); tips of fingers wider than tips of toes, first finger much smaller than other fingers, its tip only scarcely wider than the penultimate phalanx; relative length of fingers 3>2=4>1; third toe clearly longer than the fifth, tip of first toe slightly smaller than tip of the fifth toe, tips of remaining toes clearly wider than that of first and fifth toe; all finger and toe tips with terminal grooves; relative length of toes 4>3>5>2>1, all subarticular tuber- cles as well as metatarsal and metacarpal tubercles not or only scarcely developed. With exception of some tuber- cles on flanks, all dorsal, lateral, and ventral surfaces smooth. Dorsum light brown and clearly demarcated against dark brown upper flanks, dorsal surfaces of legs non-uniform brown, chevron-shaped mark in scapular region, dorsal surface of snout lighter than remaining body; dorsal sur- faces of fingers and toes with yellowish, light brown, and dark brown pattern; lateral and dorsolateral flanks with longitudinal rows of blackish spots, a conspicuous black- ish spot present also above insertion of foreleg and behind eye; loreal region, tip of snout, and region below and be- hind eye and underneath tympanum blackish (black face mask); ventral surface of forelegs yellowish with irregu- OZFMK 236 Rainer Gtinther kHz NO & OOO 0.05 0.10 Fig. 7. mV 35 ] | 304 Ls —— — eewe at 251 erews wae vows rar 20 otosst 9 kHz Ol a 2 Ss Ay On Oi mee L ane. Fig. 8. | Power spectrum of an advertisement call of Cophixa- lus monosyllabus sp. n. Bonn zoological Bulletin 57 (2): 231—240 SSAA, ES GSS FRR ARE RRS. GR RGR SRS. Ont 0.20 Ss Wave form (above) and spectrogram (below) of an advertisement call of Cophixalus monosyllabus sp. n.. lar dark brown flecks, ventral surface of hind legs also yel- lowish with brown flecks but the latter less intense than on forelegs (ventral skin and muscle tissue of the right thigh was removed for biochemical studies); belly, chest and throat yellowish with brown pigmentation, pigmen- tation most intense on throat and chest and least intense on abdomen; region around anal opening blackish and re- gion from behind eye, through tympanum, and up to up- per arm whitish. Variation in the type series: Mensural variation for the type series is shown in Table 1. The basic colour and colour pattern elements of all preserved types are fairly uniform and very much resemble those of the holotype. Charac- teristic elements are a light brown dorsum, which is dif- ferentiated from dark brown upper flanks, a dark brown and irregularly pronounced interocular band, a dark chevron or W-shaped mark in the scapular region, a dark face mask, a blackish spot behind eye and above inser- tion of fore leg, a blackish throat which fades posterior- ly into a diffuse dark brown reticulum, and a pale dorsal surface of the snout which is the palest part of all the dor- sal surfaces. Only one specimen (ZMB 74996) has a whitish middorsal line from snout tip to anal opening and which continues on to the posterior thighs. The basic colour in life varies from créme or grey to light brown. Dorsum rather uniform brownish (Fig. 4) or grey (Fig. 5); conspicuous is a blackish or dark brown chevron or W-shaped mark in the scapular region, an irregular dark brown interocular band between posterior parts of eyes, and a créme or light brown coloured dorsal part of snout. Lower flanks are mostly lighter than the remaining later- al areas (Fig. 4). Upper flanks may be of nearly the same ©ZFMK New Cophixalus from western New Guinea Fig. 9. brown or grey colour as on the dorsum. Conspicuous are a blackish spot behind the eye, another blackish spot above the foreleg insertion, and some blackish spots at the bor- der between the dorsum and flanks. It is notable that the dorsum in all preserved specimens is clearly lighter than the upper flanks, whereas the dorsum and upper flanks in most living specimens differed in colour only slightly. Lo- real region in all specimens entirely or predominantly black. In some specimens this black area continues to be- low the eye and extends up to the upper arm, in others this black area ends below the eye. The inner margin of the “upper eyelid” is whitish in most specimens, this colour merging in a broad and light postocular band. While there are no or only a few tubercles in the preserved specimens, most living specimens exhibited tubercles on the flanks and extremities (rarely on the dorsum). Many of these tubercles have a blackish base and a orange-red cap. Most, and the largest, tubercles are arranged in dor- solateral rows. The yellowish spot posterior of the chevron sign in ZMB 74999 (Fig. 5), which faded to a white spot in preservative, is obviously an exception. Or- ange-red areas were also found on the forelegs of some specimens. The fine whitish middorsal line in the living Bonn zoological Bulletin 57 (2): 231-240 tO Ww — Habitat of Cophixalus monosyllabus sp. n. in the Fakfak Mountains on the Bomberai Peninsula, 700 m a.s.1. specimen in Fig. 5 disappeared completely in fixative. Dorsal sides of legs similarly coloured as other dorsal and dorsolateral body parts. Ventral sides of forearms créme- coloured, its anterior and posterior part covered with ir- regular dark spots. Throat and chest in all specimens dark- er than on the remaining ventral surfaces. These dark ven- tral areas are solidly or discontinuously black or dark brown. Abdomen and ventral sides of hind legs show greater light areas covered by a more or less dense retic- ulum of grey-brown. Weakest pigmentation was common- ly on the posterior abdomen (Fig. 6). Iris yellow-red and nerved by a dense net of blackish lines. Osteology. One cartilage-bone preparation (ZMB 74997) did not show remarkable differences from that of Cophix- alus tetzlaffi (see Giinther 2003). Vocalisation. Most calling activities were recorded dur- ing rain and damp weather from dusk to 9 p.m. All calls were recorded at temperatures of approximately 21°C. Calls are uttered in series lasting several minutes. The shortest time between two successive calls was about 3 s. Each call consists of a single unpulsed and finely tuned note (Fig. 7). Fifty-six calls (notes) from two males had ©ZFMK 238 Rainer Gtinther ~ Fig. 10. An undescribed Cophixalus species from the Fakfak Mountains, with a 16.9 mm snout-urostyle-length, which at first glance resembles Cophixalus misimae recently described by Richards & Oliver (2007) from Misima Island, Louisiade Archipela- go, Papua New Guinea. a mean length of 196 ms, with a minimum of 173 ms and a maximum of 224 ms. Most notes start with a sharp in- crease in amplitude, and the sound volume may remain constant during the entire note but may also change, with the greatest sound volume mostly near the end. The end of the note occurs more gradually and its exact cessation is fairly difficult to identify (Fig. 7). The dominant fre- quency 1s approximately 2.8 kHz (Fig. 8), the fundamen- tal frequency is approximately 1.4 kHz, and the first (and most pronounced) upper harmonic band is at about 4.2 kHz. Distribution. The new species lives on slopes and in val- leys of the southern part of the Fakfak Mountains. We found it along the Fakfak town-Kokas road at elevations of from 250 to 700 m a.s.l. Whether it also occurs in the northern part of the Fakfak Mountains remains to be de- termined. Habitat and habits. Cophixalus monosyllabus sp. n. lives mostly in the understory (bushes, young trees, and herbs) of taller trees but also in shrubbery without a canopy cov- er (Fig. 9). The frogs perched mainly on or between liv- ing or dead leaves at heights of from one to three meters above the ground. The species is common: we heard sev- eral hundred males during a walk of three kilometres along the Fakfak-Kokas road. Some males called at distances of Bonn zoological Bulletin 57 (2): 231—240 only two m from one another. At favoured places about ten males could be heard calling from one point. For bio- geographical reasons it seems worthwhile to mention that at elevations of between 500 and 700 m a.s.l., C. mono- syllabus sp. n. occurs syntopically with C. tetzlaffi, C. tri- dactylus, and another obviously new Cophixalus species Cophixalus balbus ZMB 62594 (RG 7434) Cophixalus balbus ZMB 62596 (RG 7487) Cophixalus balbus ZMB 62597 (RG 7502) Cophixalus tetzlaffi ZMB 62598 (RG 7144) Cophixalus tetzlaffi ZMB 74988 (RG 7839) Cophixalus monosyllabus ZMB 74994 (RG 7889) Cophixalus monosyllabus ZMB 74997 (RG 7915) Fig. 11. Bayesian inference phylogram of 16S rRNA. Numbers on branches denote posterior probabilities. OZFMK New Cophixalus from western New Guinea 239 (Fig. 10). Ecological differences between these four species are the following: C. monosyllabus sp. n. occurs at from 250 to 700 m a.s.I. and its calling sites are at be- tween one and three metres above ground; C. tetzlaffi oc- curs at from 400 to 900 m a.s.l. (top of the mountains) and its calling sites are on structures up to one m above the ground; C. tridactylus was found at from 500 to 900 m a.s.l. and its calling sites are on the ground; and the ob- viously new species was found at 860 m a.s.]. in humus soil below the ground surface. Etymology. The Latin word “monosyllabus” is derived from the Greek composite adjective “monosyllabos” meaning one syllable or monosyllabic, and refers to the advertisement call of the new species which consists of only one uniform note. I dedicate this new species to my dear colleague of many years, Prof. Dr. Wolfgang Bohme, to acknowledge his extraordinary contributions to herpetological science and on the occasion of his re- tirement from official service, although it is well known that Wolfgang is by no means monosyllabic but rather is very eloquent. Molecular evidence. According to B. Stelbrink and T. von Rintelen (pers. comm., July 2010) DNA isolation and PCR were done using the protocol of Kohler & Giinther (2008). Forward and reverse strands were aligned using Codon- Code Aligner v. 3.0.3 (CodonCode Corporation, Dedham, MA, USA) and corrected by eye. Sequences were aligned using MAFFT (Katoh & Toh 2008) and optimized using ALISCORE (Misof & Misof 2009). Phylogenetic analy- sis (Bayesian inference) was accomplished as conducted by Giinther et al (2010). The analysis of 480 base pairs of the 16S rRNA gene re- vealed that Cophixalus monosyllabus sp. n. is clearly a sister species of C. tetz/affi and both are a sister clade of C. balbus (Fig. 11). C. tridactylus and C. humicola appear more distant in the molecular tree (see also Kohler & Giin- ther 2008), and indicate that the present genus Cophixalus most probably is polyphyletic. The genetic distance (un- corrected p-distance) between C. monosyllabus sp. n. and C. tetzlaffi is 4.3 % for the 16S rRNA gene. Comparison with other species. Cophixalus monosylla- bus sp. n. is distinct from other Cophixalus species, des- cribed up to the year 2003, in the same characters as is C. tetzlaffi (Giinther 2003). All 16 Cophixalus species des- cribed after 2003 (Hoskin 2004; Kraus & Allison 2006, 2009; Gtnther 2006; Richards & Oliver 2007) differ clearly from C. monosyllabus sp. n. in body size and al- so in their advertisement calls. The only species with which C. monosyllabus sp. n. can be confused morpho- logically is C. tetzlaffi, especially as both species occur syntopically. Bonn zoological Bulletin 57 (2): 231-240 .070 .065 .060 O55 050 O45 A B Fig. 12. Box-Whisker-Plot of the ratio “diameter of disc of fourth toe/snout-urostyle-length” (F3D/SUL) in Cophixalus tetz- laffi (A) compared to that of Cophixalus monosyllabus sp. n. (B). ©ZFMK 240 Rainer Gunther I compared the measurements of ten male C. monosyllabus sp. n. with that of eight male C. fetzlaffi and found the fol- lowing differences: with a mean body size (SUL) of 23.0 mm (range 20.6—24.3 mm), C. monosyllabus sp. n. 1s somewhat larger than C. fetzlaffi (mean 21.4 mm, range 19.5—22.6 mm), Student’s t-test revealed a significant dif- ference with t=2.98 and P=0.0046 in this character; C. monosyllabus sp. n. has significantly shorter tibiae than C. tetzlaffi (mean of TL/SUL in C. monosyllabus sp. n. 0.50, that in C. tetzlaffi 0.52, t= 3.39, P=0.0019); C. mono- syllabus sp. n. has a longer third finger than C. tetzlaffi (mean of F3L/SUL in the former 0.27, in the latter 0.26, t=3.01, P=0.0041; C. monosyllabus sp. n. has a wider ter- minal disc on the fourth toe than C. fetzlaffi (mean of T4D/SUL in the former 0.054, in the latter 0.046, t=3.11, P=0.0094); C. monosyllabus sp. n. has a wider terminal disc on first toe than C. tetzlaffi (mean of T1D/SUL in the former 0.027, in the latter 0.018, t=5.63, P=0.00002); C. monosyllabus sp. n. has a wider terminal disc on first fin- ger than C. tetzlaffi (mean of F1D/SUL in the former 0.021, in the latter 0.015, t=5.36, P=0.00003) and, most significantly, C. monosyllabus sp. n. has a wider terminal disc on the third finger than C. tetzlaffi (mean of F3D/SUL in the former 0.063, and in the latter 0.051, t=6.14, P=0.000007) (Fig. 12). There are continuous dorsolater- al skin ridges in C. ftetzlaffi, but discontinuous dorsolat- eral skin glands in C. monosyllabus sp. n. Apart from these morphological differences, and most im- portant for species differentiation, are the advertisement calls: C. monosyllabus sp. n. utters a single peeping note with a mean duration of 196 ms (range 173-224 ms), while the call of C. tetzlaffi consists of three to four peep- ing notes with a mean note duration of 422 ms (range 347-518 ms). Acknowledgements. Field work and collection of voucher speci- mens was permitted by representatives of Belai Besar Konser- vasi Sumber Daya Alam (KSDA), Sorong, Papua Province of Indonesia (PPI). Marthinus Kapisa (Biak/PPI), Andreas, Frank, and Apner Piahar (Kampung Lusiperi near Fakfak Town/PPI), and Christian Bergmann (Berlin/Germany) helped during field work. The Family Piahar also permitted the collection of frogs on their property. Rudolf Arndt (Pomona, New Jersey, USA) carefully read my draft and made a number of helpful comments. B. Stelbrink and T. von Rintelen (ZMB) “constructed” the mo- lecular tree. Elisa Forster (Ztirich, Switzerland) prepared Figures 2 and 3, and Frank Tillack (ZMB) provided technical help. To all of them I am deeply grateful. Bonn zoological Bulletin 57 (2): 231-240 REFERENCES Dingerkus G, Uhler LD (1977) Enzyme clearing of alcian blue stained whole small vertebrates for demonstrating cartilage. Stain Technology 52: 229-232 Frost DR (2010) Amphibian Species of the World: an online reference, version 5.4 (8 April 2010). American Museum of Natural History, New York. Available from: http://research.amnh.org/herpetology/amphibia/index.php Giinther R (2003) First record of the microhylid frog genus Cophixalus from western Papua, Indonesia, with descriptions of two new species. Herpetozoa 16 (1/2): 3-21 Giinther R (2006) Two new tiny Cophixalus species with reduced thumbs from the west of New Guinea (Anura: Microhylidae). Herpetozoa 19 (1/2): 59-75 Giinther R, Stelbrink B, Rintelen T von (2010). Oninia senglaubi, another new genus and species of frog (Amphibia, Anura, Mi- crohylidae) from New Guinea. Zoosystematics and Evolution 86 (2): 245-256 Hoskin CJ (2004) Australian microhylid frogs (Cophixalus and Austrochaperina): phylogeny, taxonomy, calls, distributions and breeding biology. Australian Journal of Zoology 52: 237— 269 Katoh K, Toh H (2008) Recent developments in the MAFFT multiple sequence alignment program. Briefings in Bioinfor- matics 9: 286-298 Kohler F, Giinther R (2008) The radiation of microhylid frogs (Amphibia: Anura) on New Guinea: A mitochondrial phyloge- ny reveals parallel evolution of morphological and life histo- ry traits and disproves the current morphology-based classi- fication. Molecular Phylogenetics and Evolution 47: 353-365 Kraus F, Allison A (2006) Three new species of Cophixalus (Anura: Microhylidae) from southeastern New Guinea. Her- petologica 62: 202—220 Kraus F, Allison A (2009) New species of Cophixalus (Anura: Microhylidae) from Papua New Guinea. Zootaxa 2128: 1-38 Misof B, Misof K. (2009) A Monte Carlo approach successful- ly identifies randomness in multiple sequence alignments: A more objective means of data exclusion. Systematic Biology 58: 21-34 Richards SJ, Oliver PM (2007) A new species of Cophixalus (Anura: Microhylidae) from Misima Island, Papua New Gui- nea. Pacific Science 61 (2): 279-287 Received: 27.VII.2010 Accepted: 11.X.2010 OZFMK Bonn zoological Bulletin | Volume 57 Issue 2 pp. 241-255 Bonn, November 2010 | High mitochondrial sequence divergence meets morphological and bioacoustic conservatism: Boophis quasiboehmei sp. n., a new cryptic treefrog species from south-eastern Madagascar Miguel Vences !, Jérn Kohler 2, Angelica Crottini !3 & Frank Glaw 4 ! Division of Evolutionary Biology, Zoological Institute, Technical University of Braunschweig, Spielmannstr. 8, D-38106 Braunschweig, Germany; E-mail: m.vences@tu-bs.de ? Department of Natural History — Zoology, Hessisches Landesmuseum Darmstadt, Friedensplatz 1, D-64283 Darmstadt, Germany 3 Sezione di Zoologia e Citologia, Dipartimento di Biologia, Universita degli Studi di Milano, Via Celoria 26, I-20133 Milano, Italy 4 Zoologische Staatssammlung Miinchen, Miinchhausenstr. 21, D-81247 Miinchen, Germany Abstract. We describe a new species of treefrog from Madagascar that is highly similar in external adult morphology, bioacoustics and colouration to Boophis boehmei but differs from this species by a remarkable differentiation in a frag- ment of the mitochondrial 16S rRNA gene. A more detailed analysis revealed that this differentiation is concordant with the pattern in two nuclear genes (Rag! and POMC) which show no hapiotype sharing of the new species with B. boehmei, and with a consistent difference in tadpole morphology (third lower row of labial keratodonts reduced in length in the new species). We conclude that concordance between these independent characters indicates two independent evolution- ary lineages that should best be considered as separate species, despite their similar adult morphology. The new species, Boophis quasiboehmei sp. n., is so far known only from an area in the southern central east and south-east of Madagas- car, south of the Mangoro river, while B. boehmei is known only from the area around Andasibe north of the river Man- goro. Preliminary data indicate that this group of treefrogs contains several more cryptic species, and a simple explana- tion assuming the Mangoro river as a barrier being responsible for divergence between them is likely no longer tenable. Key words. Amphibia, Anura, Mantellidae, Boophis boehmei, Boophis quasiboehmei sp. n., Madagascar. INTRODUCTION Treefrogs of the genus Boophis have long been among Madagascar’s less studied amphibians, but intensified fieldwork and application of integrative taxonomy proto- cols have led to a steep increase of knowledge (Blommers- Schlosser 1979; Cadle 2003; Glaw & Vences 2007; Glaw et al. 2010). Many Boophis species call from high posi- tions in the vegetation and intensive nocturnal searches for calling males are needed to find them. Consequently, many species have been described on the basis of only small series or even single individuals, and females are often unknown. Furthermore, many species of Boophis are known to be morphologically very similar and a diagno- sis based on external morphology alone is often unreli- able (Glaw et al. 2001; Vences et al. 2008). However, be- cause the advertisement calls of these species are usual- ly loud and species-specific (Vences et al. 2006), the Bonn zoological Bulletin 57 (2): 241-255 integration of bioacoustics into their taxonomy has led to an improved understanding of Boophis species diversity. Together with an initial screening of molecular diversity, this has led to the description of many new species of Boophis (e.g., Andreone 1993, 1996; Andreone et al. 1995; Cadle 1995; Glaw & Thiesmeier 1993; Glaw & Vences 1992, 1994, 1997b, 2002; Glaw et al. 2001, 2010; Koh- ler et al. 2007, 2008; Vallan et al. 2003, 2010; Vences & Glaw 2002, 2005; Vences et al. 2010; Wollenberg et al. 2008) and the identification of a large number of addition- al, yet undescribed candidate species (Vieites et al. 2009). Furthermore, tadpoles of Boophis are among the most commonly encountered anuran larvae in Malagasy rain- forest streams (Vences et al. 2008), and a large number of them have recently been described (e.g., Raharivololo- niaina et al. 2006; Randrianiaina et al. 2009a, b). ©ZFMK 242 Miguel Vences et al. Taking the latest species descriptions into account, the genus Boophis, classified in the endemic Malagasy-Co- moroan family Mantellidae, currently comprises 71 de- scribed species. The genus is monophyletic and composed of two main clades that correspond to mainly stream- breeding (subgenus Boophis) and pond-breeding species (subgenus Sahona), respectively (Glaw & Vences 2006, 2007). The stream breeders are further divided into eight phenetic species groups. Most of these species groups probably are monophyletic units although some are not (particularly the Boophis majori group). The Boophis goudoti species group contains 13 small to large species of largely arboreal frogs that are mainly dis- tributed in the rainforests and highlands of Madagascar. A subgroup of small-sized species is characterized by colourful eyes, usually with red iris colour and a bluish iris periphery (Glaw & Vences 1997a, b). Several of these species such as Boophis boehmei, B. burgeri, B. reticula- tus, and B. rufioculis are known to occur at the same lo- cality in the Andasibe region in the northern central east of Madagascar and B. reticulatus, B. sp. aff. rufioculis and B. sp. aff. boehmei (= B. sp. 8 and B. sp. 16 of Vieites et al. 2009) in Ranomafana National Park in the southern central east. Of the various confirmed candidate species in the B. goudoti group (Glaw & Vences 2007; Vieites et al. 2009), four have recently been described (or older names were resurrected for them) on the basis of molec- ular, morphological, and/or bioacoustic differences (Glaw et al. 2010). However, no taxonomic conclusions have so far been drawn for the two candidate species from the Ra- nomafana region mentioned above (B. sp. 8 and B. sp. 16), mainly because of their high morphological similarity to Boophis rufioculis and to B. boehmei, respectively. Boophis boehmei is the smallest species in the B. goudoti group and has been originally described from Andasibe, where it is rather common (Glaw & Vences 1992). Pop- ulations from more southern localities, initially allocated to this species (Ranomafana region and Andohahela) turned out to be genetically highly divergent (Vieites et al. 2009) and have therefore been considered as Boophis sp. aff. boehmei (Glaw & Vences 2007) or B. sp. 16 (Vieites et al. 2009), although no reliable morphological or bioacoustic difference between them had been ob- served. The recent discovery of differences in the tadpole labial tooth row arrangements of Boophis boehmei and Boophis sp. 16 (Randrianiaina et al. 2009b) prompted us to undertake a more detailed comparison. On the basis of high mitochondrial divergences, consistent differences in two nuclear genes, constant differences in tadpole mor- phology, and subtle differences in iris colour, we conclude that the central south-eastern populations indeed consti- tute a distinct species which we describe herein as Boophis quasiboehmei. It is however worth to note that B. boehmei Bonn zoological Bulletin 57 (2): 241-255 and the newly described species are indeed among the morphologically and bioacoustically most cryptic species pairs so far discovered in Madagascar. MATERIALS AND METHODS Frogs were collected at night by opportunistic searching, using torches and head lamps. Specimens were euthanized in a chlorobutanol solution, fixed in 95% ethanol, and pre- served in 70% ethanol. Locality information was record- ed with GPS receivers. Specimens were deposited in the collection of Université d’ Antananarivo, Département de Biologie Animale, Antananarivo (UADBA), Zoologisches Forschungsmuseum Alexander Koenig, Bonn (ZFMK), and the Zoologische Staatssammlung Munchen (ZSM). FGMV, FGZC and ZCMV refer to F. Glaw and M. Vences field numbers. Terminology for biogeographic regions of Madagascar follows Boumans et al. (2007). Morphological measurements (in millimetres) were all done by M. Vences with a digital caliper (precision 0.01 mm) to the nearest 0.1 mm. Used abbreviations are: SVL (snout—vent length), HW (greatest head width), HL (head length), ED (horizontal eye diameter), END (eye—nostril distance), NSD (nostril-snout tip distance), NND (nos- tril-nostril distance), TD (horizontal tympanum diameter), TL (tibia length), HAL (hand length), HIL (hindlimb length), FOL (foot length), FOTL (foot length including tarsus), FORL (forelimb length), and RHL (relative hindlimb length). Terminology and description scheme follow Glaw et al. (2010). Webbing formulae follow Blommers-Schloésser (1979). Statistical analyses were per- formed with Statistica software (Statsoft Corp., Tulsa, USA). Vocalizations were recorded in the field using different types of tape recorders (Sony WM-D6C, Tensai RCR- 3222) and external microphones (Sennheiser Me-80, Vi- vanco EM 238), and an Edirol R-09 digital recorder with internal microphones and saved as uncompressed files. Recordings were sampled (or re-sampled) at 22.05 kHz and 16-bit resolution and computer-analysed using the software CoolEdit 98. Frequency information was ob- tained through Fast Fourier Transformation (FFT; width 1024 points). Spectrograms were obtained at Hanning window function with 256 bands resolution. Temporal measurements are given as range, with mean + standard deviation in parentheses. Terminology in call descriptions follows Kohler (2000). Two different molecular data sets were studied: First, we analyzed sequences of the mitochondrial 16S tRNA gene of around 500 bp from all Boophis goudoti OZFMK Boophis quasiboehmei sp. n., a new cryptic treefrog species from south-eastern Madagascar Table 1. Primer sequences and PCR conditions used in the present study. PCR conditions start with temperature (in °C) of each step followed by the time in seconds. Gene Primer name Sequence (5’ > 3’) Source PCR conditions BDNF — BDNF DRV | ACCATCCTTTTCCTKACTATGG Vieites et al. (2007) 94(120), [94(20), 57(45), BDNF — BDNF DRV 1 CTATCTTCCCCTTTTAATGGTC Vieites et al. (2007) 72(120) 39], 72(600) Ragl Amp F2 ACNGGNMGICARATCTTYCARCC s. Chiari et al. (2004) = 94(120), [94(20), 50(50), Ragl Amp R2 GGTGYTTYAACACATCTTCCATYTCRTA — s. Chiari et al. (2004) = 72(180) x 45], 72 (600) POMC POMCDRVFI ATATGTCATGASCCAYTTYCGCTGGAA Vieites et al. (2007) 95(120), [95(60), 58(60), POMC POMCDRVRI GGCRITYTTGAAWAGAGTCATTAGWGG _ Vieites et al. (2007) 72(90) x 35], 72(600) group species and candidate species with reddish iris colour as obtained by Vieites et al. (2009), Randrianiaina et al. (2009b) and StrauB8 et al. (2010). After alignment and removal of incomplete sections at its beginning and end the data set for analysis had a length of 479 bp. Unparti- tioned Bayesian inference searches were performed. The best model of evolution (GTR+G) was determined by AIC in MrModeltest (Nylander 2002). Bayesian analyses were performed with MrBayes 3.1.2 (Ronquist & Huelsenbeck 2003). Two runs of 10 million generations (started on ran- dom trees) and four incrementally heated Markov chains (using default heating values) each, sampling the Markov chains at intervals of 1000 generations were used. The last 5001 trees were retained post burn-in and summarized to generate the majority rule consensus tree. Second, we used tissue samples of four and one Boophis boehmei from Andasibe and An’ Ala, respectively, and four and two tissue samples of B. quasiboehmei from Sa- hamalaotra (=Samalaotra) and Ambohitsara (Tsitolaka for- est) for newly determining DNA sequences of various nu- clear genes. Toe clips or leg muscle tissue samples (pre- served in 95% ethanol) were used for DNA extraction. To- tal genomic DNA was extracted from the tissue samples using proteinase K digestion (10 mg/ml concentration) fol- lowed by a standard salt extraction protocol (Bruford et al. 1992). We amplified fragments of three genes from the nuclear DNA (nuDNA): brain-derived neurotrophic fac- tor (BDNF), recombination activating gene | (Rag1), and pro-opiomelanocortin (POMC). Standard Polymerase chain reactions were performed in a final volume of 11 ul and using 0.3 pl each of 10 pmol primer, 0.25 ul of to- tal dNTP 10 mM (Promega), 0.08 ul of 5 U/ml GoTagq, and 2.5 ul 5X Green GoTaq Reaction Buffer (Promega). Primers and detailed PCR conditions are provided in Table 1. PCR products were then purified through QIAquick pu- rification kit (Qiagen) according to the manufacturer’s in- struction. Purified PCR templates were sequenced on an automated DNA sequencer (Applied Biosystems ABI 3130XL). Chromatographs were checked and sequences Bonn zoological Bulletin 57 (2): 241-255 were edited using CodonCode Aligner (v. 2.0.6, Codon Code Corporation). All newly determined sequences have been deposited in GenBank (HQ380132-HQ380172). Haplotypes of POMC data were inferred using the PHASE algorithm (Stephens et al. 2001) implemented in DnaSP software (Version 5.10.3; Librado & Rozas 2009). Hap- lotype network reconstruction of phased sequences of the POMC (Fig. 2A) and Rag! (Fig. 2B) fragments were per- formed using the software TCS, version 1.21 (Clement et al. 2000). This software employs the method of Temple- ton et al. (1992) and it calculates the number of mutation- al steps by which pairwise haplotypes differ, computing the probability of parsimony for pairwise differences un- til the probability exceeds 0.95 (no manual adjustment of threshold was necessary). RESULTS A detailed analysis of all available 16S rRNA sequences of adults and tadpoles assigned to B. boehmei (GenBank accession numbers GQ904739-GQ904746, DQ792470-DQ792471, AY341717, AY848560— AY 848562) and the candidate species B. sp. 16 (sensu Vieites et al. 2009) (accession numbers GQ904717—GQ904738, AY848529—AY 848536) con- firmed that these two forms are genetically highly diver- gent. Depending on the length of the sequence available, the uncorrected pairwise distances were between 8.8% and 11.0% (note that these values are higher than the 6.8% re- ported by Vieites et al. (2009) because of different lengths of the sequences, with a different proportion of hypervari- able sites included in the analysis). Next to single substi- tutions we also detected one major insertion of seven nu- cleotides in the candidate species which in this extent was not present in any of the related species of Boophis (Fig. 1). Pairwise divergences were 0.0—-0.9% within B. boehmei, 0.0—0.5% within specimens of B. sp. 16 from the Ranomafana region, and 3.6-4.9% between the single available sequence of B. sp. 16 from Andohahela ©ZFMK 244 Miguel Vences et al. Boophis axelmeyeri (Tsaratanana - DQ118669) Boophis rufioculis (An'Ala - DQ003334) U——_______-. Boophis sp. 41 (Mahasoa - FJ559156) ** Boophis sp. 8 - aff. rufioculis (Maharira - ZCMV 235 - AY848535) Boophis sp.8 - aff. rufioculis (Antoetra - FAZC 11465 - AY848553) Lr Boophis sp. 8 - aff. rufioculis (Antoetra - FAZC 11451 - AY848551) “ Boophis sp.8 - aff. rufioculis (Antoetra - FAZC 11452 - AY848552) Boophis sp. 40 (Mahasoa - FJ559155) lime Boophis boehmei [Ca43 HM631885] (Sahafina - PSG 418) Lr Boophis boehmei [Ca43 HM631885] (Sahafina - PSG 313) “ Boophis boehmei [Ca43 HM631885] (Sahafina - PSG 417) *k* Boophis boehmei (Andasibe - LR 167 - DQ792471) re Boophis boehmei (Andasibe - FGMV 2001.1206 - AY848560) Boophis boehmei (Andasibe - MVTIS 2002G39 - AY848562) [ Boophis boehmei (Andasibe - LR 145 - DQ792470) Lp Boophis boehmei (Andasibe - FGMV 2001.1205 - AY848559) ** L— Boophis boehmei (Andasibe - FGMV 2001.1205 - AY341717) Lp Boophis boehmei (Andasibe - MVTIS 2002G38 - AY848561) “ Boophis boehmei (An'Ala - ZCMV 3508 - GQ904744) ** Boophis boehmei (An'Ala - ZCMV 3571 - GQ904746) “ Boophis boehmei (An'Ala - ZCMV 3482 - GQ904739) Boophis boehmei (An'Ala - ZCMV 3445 - GQ904740) Ly Boophis boehmei (An‘Ala - ZCMV 3555 - GQ904745) “ Boophis boehmei (An'Ala - ZCMV 3458 - GQ904741) Boophis quasiboehmei (Andohahela - FGZC 236 - AY848529) Boophis quasiboehmei (Ranomafana - ZCMV 324 - AY848536) “ Boophis quasiboehmei (Ranomafana - ZCMV 2690 - GQ904734) Boophis quasiboehmei (Ranomafana - FGMV 2002.327 - AY848534) “ Boophis quasiboehmei (Ranomafana - ZCMV 3624 - GQ904729) -— Boophis quasiboehmei (Ranomafana - ZCMV 3634 - GQ904731) “ Boophis quasiboehmei (Ranomafana - ZSM 1153/2007 - GQ904725) *k* Boophis quasiboehmei (Ranomafana - FGMV 2002.328 - AY848533) “ Boophis quasiboehmei (Ranomafana - FGMV 2002.324 - AY848530) L— Boophis quasiboehmei (Ranomafana - ZCMV 2688 - GQ904733) ‘ Boophis quasiboehmei (Ranomafana - ZCMV 3767 - GQ904728) Boophis quasiboehmei (Ranomafana - FGMV 2002.325 - AY848531) el “ Boophis quasiboehmei (Ranomafana - ZSM 752/2007 - GQ904719) |. Boophis quasiboehmei (Ranomafana - ZSM 1370/2007 - GQ904724) 0.1 “ Boophis quasiboehmei (Ranomafana - ZSM 1010/2007 - GQ904721) Boophis quasiboehmei (Ranomafana - ZCMV 4083 - GQ904726) “ Boophis quasiboehmei (Ranomafana - ZCMV 3045 - FJ559139) Boophis quasiboehmei (Ranomafana - FGMV 2002.326 - AY848532) . Boophis quasiboehmei (Ambohitsara - ZCMV 4937 - GQ904735) Boophis quasiboehmei (Ranomafana - ZSM 684/2007 - GQ904718) Lr Boophis quasiboehmei (Ranomafana - ZSM 932/2007 - GQ904720) “ Boophis quasiboehmei (Ranomafana - ZCMV 4491 - GQ904717) DQ792471 (NTAAA ET ARTE RETA T RATE 0Q792470 |AtAAAT TART ET TAATBATEGRAGA AY848562 [ATAAATEAR TDD ETAT EATE AY848561 (MTARATTART TED DARTRATEER AEN AY848560 [ATARAT TAA TEED TAATATEERABR------- AY848559 (AGARATTA ATED EM AERATEG HABA. ------ AY341717 HeeeeeeenraaHAnlaniiiata aS GQ904746 |ATAAA TT ANTE T TTA R EC RAGA 6Q904745 (ATEAREERE RADHA REREAD EAREER- ------ GQ904741 |AEAAAEANTONETANT HATO GEREN GQ904740 (TMM N TANT TET TAN TAREE ERAGK GQ904739 ATMA TE AATTTTTA ATE AY848536 (MTAAATTAM TERT EAATEAGE ERAT ABRRRATAAT CRATERS AY848534 (KTAAATTAAT TET OMATRAGEER ADAG CAO HOATOAEORESEAAE AY848533 |ATAAATTARTTTRRAATEARERRATAGEE RATAN TERA TEAM CHARGE ANDRE AECERREEE) AY848532 (HTAMATTAAT EDD EMATRARE ERAT ACR RRATANT ERAT OAAGRAAG RANA TT AGRE TERED) AY848531 (ATMAATTAATTTTEAATPARRRRAT AGG ERAT ARTE RA DENNER ARECRAB TA AY848530 (MPAAATTAAT TDD GAATHAREE RAT AERORATAATERARGAAGRAAGCAABET Bi AY848529 (MHMAATTAMTEDEDAATRAREE RAT AERGRATAAT ERAT ERAGE GAMERTAG TO REE Fig. 1. Phylogenetic tree of species and candidate species of the Boophis goudoti group with red iris colour, obtained using Bayesian inference based on DNA sequences of the mitochondrial 16S rRNA gene (alignment length 479 bp). Bayesian posterior values >0.95 symbolized by a single asterisk, of 0.99-1.00 by two asterisks. For each sequence, locality, voucher number and Genbank number are given in parentheses. Boophis goudoti was used as outgroup (not shown). Note that the deeper phylogenetic relation- ships shown are not reliable due to the limited amount of sequence information used in the analysis, and according to an unpub- lished multi-gene data set of K. C. Wollenberg, B. boehmei and B. quasiboehmei are probably sister groups. The alignment in the lower part of the figure shows a section of the 16S alignment, with sequences of Boophis boehmei (upper 13 sequences; numbers to the left are Genbank accession numbers) and Boophis quasiboehmei (lower seven sequences). The insertion of seven nucleotides is a synapomorphy of all B. guasiboehmei specimens for which a sequence was obtained, and in this extent is lacking also in all other species of the B. goudoti group. Bonn zoological Bulletin 57 (2): 241-255 ©ZFMK Boophis quasiboehmei sp. n., a new cryptic treefrog species from south-eastern Madagascar 245 10R 6R Fig. 2. Boophis quasiboehmei ® Sahamalaotra = Ambohitsara Boophis boehmei Andasibe = An'Ala fe— R 1\2\ 358 SS Haplotype networks of the nuclear POMC (A) and Rag! (B) genes fragments in B. boehmei and B. quasiboehmei, each from two different localities. Haplotypes per each individual were inferred using the Phase algorithm. The networks show com- plete absence of haplotype sharing among the two taxa. (AY848529) and those from Ranomafana. Genetically identified specimens assigned to B. boehmei were from Andasibe and An’ Ala. Specimens from Sahafina (Gehring et al. 2010) had quite divergent DNA sequences and their status is unclarified, but they clustered with B. boehmei (Fig. 1). Following the scheme suggested by Padial et al. (2010), this population was considered a new unconfirmed candidate species Boophis boehmei [Ca43 HM631885] by Gehring et al. (2010). Probably, specimens from Ankeni- heny for which no molecular data are available belong to this species as well. Specimens assigned to B. sp. 16 were from the Ranomafana area (including Ambatovory, Sa- hamalaotra, Imaloka, Kidonavo, Vohiparara) and Ambo- hitsara, as well as from Andohahela. Besides a simple assessment of molecular divergences be- tween Boophis sp. 16 and B. boehmei it is also necessary to comment on its phylogenetic position. The analysis of Vieites et al. (2009) placed B. boehmei with B. sp. 8 from Ranomafana and B. sp. 40 from Mahasoa forest, and the clade made up by these species was sister to B. sp. 16. Our Bonn zoological Bulletin 57 (2): 241-255 analysis (Fig. 1) included sequences of all these taxa and confirmed the phylogenetic relationships suggested by Vieites et al. (2009). However, an unpublished analysis based on multiple mitochondrial genes by K.C. Wollen- berg instead suggested a probable sister-group relationship between B. boehmei and B. sp. 16, confirming that the 16S rRNA gene alone as used here is insufficient to clarify the phylogeny among Boophis species. Altogether, the phy- logenetic relationships among all these species require a much more detailed analysis which however is beyond the scope of the present paper. The results of the mitochondrial marker indicate no or lim- ited gene flow between B. boehmei and B. sp. 16. This re- sult was corroborated by the analysis of two nuclear mark- ers (Fig. 2; the conserved BDNF gene showed no varia- tion). 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Factor 1 Factor 2 Factor3 Factor 4 SVL 0.535637 0.024396 0.278024 0.432915 HW 0.401178 -0.052815 0.722325 0.434988 HL 0.283729 = -0.059521 0.901536 -0.000783 TD -0.451112 0.171742 0.508941 0.315946 ED 0.042109 0.184460 0.248786 0.727413 END 0.263940 0.645782 0.593063 0.078554 NSD -0.058373 0.886713 -0.111786 0.323847 NND 0.169004 0.159007 -0.021443 0.796064 FORL 0.776970 = -0.189957 0.113991 0.209375 HAL 0.936216 0.015471 0.113066 -0.107870 HIL 0.852890 = -0.051712 0.150553 0.154111 FOTL 0.932469 0.147455 0.044485 0.068645 FOL 0.784388 0.333820 0.095038 0.209488 TIBL 0.743402 0.023610 0.266693 0.084073 Eigenvalue 5.977193 2.369640 1.412833 0.905600 % Variance 42.69424 16.92600 10.09166 6.46857 colouration. All specimens of B. boehmei had a bright red outer iris area and a brownish inner iris area, whereas B. sp. 16 had no such bright red colour but orange, either as a more or less uniformly orange iris or as an orange out- er iris area. In a search for a possible morphometric differentiation, we carried out a Principal Component Analysis on the ba- sis of measurements in Table 2 (males only). The analy- sis resulted in three factors with Eigenvalues greater than 1 (Table 3) which together explained 70% of the total vari- ation. Because size of specimens was similar, the first fac- tor was not representative mainly of body size, but of rel- ative limb length; the highest factor loadings were for vari- ables associated with limb length (Table 3). Factors 2 and 3 were associated with the shape of the head: Factor 2 with END and NSD, and Factor 3 with mainly HW and HL. While Factor | resulted only marginally in a trend of sep- aration of the two species (not shown), Factors 2 and es- pecially 3 separated most specimens of B. boehmei vs. B. sp. 16 (Fig. 3). However, univariate analyses on the ba- sis of the variables with highest factor loadings did not result in a convincing separation (not shown), indicating that morphometric data cannot serve as diagnostic char- acters to separate these two forms. Bonn zoological Bulletin 57 (2): 241—255 The most convincing diagnostic character comes from tad- pole morphology and has been described in detail by Ran- drianiaina et al. (2009a, b): all tadpoles of Boophis sp. 16 (from Ranomafana and Ambohitsara; N = 75) examined had a short (or completely absent) third posterior row of labial keratodonts (P3), whereas in B. boehmei (from lo- calities Andasibe and An’ Ala) this row was slightly short- er than in other species of the B. goudoti group, but still much longer than in B. sp. 16, with no overlap in num- bers of labial keratodonts in P3 and almost no overlap in relative length of P3 (Fig. 4). Given this constant difference in tadpole morphology which fully correlates with high mitochondrial divergences (among the highest observed between closely related man- tellid frog species), and with fully separated haplotypes in two nuclear genes, we conclude that B. boehmei and B. sp. 16 constitute two separate and independent evolu- tionary lineages. Therefore, they should best be consid- ered as distinct species, although cryptic in adult morphol- ogy and advertisement calls. In the following we thus de- scribe B. sp. 16 as a new species. Boophis quasiboehmei sp. n. (Figs 5—6) Holotype. ZSM 227/2006 (field number ZCMV 3045), adult male (Fig. 5), collected at Ambatovory, at the edge of Ranomafana National Park, south-eastern Madagascar, 21°14,279’ S, 47°25,487’ E, 966 m a.s.l., on 26 February 2006 by M. Vences, Y. Chiari, T. Rajoafiarison, E. Raje- riarison, P. Bora and T. Razafindrabe. Paratypes. ZFMK 59881-59882, two adult males, col- lected in the Ranomafana region, south-eastern Madagas- car, in December 1994 by M. Burger; ZSM 715/2003 (FG/MV 2002-0363), one adult male, collected at Vo- hiparara (close to the Kidonavo bridge), Ranomafana Na- tional Park, 21°13’ S, 47°22’ E, ca. 1000 m a.s.1., on 20 January 2003, by F. Glaw, M. Puente, L. Raharivololoni- aina, M. Thomas and D. R. Vieites; ZSM 228/2006 (ZCMV 3051), ZSM 229/2006 (ZCMV 3069), and ZSM 230/2006 (ZCMV 3070), three adult males, from same lo- cality and with same collectors and collection date; ZSM 224/2006 (ZCMV 2988), male, collected at Sahamalao- tra, Ranomafana National Park, south-eastern Madagas- car, 21°14.113’ S, 47°23.767’ E, south-eastern Madagas- car, on 25 February 2006 by M. Vences, Y. Chiari, T. Ra- joafiarison, E. Rajeriarison, P. Bora and T. Razafindrabe; ZSM 226/2006 (ZCMV 2951), male, collected at Imalo- ka, Ranomafana National Park, south-eastern Madagas- car, 21°14,527’ S, 47°27,909’ E; 1020 m a.s.1., on 23 Feb- ruary 2006 by Y. Chiari, P. Bora, T. Rajoafiarison, E. Ra- jeriarison, and T. Razafindrabe; ZSM 231/2006 (ZCMV OZFMK Boophis quasiboehmei sp. n., a new cryptic treefrog species from south-eastern Madagascar 249 Fig. 5. Dorsolateral (A) and ventral (B) views of the male holotype of Boophis quasiboehmei sp. n. (ZSM 227/2006) in life. 3360), male, collected at Ranomena, 21°12,736’ S, 47°26,010’ E, Ranomafana National Park, south-eastern Madagascar, on 28 February 2006, M. Vences, Y. Chiari, T. Rajoafiarison, and E. Rajeriarison; ZSM 232/2006 (ZCMV 3374) from the Ranomafana region, perhaps col- lected at Ranomafanakely river but without precise col- lecting data; ZSM 2322/2007 (ZCMV 5948), male, col- lected at Sahamalaotra, Ranomafana National Park, south-eastern Madagascar, 21°14.113’S, 47°23.767’ E, on 5 March 2007 by M. Vences, A. StrauB, R. D. Randriani- aina, and K. C. Wollenberg. Bonn zoological Bulletin 57 (2): 241-255 Diagnosis. Assigned to the genus Boophis based on the presence of an intercalary element between ultimate and penultimate phalanges of fingers and toes (verified by ex- ternal examination), presence of nuptial pads and absence of femoral glands in males, and overall similarity to oth- er Boophis species. Assigned to the Boophis goudoti group because of its brownish ground colour, presence of der- mal flaps or tubercles on heels and elbows, presence of white tubercles ventrally of the cloacal opening, presence of a sharp canthus rostralis, absence of red skin colour, and molecular phylogenetic relationships. ©ZFMK 250 Miguel Vences et al. Fig. 6. Specimens of Boophis quasiboehmei sp. n. in life: (A) frontal and (B) ventral views of a male from Ambohitsara (field number ZCMV 5867); (C) dorsolateral and (D) ventral views of a male from Andohahela (deposited in UADBA); (E) male from Ranomafana (deposited in UADBA); (F) male paratype ZFMK 59882 from Ranomafana (photo by M. Burger). Together with B. boehmei, the smallest species in the Boophis goudoti group characterized by a deviant oral morphology of the tadpole which is unknown from any other Boophis species. Boophis quasiboehmei sp. n. dif- fers from all described species in the B. goudoti group by substantial genetic differentiation (> 6% pairwise diver- gence in a fragment of the 16S rRNA gene) and further- Bonn zoological Bulletin 57 (2): 241—255 more from B. goudoti, B. obscurus, B. periegetes, B. madagascariensis, B. roseipalmatus, B. brachychir, B. entingae, B. rufioculis, B. burgeri, B. reticulatus, B. ax- elmeyeri, and B. spinophis by smaller size (SVL of adult males 28-31 mm versus 31—82 mm) and bioacoustic dif- ferentiation (see Vences et al. 2006 for details). B. quasi- boehmei sp. n. is most similar to B. boehmei and differs ©ZFMK Boophis quasiboehmei sp. n., a new cryptic treefrog species from south-eastern Madagascar 251 Fig. 7. Male specimens of Boophis boehmei from Andasibe in life: (A) paratype ZSM 563/1999 (originally ZFMK 53643); (B) paratype ZFMK 52637. from this species by an orange (versus red) outer iris ring, by a very short third posterior keratodont row in the tad- pole, consisting of only 0-15 keratodonts (versus 23-63 keratodonts in B. boehmei, see Randrianiaina et al. 2009b), and substantial genetic differentiation. Description of holotype. Adult male in excellent state of preservation, muscles of right thigh removed as DNA tis- sue sample. SVL 26.7 mm. Body moderately slender; head slightly longer than wide, wider than body; snout point- ed in dorsal view, obtuse to acuminate in lateral view; nos- trils directed laterally, eqidistant to eye and to tip of snout; canthus rostralis sharp, straight in dorsal view from eye to nostril, slightly curved from nostril to tip of snout; lo- real region slightly concave; eye large; tympanum distinct, rounded, TD 54% of ED; supratympanic fold narrow, prominent; vomerine odontophores distinct, well separat- ed in two slightly elongated patches, positioned median between choanae; choanae medium-sized, rounded. Tongue distinctly bifid and free posteriorly. Arms mod- erately slender; a small pointed dermal appendage on el- Bonn zoological Bulletin 57 (2): 241-255 bow; subarticular tubercles single, round; inner palmar tu- bercle poorly recognizable; fingers poorly webbed and without lateral dermal fringes; webbing formula 1(—), 21(—), 2e(1), 3111.5), 3e(1.5), 4(1); relative length of fin- gers 1<2<4<3 (finger 2 distinctly shorter than finger 4); finger discs enlarged. Hind limbs slender; a pointed der- mal appendage on heel; tibiotarsal articulation reaching widely beyond snout tip when hind limb is adpressed along body; lateral metatarsalia separated by webbing; in- ner metatarsal tubercle medium-sized, distinct, elongat- ed; no outer metatarsal tubercle; toes moderately webbed; webbing formula 1(0), 2i(1), 2e(0), 31(1), 3e(0), 41(2), 4e(2), 5(0.75); relative length of toes 1<2<3=5<4; toe discs enlarged. Skin smooth on dorsal surfaces, smooth on throat and chest, coarsely granular on belly, rather smooth on ventral surface of thighs, prominent scattered tubercles around cloaca. A worm-like parasite (possibly a nematode) apparently tried to escape when the frog was preserved and sticks in the left nostril. Measurements (in mm): SVL 26.7, HW 10.6, HL 11.2, ED SH/REND 23. NSD23.NND)3:3, 0D) 2.0, Th 15.2, HAL 9.1, FOL 11.1, FOTL 20.8. After almost four years in preservative, ground colour of upper surface of head, dorsum and limbs greyish brown, with few irregularly scattered and indistinct darker mark- ings; supratympanic fold and tympanic region not distinct- ly coloured; upper lip creamy white; dorsal surfaces of thigh, shank, tarsus and external toe, as well as lower arm, hand and external finger with distinct dark brown cross- bands; flanks brown with small pale white spots and dots, forming a reticulated pattern; several whitish dots below the cloaca, but no additional single white tubercles in the cloacal region; posterior surfaces of thighs greyish pale brown with beige reticulation on the proximal part, light brown without reticulations in the distal part; ventral sur- face creamy beige, with some pale greyish mottling along the lower jaw, the lower arms, hands and feet. In life, ground colour of upper surface of head, dorsum and legs light brown (slightly darker on the head), with few irregularly scattered yellowish spots on the back and scattered dark dots on back and more densely on the lat- eral parts of the head; flanks with reticulated pattern of brown, yellow and white; upper surfaces of hands and feet mottled with brown and yellowish; outer edge of tarsus with thin white line and white tarsal tubercle, outer edge of lower arm with white tubercle; two irregular rows of white tubercles on shank; dorsal surfaces of limbs with moderately distinct brown crossbands; posterior surfaces of thighs white, numerous white tubercles around the cloa- ca and uniformly brown posteriorly. Throat, chest and ven- ter creamy white; two irregular bluish spots on throat. Ven- tral surfaces of limbs only partially with whitish pigment, ©ZFMK 252 Miguel Vences et al. largest parts of thighs, shanks, hands and feet without white pigment. Outer iris almost uniformly bright orange, broadened above; inner iris ring brownish with some ves- sel-like brown reticulation; iris surrounded by a black ring; posterior iris periphery blue. Variation. All paratypes were similar to the holotype in general morphology. For measurements, see Table 2. Male SVL ranged from 26.7—29.3 in the Ranomafana region, and was 30.8 mm in one specimen from Midongy. No fe- males are known. Colouration was relatively constant in various localities of Ranomafana National Park, and in Ambohitsara (Fig. 6). The rather uniform orange eye colouration in life was typical for most specimens although at Andohahela (Fig. 6C) specimens tentatively assigned to this species had a more reddish eye colour. Distribution. Besides different sites in (1) the Ra- nomafana region, the species is also known from (2) Tsi- tolaka forest near Ambohitsara, about 30 km from Ra- nomafana, and was tentatively identified from (3) Befo- taka-Midongy Reserve (specimen ZSM 178/2006), and from (4) Andohahela National Park (Col Tanatana, 24°44’ S, 46°50’ E, 750 m a.s.l.). in the extreme south- east of Madagascar (GenBank accession number AY848529; specimen FGZC 236, deposited in UADBA). Natural history. At Ranomafana National Park, Boophis quasiboehmei sp. n. was one of the most common species of frogs and its larvae occurred in 29 out of 30 streams surveyed for tadpoles (Randrianiaina et al. 2009b; StrauB et al. 2010). Adult specimens, however, were less com- monly found, and in some areas occurred only in some densely clustered demes along small stretches of the streams. Males were observed calling at night from perch heights of 2-3 m from bushes and trees close to streams in primary as well as degraded rainforest. Vocalization. Generally, calls of Boophis quasiboehmei sp. n. exhibit a characteristic structure, consisting of short to moderately long pulsatile notes. However, the pattern of emission of these notes is highly variable and mostly irregular. Sometimes, notes are combined to regular se- ries (2—6 notes), with the initial note being longer than sub- sequent secondary notes. The calls emitted by the holo- type (Fig. 8) and recorded on 26 February 2006 at Am- batovory have the main frequency distributed between 2100 and 3400 Hz, with additional frequency bands of lower amplitude at 5500-6000 Hz and 8100-8900 Hz. Nu- merical parameters for the holotype calls are as follows (range followed by mean + standard deviation): duration of note series, 335-736 ms (519 + 203; n = 3); number of notes per series, 3-6 (4.3 + 1.5; n = 3); note duration (including initial notes within series), 66-79 ms (72.1 + 4.6; n = 8), duration of secondary notes within series, Bonn zoological Bulletin 57 (2): 241-255 Frequency (kHz) No Relative amplitude © 0 250 500 750 1000 Relative amplitude 0 25 50 75 100 Time (ms) Fig. 8. Spectrogram, corresponding waveform, and expanded waveform (bottom) of the initial note of a regular note series emitted by the holotype of Boophis quasiboehmei sp. n. Record- ing obtained on 26 February 2006 at Ambatovory, Ranomafana National Park. 20-34 ms (24.9 + 4.6; n = 8); pulses/note, 5-19 (12.6 + 6.2; n= 15); inter-note intervals, 97-125 ms (109.9 + 7.5; n= 10); dominant frequency, 2680-2963 Hz (2807 + 86; n= 10). A short sequence with three notes recorded on 1 March 1996 at Ranomafana (Vences et al. 2006, CD 1, track 66) has the following parameters: duration of note series, 373 ms, notes/series, 3; note duration, 18—58 ms; pulses/note, 5—12; inter-note intervals, 139-142 ms; dominant frequen- cy, 2550-2637 Hz. Calls of B. quasiboehmei sp. n. from Ambohitsara recorded on 3 March 2007 generally agree in structure with those emitted by the holotype, although they have shorter note duration and more variable, distinctly longer inter-note intervals. Numerical parameters are as follows: duration of note series, 527 ms (n = 1); number of notes per series, 6 (n= 1); note duration (including initial notes within series), 22-47 ms (35.2 + 7.3; n= 13); pulses/note, 4—12 (7.9 +2.5; n= 18); inter-note intervals, 475—942 ms (724.4 + 138.8; n= 16); dominant frequency, 2293-2572 Hz (2465 x 90; n= 9). Comparative call data. The morphologically most sim- ilar species, Boophis boehmei, has an almost identical call compared to that of B. quasiboehmei sp. n. A re-analysis of calls of B. boehmei from Andasibe (type locality) recorded on 12 January 1992 at 23°C (Fig. 9) revealed the following parameters: duration of note series, 455—530 ms ©OZFMK Boophis quasiboehmei sp. n., a new cryptic treefrog species from south-eastern Madagascar 253 Relative amplitude © 0 250 500 750 1000 Relative amplitude 0 25 50 75 100 Time (ms) Fig. 9. Spectrogram, corresponding waveform, and expanded waveform (bottom) of the initial note of a regular note series emitted by Boophis boehmei. Recording obtained on 12 Janu- ary 1992 at Andasibe (air temperature 23°C). (n = 2); number of notes per series, 3—4 (n = 2); note du- ration, 27-106 ms (62.6 + 23.0; n = 11); pulses/note, 10—24 (15.7 + 4.8; n= 9); inter-note intervals, 93-157 ms (125.0 + 29.6; n =5); dominant frequency, 2640-3177 Hz (2835 + 165.6; n= 8). A second recording from the type locality of B. boehmei recorded on 7 December 2001 at 24.8°C (Vences et al. 2006, CD 1, track 64) differs from the one described above by longer inter-note intervals. Numerical parameters are as follows: note duration, 34-98 ms (62.7 + 19.7; n= 18); pulses/note, 13—23 (16.7 + 3.6; n = 11); inter-note inter- vals, 591-1070 ms (766.1 + 210.0; n =7); dominant fre- quency, 2360-2980 Hz (2760 + 198; n = 12). In this recording, a single regular series composed of 6 notes 1s present, exhibiting note durations of 3444 ms and inter- note intervals within the series of 61—85 ms. A call recording of B. boehmei from Ankeniheny record- ed on 20 March 1994 at 22°C air temperature showed note duration of 16—61 ms, inter-note intervals of 162—164 ms and a dominant frequency of 2500-2800 Hz. In conclusion, there are no temporal or spectral call char- acters that distinguish B. boehmei from B. quasiboehmei sp. n. (see above). Etymology. The specific epithet is a combination of the Latin word ‘quasi’, meaning ‘almost’, and a patronym for Wolfgang Bohme (ZFMK). It refers to the impressively Bonn zoological Builetin 57 (2): 241-255 cryptic morphological and bioacoustic similarity of the new species to Boophis boehmei. DISCUSSION The initial detection of a probable species status of Boophis quasiboehmei sp. n. was based on its large diver- gence in a single marker of mitochondrial DNA. Due to the extent of this divergence (>6% to all described species), Vieites et al. (2009) deviated slightly from their usual rationale and listed this species as confirmed can- didate species, despite the lack of concordant indications by independent taxonomic characters. Although the work protocol of integrative taxonomy proposed by Padial et al. (2010) would allow for the description of species based on single characters if these are deemed to be sufficient- ly indicative of the existence of independent evolution- ary lineages, we do not recommend this procedure. In- stead, we only decided to formally describe B. quasi- boehmei sp. n. as new species once that independent and congruent evidence of various taxonomic characters had accumulated, even if those were subtle at first view: (1) a weak and not fully constant difference in adult eye colouration, (2) a slight tendency of morphometric differ- entiation detectable only by multivariate techniques, (3) a constant difference in tadpole morphology, and (4) con- cordance between three independent molecular markers (two nuclear and one mitochondrial). The molecular con- cordance alone would be sufficient for species recogni- tion under the genealogical concordance method of phy- logenetic species recognition, GCPSR (Avise & Ball 1990), but the further strict concordance with one mor- phological character (tadpole labial keratodonts) provides amore convincing evidence, especially because it is based on large series of individuals (Randrianiaina et al. 2009a, b). We are therefore convinced that Boophis quasiboehmei sp. n. and B. boehmei are to be considered as distinct species under an evolutionary or general lineage species concept (de Queiroz 2007). Among the various mechanisms of species diversification discussed for Madagascar (Vences et al. 2009), two (the watershed and the river barrier mechanism) might apply to the species pair B. boehmei and B. quasiboehmei sp. n. that occur in two different neighbouring centres of en- demism (CE2 and CE3) as defined by Wilmé et al. (2006), and because these two CEs are divided by the Mangoro river that has been invoked as an important river barrier in eastern Madagascar (see Vences et al. 2009). Discern- ing between these hypotheses is difficult, but both are con- tradicted by the fact that B. quasiboehmei sp. n. also oc- curs in Andohahela, which is in a different CE (CE5) and separated by a further large river barrier (the Mananara river). Also, the fact that numerous other red-eyed ©ZFMK treefrog species and candidate species have been already identified from eastern Madagascar (see Vieites et all. 2009: B. axelmeyeri, B. rufioculis, B. sp. 8, B. sp. 40, B. sp. 41), several of which appear to be microendemic to small areas while others might be more widespread, in- dicates a more complex situation. Only a more compre- hensive study of this group, with assessments of the sta- tus of all candidate species and their phylogenetic rela- tionships, and a more detailed analysis of their distribu- tion, will significantly contribute to the understanding of the diversification mechanisms that may have lead to this surprising morphological cryptic diversity. However, the fact that the phylogenetic position of B. boehmei and B. quasiboehmei 1s unclarified should not be interpreted as casting doubts on the species status of B. quasiboehmei since this new species 1s differentiated from topotypical B. boehmei by a high genetic differentiation and tadpole mouthparts, and from all other nominal species in the B. goudoti group by a high genetic differentiation, tadpole mouthparts, adult morphology, and advertisement calls. However, clarifying the phylogenetic relationships of all species and candidate species will be important to under- stand the status of the various UCS and CCS in the group and to be able to provide formal descriptions of those for which the data will confirm the status as distinct species. Additional data still missing at this time are on tadpole morphology of the populations from Andohahela, M1- dongy, Sahafina and Mahasoa that herein we have as- signed in a preliminary way to B. quasiboehmei (Ando- hahela, Midongy) or different candidate species (Sahafi- na, Mahasoa). At Ranomafana National Park, Boophis quasiboehmei sp. n. was commonly encountered at least in its tadpole stage, and its occurrence was confirmed at Andohahela Nation- al Park and tentatively in Befotaka-Midongy National Park. Although we never observed the species in second- ary vegetation formations, it appears to be tolerant to some degree of rainforest degradation. The relatively large dis- tribution area (from Ranomafana to Andohahela), its oc- currence in at least two protected areas and large area of occupancy at least in the Ranomafana area lead us to pro- pose an IUCN red list status of Least Concern for this new- ly described species (compare Andreone et al. 2005, 2008). Acknowledgements. For assistance in the field we are indebt- ed to Ylenia Chiari, Parfait Bora, Emile Rajeriarison, Theo Ra- joafiarison, Tokihery Razafindrabe, Axel StrauB, and Katharina C. Wollenberg. Marius Burger provided a photo and collected specimens of the new species. Roger-Daniel Randrianiaina kind- ly contributed the tadpole photographs. Meike Kondermann and Gabriele Keunecke helped with lab work. This study was made possible by collaboration agreements of the author’s institutions with the Université d’ Antananarivo Département de Biologie An- imale (UADBA) and the Association Nationale pour la Gestion des Aires Protegées. 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A tadpole’s goatee helps to distinguish two cryptic treefrog species of the genus Boophis. Herpetol- ogy Notes 2: 165-173 Bonn zoological Bulletin 57 (2): 241-255 Raharivololoniaina L, Grosjean S, Rasoamampionona Raminosoa N, Glaw F, Vences M (2006) Molecular identifi- cation, description and phylogenetic implications of the tad- poles of 11 species of Malagasy treefrogs, genus Boophis. Journal of Natural History 40: 1449-1480 Ronquist F, Huelsenbeck JP (2003) MrBayes 3: Bayesian phy- logenetic inference under mixed models. Bioinformatics 19: 1572-1574 Stephens M, Smith NJ, Donnelly P (2001) A new statistical method for haplotype reconstruction from population data. American Journal of Human Genetics 68: 978-989 StrauR® A, Reeve E, Randrianiaina R, Vences M, Glos J (2010) The world’s richest tadpole communities show functional re- dundancy and low functional diversity: ecological data on Madagascar’s stream-dwelling amphibian larvae. 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A preliminary evaluation of survey techniques and identification reliability of Malagasy amphibians. Mono- grafie d. Mus. Regio. di Scienze Natur. di Torino 45: 233-253 Vences M, Glaw F (2002) Two new treefrogs of the Boophis rap- piodes group from eastern Madagascar. Tropical Zoology 15: 141-163 Vences M, Glaw F (2005) A new cryptic frog of the genus Boophis from the northwestern rainforests of Madagascar. African Journal of Herpetology 54: 77—84 Vences M, Glaw F, Marquez R (2006) The Calls of the Frogs of Madagascar. 3 Audio CDs and booklet, 44 pp. Alosa-Fono- zoo, Barcelona Vences M, Wollenberg KC, Vieites DR, Lees DC (2009) Mada- gascar as a model region of species diversification. Trends in Ecology and Evolution 24: 456-465 Vieites DR, Min MS, Wake DB (2007) Rapid diversification and dispersal during periods of global warming by plethodontid salamanders. Proceedings of the National Academy of Sci- ences of the U.S.A. 104: 19903-19907 Vieites DR, Wollenberg KC, Andreone F, Kohler J, Glaw F, Vences M (2009) Vast underestimation of Madagascar’s bio- diversity evidenced by an integrative amphibian inventory. Proceedings of the National Academy of Sciences of the U.S.A. 106: 8267-8272 Wilmeé L, Goodman SM, Ganzhorn JU (2006) Biogeographic evolution of Madagascar’s microendemic biota. Science 312: 1063-1065 Wollenberg KC, Andreone F, Glaw F, Vences M (2008) Pretty in pink: a new treefrog species of the genus Boophis from north-eastern Madagascar. Zootaxa 1684: 58-68 Received: 30.VI.2010 Accepted: 10.1X.2010 ©ZFMK (oy an 08 4 SA Sh anh ' ‘ cre : a8, Th | wei » < i Aimar Ndi eek. «thal Me onal : oy any i va “Hs y : j vee 6 M01) 04 epee 4 opty ae igiy (boy) ae : ' ; ry a mT an. Aa A diy m4 i | i & j ; | | i : : ’ e ’ | | a) cg | | / i) ; 1 | j 7 ; 4 i Re ‘ = cane | i aA iN A fo hoy b Bie ois te t a j Y 7 ' 4 7 inal 1 hf i ul | { | LS Ue Wvpatllh Seige : ; aay ‘i i {) i r Bonn zoological Bulletin Volume 57 Issue 2 | pp. 257-266 Bonn, November 2010 A new species of Pachydactylus (Squamata: Gekkonidae) from the Otavi Highlands of northern Namibia Aaron M. Bauer Department of Biology, Villanova University, 800 Lancaster Avenue, Villanova, Pennsylvania 19085, USA; E-mail: aaron.bauer@villanova.edu Abstract. A new species of the “northwestern clade” of Pachydactylus 1s described from the Otavi Highlands of north- eastern Namibia. It is distinguishable from all other members of this clade and from the superficially similar members of the Pachydactylus serval/weberi group on the basis of its inclusion of the rostral in the nostril rim, the possession of a maximum of only four undivided scansors beneath the digits of the pes, is 16 rows of strongly keeled, rounded, juxta- posed dorsal trunk tubercles, its projecting, keeled, lanceolate caudal tubercles, and its complex dorsal trunk patterning. Its probable closest relative is P. otaviensis, also form the Otavi Highlands. These are the only known endemic reptiles from this dolomitic area and their existence points both to an unappreciated area of diversity and endemism in northeast- ern Namibia and to the need for additional herpetological work in even well-known parts of the country. Key words. Gekkonidae, Pachydactylus, Namibia, new species. INTRODUCTION Pachydactylus Wiegmann, 1834 is the most species-rich genus of geckos in southern Africa, with more than 50 species currently recognized (Bauer & Lamb 2005; Bauer et al. 2006a, 2006b; Branch et al. 2010). Although all parts of the subcontinent are inhabited by members of this group, the arid zones of Namibia the adjacent portions of the Northern Cape Province of South Africa have the high- est diversity. A minimum of 35 species of Pachydactylus occur in the Republic of Namibia alone, the majority of which are endemic (Branch 1998; Bauer et al. 2002, 2006a; Branch et al. 2010). Most of these fall into one of two species-rich clades that have been previously identi- fied: the Pachydactylus serval/weberi group and the “northwestern clade” of Pachydactylus (sensu Bauer & Lamb 2005). Most species in both clades are relatively small-bodied, rupicolous species. Most members of the P. serval/weberi group are restricted to southern Namibia and the Northern Cape, with the greatest richness along the lower Orange River Valley and in the Karasberg Moun- tains, where up to five members of the group occur sym- patrically. However, five members assigned to the group have been found north of 21°S latitude: P fasciatus Boulenger, 1888 — widely distributed in northwestern Namibia east of the Namib and north of the Swakop Riv- er, P. waterbergensis Bauer & Lamb, 2003 — endemic to the immediate vicinity of the Waterberg Plateau. P. tsodiloensis Haacke, 1966 — in the Tsodilo Hills of Bonn zoological Bulletin 57 (2): 257-266 northwestern Botswana, and P. otaviensis Bauer, Lamb & Branch, 2006 and an undescribed species (‘Pachydacty- lus sp. 2’, Bauer et al. 2006a) — both from the Otavi High- lands (Otaviberge) of northeastern Namibia. Bauer & Lamb (2005) and Bauer et al. (2006a) used molecular phy- logenetic data to confirm that the first three of these species comprise a monophyletic group that is sister to the rest of the P. serval/weberi group. However, recent mul- ti-gene phylogenetic analyses incorporating all but one of the recognized species of Pachydactylus (Heinicke, Jack- man & Bauer, unpublished) have demonstrated that P. otaviensis is not a member of the P. serval/weberi clade (these phylogenetic results will be presented in their en- tirety elsewhere), but rather part of the “northwestern clade”, which otherwise comprises ten morphologically diverse species that are widely distributed in Namibia and southern Angola, with a single species, P.- punctatus Pe- ters, 1854, extending southwards into South Africa and east to the Indian Ocean coast of Mozambique (Bauer & Branch 1995). Excluding P. punctatus, P. otaviensis has the easternmost distribution of any member of the “northwestern clade”, being known only from the farms Uithoek and Varianto, both in the Tsumeb District, Oshikoto Region in the east- ern Otavi Highlands (quarter degree square 1917Bc; Fig. 1). A second species from the Otavi Highlands was sig- ©ZFMK bo Nn oo Aaron M. Bauer Table 1. Mensural and labial scale data for the type series of Pachydactylus boehmei sp.n. Abbreviations as in Materials and Me- thods, all measurements in mm. Holotype MCZ R184884 MCZ R184880 Sex female female SVL 44.4 43.21 ForeaL 6.6 6.8 CrusL 7.9 79 TailL (total) 44.5 6.9 TailL (regen.) = BR TailW 4.8 4.7 TrunkL 21.0 18.8 HeadL 13.7 14.1 HeadW 8.3 8.3 HeadD 5.0 5.3 OrbD 315 3.8 EyeEar 3.4 Bill SnEye 4.9 5.0 NarEye Sai) 329. Interorb 44 4.3 EarL 1.0 0.9 Internar 1.4 1.1 Supralab. (L/R) 10/9 11/10 Infralab. (L/R) TT 8/8 Paratypes MCZ R184881 MCZ R184882 MCZ R184883 female male female 44.0 35.4 34.8 6.6 5.4 4.7 8.6 ae) 5.9 39.0 35.8 30.3 33.4 — 21.6 47 3.9 3.5 19.0 15.6 13.8 13.0 11.5 12.0 9.0 6.9 V2 5.3 3.8 4.0 3.8 2.8 3.3 3.2 2.9 2.5 4.7 4.2 4.2 3.6 2.9 3.0 3.9 3.6 3.0 1.0 0.7 0.7 1.3 1.0 1.2 10/10 9/9 10/10 7/8 8/8 9/8 naled by Bauer et al. (2006a) as “Pachydactylus sp. 2”, but was not described as it was known only from one ju- venile and one hatchling, making meaningful comparisons with other species difficult. However, Bauer et al. (2006a) noted that it exhibited some features shared with the PR. weberi complex sensu stricto and others with the P. serval complex, and that it possessed a unique and di- agnostic juvenile color pattern. Subsequent field work on the Farm Uisib has yielded a series of adult specimens of this species, permitting its description. Ongoing molecu- lar phylogenetic work verifies that it too is, in fact, cor- rectly assigned to the “northwestern clade” of Pachydact- lyus. MATERIALS AND METHODS The following measurements were taken with Brown and Sharpe Digit-cal Plus digital calipers (to the nearest 0.1 mm) as per Bauer et al. (2006a): snout-vent length (SVL; from tip of snout to vent), crus length (CrusL; from base of heel to knee); tail length (TailL; from vent to tip of un- regenerated tail), tail width (TailW; measured at base of tail); axilla to groin length (TrunkL); head length (HeadL; distance retroarticular process of the jaw and snout-tip), head width (HeadW; measured at angle of jaws), head depth (HeadD; maximum height of head, from occiput to Bonn zoological Bulletin 57 (2): 257-266 throat), ear length (EarL; longest dimension of ear); fore- arm length (ForeaL; from base of palm to elbow); orbital diameter (OrbD), nostril to eye distance (NarEye; distance between anteriormost point of eye and nostril), snout to eye distance (SnEye; distance between anteriormost point of eye and tip of snout), eye to ear distance (Eye- Ear; distance from anterior edge of ear opening to poste- rior corner of eye), and interorbital distance (Interorb; shortest distance between left and right superciliary scale rows). Scale counts and external observations of morphology were made using a Nikon SMZ-1000 dissecting micro- scope. Comparisons were made with museum material (see Appendix) representing all species in the Pachydacty- lus serval/weberi group and the “northwestern clade” of Pachydactylus (sensu Bauer & Lamb 2005; Bauer et al. 2006a). Standard codes for museum collections follow Leviton et al. (1985) except as noted (*): California Acad- emy of Sciences, San Francisco (CAS), Museum of Com- parative Zoology, Harvard University, Cambridge, USA (MCZ), National Museum of Namibia, Windhoek (NMN*), Naturmuseum und Forschungsinstitut Sencken- berg, Frankfurt am Main (SMF), South African Museum, Cape Town (SAM), Transvaal Museum, Pretoria (TM), Zoologisches Forschungsmuseum Alexander Koenig, Bonn (ZFMK). OZFMK A new Pachydactylus from northern Namibia 259 Fig. 1. Map of Namibia and surrounding countries illustrating the type locality of Pachydactylus boehmei sp. n. (red star) and of the related P. ofaviensis (blue circle) in the Otavi Highlands of northeastern Namibia. Satellite image from NASA MODIS sensor (available at http://visibleearth.nasa.gov). RESULTS Pachydactylus boehmei sp. n. Pachydactylus sp. 2 Bauer, Lamb & Branch (Bauer et al. 2006a: 684) Holotype. MCZ R184884 (Figs 2—3): adult male; Namib- ia, Otjozondjupa Region, Grootfontein District, Farm Uis- ib, 19°33’06”S, 17°14711”E, 1400 m a.s.l. coll. A.M. Bauer, J. Marais, T. Jackman, and W.R. Branch, 15 Sep- tember 2006. Paratypes. MCZ R184880—81 (adult females), 184883 (subadult/adult female), MCZ R184882 (subadult/adult male), same data as holotype. Bonn zoological Bulletin 57 (2): 257-266 Additional material. TM 84999, 85005; Namibia, Otjo- zondjupa Region, Grootfontein District, Farm Uisib (IGS WS, LIN), Diagnosis. Snout-vent length to at least 44.4 mm. A mod- erate-sized Pachydactylus with a depressed body form. Trunk with 16 rows of enlarged, keeled tubercles, grad- ing into prominent conical scales on flanks (Figs 2-4). Or- bital diameter as great as eye-ear distance. Rostral partic- ipating 1n nostril rim. Dorsal surface of thighs and shanks covered by enlarged conical to keeled scales. Tail with keeled lanceolate tubercles restricted to one scale row per tail segment. Dorsal pattern with an occipital-nuchal loop, a “V”-shaped band on posterior of neck, a transverse bar anterior to hindlimb insertion, and a series of oval mark- ings or fusions thereof on the trunk (Figs 2—5). Among its congeners P. boehmei sp n. is superficially sim- ilar to some members of the P. weberi group, but can be distinguished from these by its inclusion of the rostral in the nostril rim and the possession of a maximum of only four undivided scansors beneath the digits of the pes (ver- sus at least five on some digits). Among other members of the “northwestern clade” of Pachydactylus it may be differentiated from P. bicolor Hewitt, 1926, P. punctatus, P. scherzi Mertens, 1954, and P. caraculicus FitzSimons, 1959 by its tuberculate (versus atuberculate) dorsum, from P. angolensis Loveridge, 1944 by its inclusion of the ros- tral and first supralabial in the nostril border (versus both excluded), from P. oreophilus McLachlan & Spence, 1967 by its smaller size (maximum SVL < 45 mm versus 57 mm), and lower number of subdigital lamellae (4 versus 5—6 undivided lamellae), from P. gaiasensis Steyn & Mitchell, 1967 by its smaller size (maximum SVL < 45 mm versus 68 mm), lower number of subdigital lamellae (4 versus 5—7 undivided lamellae), longer tail (slightly longer than SVL versus less than SVL), and lack of a ver- tebral stripe, from P. sansteynae Steyn & Mitchell, 1967 by its much larger dorsal tubercles (4-10 times larger than other dorsal scales versus less than twice size of dorsal granules) and presence (versus absence) of tubercles on the parietal region, from P. parascutatus Bauer, Lamb & Branch, 2002 by its larger size (to 44.4 mm versus < 40 mm SVL) and presence of a pale dorsal collar (versus no collar), and from P. scutatus Hewitt, 1927 by is juxtaposed (versus imbricating) keeled dorsal scales, enlarged coni- cal (versus small and granular) flank scales, projecting lanceolate (versus flattened and rounded to oval) caudal tubercles, and its complex dorsal trunk patterning (versus patternless or with small, scattered dark markings. Pachy- dactylus boehmei sp n. is most similar to the geographi- cally proximal P. otaviensis, but may be distinguished from this form by its inclusion (versus exclusion) of the rostral in the nostril rim, the presence of 4 (versus 5) lamel- lae beneath digit IV of the pes, 16 (versus 18) longitudi- ©OZFMK 260 Aaron M. Bauer Fig. 2. Holotype of Pachydactylus boehmei sp. n., MCZ R184884. Scale bar = 10 mm. nal rows of keeled dorsal tubercles, and differences in col- or pattern. Description of holotype. Adult female. Snout-vent length (SVL) 44.4 mm. Body relatively depressed, elon- gate (TrunkL/SVL ratio 0.46). Head elongate, large (HeadL/SVL ratio 0.31), relatively narrow (HeadW/HeadL ratio 0.61), depressed (HeadH/HeadL ra- tio 0.36), distinct from neck. Lores inflated; interorbital region flat. Snout short (SnEye/HeadL ratio 0.36, longer than eye diameter (OrbD/SnEye ratio 0.71); scales on Bonn zoological Bulletin 57 (2): 257-266 snout enlarged, smooth, slightly domed, roughly hexag- onal; scales on snout much larger than those of interor- bital region and parietal table. Eye moderately large (OrbD/HeadL ratio 0.25); orbits without extra-brillar fringes; posterior superciliary scales bearing five small spines; pupil vertical, with crenelated margins. Ear open- ing oval, small (EarL/HeadL ratio 0.07), round; eye to ear distance approximately equal to diameter of eyes (Eye- Ear/OrbD ratio 0.97). Rostral approximately 50% as deep (0.9 mm) as wide (1.9); no rostral groove; contacted by two enlarged supranasals and first supralabials; nostrils oval, each surrounded by two postnasals, one supranasal, first supralabial, and rostral; supranasals in broad contact; dorsal postnasals separated by two granules from one an- other; nostril rims weakly inflated; 1—2 rows of scales sep- arate orbit from supralabials; mental rectangular, only slightly wider anteriorly than posteriorly, approximately 1.6 times deeper (1.8 mm) than wide (1.1 mm); no en- larged postmentals or chin shields. Enlarged supralabials to angle of jaws 9(R)—10(L), 8 to mid-orbit, several gran- ular scales along labial margin to rictus; enlarged infral- abials 7; interorbital scale rows between superciliary scale rows (at midpoint of orbit) 30, 8 across narrowest point of frontal bone. Enlarged conical tubercles present from posterior border of orbit and occiput posteriorly; dorsal trunk tubercles large (4-10 times size of adjacent scales), rounded, with a strongly developed median keel, forming approximate- ly 16 longitudinal rows; tubercles largest on dorsolateral surfaces of trunk, smaller along vertebral midline, and grading into enlarged conical scales on flanks; each en- larged tubercle surrounded by rosette of smaller pyrami- dal scales, some also keeled, larger keeled tubercles typ- ically separated from one another by a single smaller scale; ventral scales flattened, subimbricate, becoming somewhat larger posteriorly, approximately 40 between lowest tu- bercular rows at midbody; non-tuberculate scales on dor- sum at midbody similar in size to those on ventrum at same level; gular granules less than one half size of ven- tral scales of chest, increasing abruptly in size on throat. No precloacal or femoral pores. Scales on palm, sole, and ventral surface of forelimb small, smooth, granular, jux- taposed; scales on ventral aspect of hindlimbs smooth, jux- taposed to subimbricate; scales on dorsal aspect of fore- limb heterogeneous, with midsized conical to keeled tu- bercles intermixed with smaller granular to conical scales; scales on dorsum of thigh and crus greatly enlarged, conical and keeled, in contact with each other or narrow- ly separated by much smaller interscales. Forelimbs moderately short, stout; forearm short (Fore- aL/SVL ratio 0.15); hindlimbs relatively short, tibia mod- erately short (CrusL/SVL ratio 0.18); digits relatively short, claws minute, stylet-like, visible only with difficul- ©ZFMK A new Pachydactylus from northern Namibia 261 <== —— = - Sea ee es ae ; Fig. 3. Life photo of holotype of Pachydactylus boehmei sp. n. Photo by Johan Marais. ty on some digits of the pes; subdigital scansors, except for distalmost, entire, present only on distal portion of toes, approximately 1.5 times wider than more basal (non-scan- sorial) subdigital scales; interdigital webbing absent. Rel- ative length of digits (manus): HI > IV > II > V > I; (pes): IV > ll > V > II >I. Subdigital scansors, exclusive of di- vided distalmost scansor (manus): I (4), II (4), II (4), 1V (4), V (4); (pes) I (4), II (4), HI (4), TV (4), V (4). Tail sub-cylindrical, clearly depressed; original tail approx- imately snout-vent length (TailL/SVL ratio 1.00); tail con- stricted basally, then expanded before tapering towards tip, distinctly segmented; each segment with 5 rows of scales dorsally and 3 ventrally, dorsal caudal , tail segment; cau- dal tubercles heterogeneous, medial tubercles more-or-less recumbent, lateral tubercles projecting, up to 8 keeled tu- bercles per row basally, decreasing to 4 on distal caudal segments; subcaudal scales smooth, imbricating, oval to rectangular; no enlarged postcloacal spurs on side of tail- base. Coloration. In preservative (Fig. 2): Ground color of dor- sum straw to yellowish brown with mid-brown markings. A broad “V”-shaped nape band and a transverse band an- terior to hindlimb insertion. Trunk bearing a series if ir- regular oval markings, darker on their edges than at their centers, 3 (left) and 4 (right) markings in paravertebral po- sition, 6 on upper left flank, last fused with transverse band, 2 on upper right flank followed by an irregular lon- gitudinal marking representing the fusion of several oval markings. An additional pair of small dark markings at an- terior face of hindlimb insertion and an additional cross- band on dorsum of posterior sacrum. Bonn zoological Bulletin 57 (2): 257-266 Head with a pale stripe from nostril to anterodorsal rim of orbit. Dark stripe along loreal region to mid-anterior of orbit, continuing from midposterior of orbit, above ear, to meet contralateral stripe to form a complete loop be- tween the occiput and nape. Crown mottled, a triangular brown marking with apex at supranasals scales extending back to anterodorsal orbital rim. Labial scales pale with diffuse speckling; grayish vertical markings on lateral edges of rostral. Limbs mottled with irregular markings. Tail with alternat- ing irregular bands of grayish-brown and mid-brown, 20 dark bands including tail tip; most caudal tubercles cream to beige. Body venter beige, soles and palms grayish, tail venter grayish-brown with irregular darker gray-brown markings scattered along length of tail. In life (Fig. 3): Background color of dorsum a pale pink- ish-gray. Labial scales, canthal stripe, and nape whitish. Dark markings yellowish-to mid brown, darkest on head and occiput. Venter white. Variation. Variation in mensural characters of the holo- type and paratypes are presented in Table 1. All paratypes share with the holotype the same number of longitudinal rows of dorsal tubercles, number of subdigital lamellae, and configuration of the scales of the nasal region. Labi- al scale numbers varied across the type series and are al- so presented in Table 1. The male paratype, MCZ R184882 has prominent precloacal spurs (Fig. 4A), each bearing two rows of enlarged, compressed, dorsally-direct- ed scales. Those of the dorsal row (5 scales on both sides) larger than those of ventral row (5 scales left, 6 scales ©ZFMK 262 Aaron M. Bauer Fig. 4. right). Color pattern variable amongst paratypes (Figs 4-5). Dark occipital and nape bands thinner in MCZ R184880, R184882 than in holotype. Dorsal oval patterns largely replaced by coalescent blotches and lines except in MCZ R184881. Dorsal pattern weakly contrasting in MCZ R184883. Etymology. Named for Prof. Dr. Wolfgang BOhme (born 21 November 1944), my longtime friend and colleague and a leading contributor to African herpetology. It is a privilege to apply this patronym to a species of one of the continent’s dominant genera on the occasion of his nom- inal retirement from his position at the Museum Alexan- der Koenig. The epithet is formed in the masculine gen- itive. Distribution. The species is known only from Farm Uis- ib in the Grootfontein District of northeastern Namibia (Fig. 1). This lies in the western portion of the Otaviberge or Otavi Highlands, 15 km northwest of the town of Otavi. The distribution of P. boehmei sp n. in the region is un- known and the closely related P. otaviensis occurs only 50 km to the northeast. These two geckos are relatively isolated from other members of the “northwestern clade” of Pachydactylus except the ubiquitous P. punctatus; the nearest known localities for P. bicolor and P. scutatus be- ing more than 200 km distant. Other rock-dwelling con- geners in other clades are also quite remote, with P. wa- terbergensis approximately 125 km to the south and P. tsodiloensis almost 400 km to the north-northeast. The Bonn zoological Bulletin 57 (2): 257-266 Life photos of paratypes of P. boehmei sp. n. (A) MCZ R184882, male — note the raised precloacal spur visible lateral to the tail base. (B) MCZ R184880, female — note the transition from rounded, keeled dorsal tubercles to enlarged conical flank scales. Photos by Johan Marais. Otavi Highlands as a whole has been poorly explored her- petologically and may harbor other isolated populations and/or endemic species of lizards. A number of endemic invertebrates and fish are already known from the Otavi- Tsumeb-Grootfontein area (Barnard et al. 1998). Natural history. The area where P. boehmei sp n. occurs is characterized as mountain savanna and karstveld (Giess 1971). The type series was collected in broadleaf savan- na on rocky dolomite hills (Fig. 6). Specimens collected by the author and colleagues were moving on rock faces or were found in large crevices or cracks between 22:30 and 00:30. The two Transvaal Museum specimens (see Additional material) referred to this species were collect- ed in the course of searching for scorpions (E. Scott & L. Prendini, pers. comm.). Barnards Namib day gecko, Rhop- tropus barnardi Hewitt, 1926, was also collected at Farm Uisib, which is one of the easternmost localities for any member of its genus. Other species observed at the type locality were the widespread Chondrodactylus turneri (Gray, 1864), Zrachylepis sulcata (Peters, 1867), and 7: punctulata (Bocage, 1872). Lygodactylus capensis (Smith, 1849) was collected at the nearby Uisib farmhouse (192332 Sa 7 ens 00uE): Two enlarged eggs are visible through the ventral body wall of the holotype collected in mid-September, suggest- ing spring breeding and hatching late in the year, corre- sponding to the rainy season. Trombiculid mites were found on the specimens, most notably in between the ©ZFMK A new Pachydactylus from northern Namibia 263 Fig. 5. scales of the tail base. In the male paratype, MCZ 184882, the infestation of mites around the tail base and scales of the precloacal spurs was particularly severe. Phylogenetic affinities. Pachydactylus boehmei sp n. is similar in habitus to the other small-bodied, tuberculate members of the “northwestern clade”. It is superficially most similar to the neighboring species P. ofaviensis, al- though the latter species lacks the rostral-nostril contact that is typical for most members of the clade. Preliminary molecular results suggest that these two species are indeed sister taxa. Conservation status. Pachydactytlus boehmei sp n. does not occur in any protected areas. At its type locality it is undisturbed and the jagged, rocky terrain precludes hu- man encroachment into its specific habitat. However, de- pending upon the extent of its actual range it may be un- der some threat from local mining activity in some places. Until such time as the species’ distribution and threats can be evaluated more fully, I recommend that it be consid- ered Data Defficient under the IUCN threat category sys- tem. Bonn zoological Bulletin 57 (2): 257-266 Paratype series of Pachydactylus boehmei sp. n. showing variation in the dorsal color pattern and degree of pattern bold- ness. Scale bar = 10 mm. DISCUSSION The discovery of this apparently range-restricted species highlights Namibia’s high biodiversity and endemism (Maggs et al. 1998). M. Griffin (1998) identified 55 rep- tile species as being strictly or primarily endemic to Namibia, but recent discoveries, particularly in Pachy- dactylus (Bauer et al. 2002, 2006a; Bauer & Lamb 2003) have increased this to approximately 70. The “northwest- ern clade” of Pachydactylus is particularly diverse along the Northern Namibian Escarpment (sensu Irish 2002), which corresponds roughly to the Kaokoveld center of Floral Endemism (Volk 1966; van Wyk & Smith 2001) and is recognized as a regional center of endemism for rep- tiles in general (Crowe 1990; Simmons et al. 1998; Grif- fin 2000). The Otavi Highlands have also been ranked as an area of high biodiversity importance (Irish 2002), but like the Waterberg to the south, the relatively low relief (a maximum of 2155 m in surrounding plains of 1200-1500 m) and accessibility to surrounding areas that promotes diversity also decreases the prospects for long- term isolation and, consequently, endemism. Thus, it is somewhat surprising that two species of Pachydactylus, ©OZFMK 264 Aaron M. Bauer Fig. 6. P. otaviensis and P. boehmei sp n., appear to be restrict- ed to this region. Bauer (1999 [2000]) emphasized the role of substrate specificity as a cladogeneic agent in Pachy- dactylus and it seems likely that dependence on microhab- itats provided by the dolomite outcrops of the Otaviberge has isolated these species from rest of the “northwestern clade”. Other groups of organisms that respond similarly to historical ecological conditions should be expected to show similar patterns of endemism and indeed this is the case in scorpions (R.E. Griffin 1998), which include many substrate specific rupicolous species, such as the bothri- urid Lisposoma josehermana Lamoral, 1979, which is largely restricted to the Otavi Highlands (Prendini 2003, 2005). Despite over 50 years of relatively intense study (e.g., Mertens 1955, 1971; Haacke 1965; van den Elzen 1978; Bauer et al. 1993; Griffin, 2000, 2003), novel herpetolog- ical taxa continue to be discovered in Namibia on a reg- ular basis. That Pachydactylus boehmei sp n. occurs in a densely-populated (by Namibian standards) agricultural district with excellent road access demonstrates that even “well known” parts of the country remain understudied. Acknowledgements. | thank the many colleagues and students who have accompanied me on trips to Namibia, and in particu- lar Johan Marais, Todd Jackman and Bill Branch, who partici- pated on the field trip on which the types of P boehmei were collected. I am also grateful to Andre Schoeman and his fami- ly, who made us welcome at Farm Uisib, and to the Ministry of the Environment and Mike Griffin, who have supported my work in Namibia for more than 20 years. Specimens were collected under Namibian Research/Collecting Permit 1068/2006. For ac- cess to comparative material I thank José Rosado, Jonathan Losos and James Hanken (MCZ), Lauretta Mahlengu and Wulf Haacke (TM), Jens Vindum (California Academy of Sciences), Mathilda Awases (NMN), Wolfgang B6hme (ZFMK), and Gun- ther K6hler (SMF). Photos were kindly provided by Johan Marais and Elizabeth Scott. This research was funded by the Na- Bonn zoological Bulletin 57 (2): 257-266 Habitat of Pachydactylus boehmei sp. n. in the Otavi Highlands: (A) View of typical dolomite hill. (B) Broken dolomi- te providing cracks and fissures as retreat sites for geckos. Photos by Elizabeth Scott. tional Science Foundation of the United States through grants DEB 0515909 and DEB 0844523 to the author. Finally, I thank Philipp Wagner for inviting me to submit this manuscript, and Wolfgang Bohme, whose illustrious career has provided the op- portunity to present these data. REFERENCES Barnard P, Bethune S, Kolberg H (1998) Biodiversity of terres- trial and freshwater habitats. Pp. 57-188 in Barnard P (ed) Bi- ological Diversity in Namibia — A Country Study. Namibian National Biodiversity Task Force, Windhoek Bauer, AM (1999 [2000]) Evolutionary scenarios in the Pachy- dactylus-group geckos of southern Africa: new hypotheses. 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Proceed- ings of the California Academy of Sciences 53: 23-36 Bauer AM, Lamb T, Branch WR (2006a) A Revision of the Pachydactylus serval and P. weberi groups (Reptilia: Squa- mata: Gekkonidae) of Southern Africa, with the description of eight new species. Proceedings of the California Academy of Sciences 57: 595-709 Bauer AM, Barts M, Hulbert F (2006b) A new species of the Pachydactylus weberi group (Reptila: Squamata: Gekkonidae) from the Orange River, with comments on its natural history. Salamandra 42: 83-92 Branch WR (1998) Field Guide to the Snakes and Other Rep- tiles of Southern A frica, 3" ed. Struik Publishers, Cape Town, 399 pp ©OZFMK A new Pachydactylus from northern Namibia 265 Branch WR, Bauer AM, Jackman TR, Heinicke M (2010) A new species of the Pachydactylus weberi complex (Reptilia: Squa- mata: Gekkonidae) from the Namib-Rand Reserve, southern Namibia. Breviora xx: in press. Crowe TM (1990) A quantitative analysis of patterns of distri- bution, species richness and endemism in southern African vertebrates. Pp. 145—160 in Peters G & Hutterer R (eds.) Ver- tebrates in the Tropics. Zoologiches Forschungsinstitut und Museum Koenig, Bonn Giess W (1971) A preliminary vegetation map of South West Africa. Dinteria 4: 5-114 Griffin M (1998) Reptile diversity. Pp. 148-154 in Barnard P (ed) Biological Diversity in Namibia — A Country Study. Namibian National Biodiversity Task Force, Windhoek Griffin M (2000) The species diversity, distribution and conser- vation of Namibian reptiles: a review. Namibia Scientific So- ciety Journal 48: 116-141 Griffin M (2003) Annotated Checklist and Provisional Nation- al Conservation Status of Namibian Reptiles. Namibia Scien- tific Society, Windhoek, [2] + 169 pp Griffin RE (1998) Species richness and biogeography of non- acarine arachnids in Namibia. Biodiversity and Conservation 7: 467-481 Haacke WD (1965) Additional notes on the herpetology of South West Africa with descriptions of two new subspecies of geck- os. Cimbebasia (11): 1-40 Irish J (2002) Namibian Mountains: Biodiversity Potential Based on Topography. Report to the Mountain Working Group of the National Biodiversity Task Force, Windhoek, 42 pp Leviton AE, Gibbs RH, Heal E, Dawson CE (1985) Standards in herpetology and ichthyology: Part 1, Standard symbolic codes for institutional resource collections in herpetology and ichthyology. Copeia 1985: 802-832 Maggs GL, Craven P, Kolberg HH (1998) Plant species richness, endemism, and genetic resources in Namibia. Biodiversity and Conservation 7: 435-446 Mertens R (1955) Die Amphibien und Reptilien Stidwestafrikas, aus den Ergebnissen einer im Jahre 1952 ausgefihrten Rei- se. Abhandlungen der Senckenbergischen naturforschenden Gesellschaft 490: 1-172 Mertens R (1971) Die Herpetofauna Stidwest-A frikas. Abhand- lungen der Senckenbergischen naturforschenden Gesellschaft 529: 1-110 Prendini L (2003) Revision of the genus Lisposoma Lawrence, 1928 (Scorpiones: Bothriuridae). Insect Systematics and Evo- lution 34: 241-264 Prendini L (2005) New records and observations on the natural history of Lisposoma elegans and L. josehermana (Scorpiones: Bothriuridae). American Museum Novitates 3487: 1-11 Simmons RE, Griffin M, Griffin RE, Marais E, Kolberg H (1998) Endemism in Namibia: patterns, processes and predictions. Biodiversity and Conservation 7: 513-530 van den Elzen P (1978) Observations sur l’herpétofaune du Wa- terberg (Namibie). Bonner zoologische Beitrage 29: 171-182 Van Wyk AE, Smith GF (2001) Regions of Floristic Endemism in Southern Africa. Umdaus Press, Hatfield, South Africa, viii + 199 pp Volk OH (1966) Die Florengebiete von Siidwestafrika. Journal der Stidwest Afrikanische Wissenschaftliche Gesellschaft 20: 25-58 Bonn zoological Bulletin 57 (2): 257-266 APPENDIX Comparative specimens examined See Bauer et al. (2006ab) for a list of P weberi group, P. otavien- sis, and P. sansteynae specimens examined, Bauer & Branch (1995) for a list of P. punctatus and P. scherzi examined, and Bauer et al. (2002) for a list of P angolensis, P. scutatus and P. parascutatus examined. Only specimens not included in these publications are listed below. For localities without precise co- ordinates quarter degree square (QDS) references have been pro- vided when possible. Each single degree square is subdivided into four quarter degrees, designated A-D (A=NW quadrant, B=NE quadrant, C=SW quadrant, D=SE quadrant). Each quar- ter degree is in turn divided into four similarly designated divi- sions, yielding a basic unit one sixteenth of a degree square, or one quarter of a degree on a side (e.g., 1915Ac represents the unit bounded by 19°15’S and 19°30’S and 15°00’S and 15°15E). All QDS references in this paper refer to degrees South and East. P. angolensis: Angola: Namibe Province: San Nicolau (1412Ab): TM 25454—55, 25459, 25476, 25478—79; Saco de Gi- raul: TM 4032830, TM 46558; Lucira (1312Dc): TM 41172, 24406, 24445, 24449; Lungo: TM 24406; Benguela Province: Hanha, 20 km N of Lobito (1213Ab); 24 km S Benguela: TM 39110—11; 30 km N of Dombe Grande: TM 41266. P. bicolor: Namibia: Erongo Region: Karibib District: 47 mi E Hentiesbaai: CAS 126210; 20 km W Karibib: MCZ R163282—83; Swakopmund District: 29.0 km N of Swakop- mund (22°25°38"S, 14°27°53”E): CAS 214576; 31.1 km N Swakopmund on Hentiesbaai Rd. (22°25’42”S, 14°27°43”E): MCZ R184218-20, 184225; Messum Crater (21°26’25.8’S, 14°13°12.9"E): CAS 214814; Kunene Region: Khorixas Dis- trict: Torrabaai Rd, 63.4 km W of Kamanjab (19°41’00"S, 14°19°10”E): CAS 223912-15; Torrabaai Rd, 58 km W of Ka- manjab (19°39°20"S, 14°21°10”E): CAS 223927-28; Torrabaai Rd, 56.7 km W of Franken entrance: CAS 176284-85: Torrabaai Rd, 37.8 km W of Franken entrance: CAS 176292—93: Kaman- jab-Torrabaai Rd, Grootberg Pass (19°50.584’S, 14°07.696’E): CAS 193675; Kamanjab-Torrabaai Rd, 59.3 km W of Kaman- jab (19°39.100’S, 14°21.335’E): CAS 193680; Torra Bay Rd, 48 km W of Kamanjab, (19°39714"S, 14°21°03”E): CAS 214661—63; Torra Bay Rd, 68 km W of Kamanjab, (19°43’00"S, 14°18°40”E): CAS 21468-89; Torra Bay Rd, 74.2 km W of Ka- manyab (19°45’°40"S, 14°17°03”E): CAS 21469396; E side of Road 3706, 19.3 km N of entrance to Palmwag: CAS 175344; Torrabaai Rd, 108.9 km W of Kamanjab: CAS 176101—08; Torrabaai Rd, 101.7 km W of Kamanjab: CAS 176116—19; Torrabaai Rd, 95.9 km W of Kamanjab: CAS 176126-31; Torrabaai Rd, 44 km W of Kamanjab: CAS 176151; Henties- baai-Uis Rd, 24 km W of Hwy C35 (21°18717”S, 14°35714”E): CAS 206959; 25 km E of Grootberg Pass: CAS 206960; ~60 km W. Kamanjab on Torra Bay Rd. (19°40°57”S, 14°19°09”E): MCZ R184919; 31.9 km E Grootberg Pass (19°40°57S, 14°19°09”E): MCZ R184197—98; 62.8 km W Kamanjab Rest Camp on Rd. to Grootberg Pass (19°40’56”S, 14°19°08”E): MCZ R185753-—55; 67.5 km W Kamanyjab on Torrabaai Rd. (19°43’00"S, 14°18°44”E): MCZ R183766; Hobatere Lodge, 2.5 km from main gate (19°18’07"S, 14°27°26”E): MCZ R184934—35; Opuwo District: Kamanjab-Ruacana Rd, 98.4 km N of Kamanjab: CAS 193719; Opuwo-Okangwati Rd, Otjivize (17°37.188 S, 13°27.535 E): CAS 193731; Outjo District: 17 mi S of Outjo: CAS 85944; Farm Franken: CAS 175347-S3, 175360-74, 176066—68, 176176—-77; Farm Franken, Haus Franken: CAS 176261—62; Farm Franken, vic. Haus Franken: ©ZFMK 266 Aaron M. Bauer CAS 176278; 62.0 km E Kamanjab, Farm Amolinda (19°48°29"S, 15°22°46”E): MCZ R185745—47; Kamanjab Rest Camp, 3 km W Kamanjab(19°37’48"S, 14°48°57”E): MCZ R184887, 18489497. P. caraculicus: Angola: Namibe Province: 36 mi. northwest of Mocamedes [Namibe]: CAS 85959; Namibia: Kunene Region: Opuwo District: Okangwati-Epupa Rd, 43.4 km N of Okang- wati: CAS 193799; 193804—05; 41.9 km N. of Okanguati on Epupa Falls Rd: CAS 206980; 32 km S Epupa Falls on Okang- wati Rd. (17°1409”S, 13°13’45”): MCZ R185767. P. gaiasensis: Namibia: Kunene Region: Khorixas District: vic. Gat-as (20°47°18"S, 14°06’44”): CAS 214626—28; 22.4 km N Ugab River on road to Gai-as (20°46°59”S, 14°06731”, 520 ma.s.l.): AMB 7568-69 (NMN), MCZ R184169-70, R184248; Gai-As (20°46°45”S, 14°04°30”E, 520 m): AMB 8484 (NMN), MCZ Z-37873 (NMN), MCZ R184181, R184192—93; Gai-As (20°46 '46'°S,14°04'29""E): MCZ R185967—75, 185979-80; “False Gai-As” (20°47°14.9"S, 14°06°44.6"E): MCZ Z- 37853-54 (NMN), MCZ R184185, R184187; 7 km E Gai-as (20°47’S, 14°07’°E): TM 68962—66; Messem Crater, 21 26.430 S, 14 13.215 E : CAS 214800; Messum Mts. (2114Ac): TM 56346; Farm Twyfelfontein: TM 42182; near Gai-as, ~20 mi N Brandberg: TM 32868—80 [paratypes]. P. oreophilus: Angola: Namibe Province: Caraculo (15°01’S, 12°40°E): TM 245 19-25, 24452; 20 km W Virei: TM 41011—15; Tambor: TM 40532—34: 6 km S Rio Coroca towards Iona: TM 40575—76; Mutiambo River on road to Lucira: TM 41088; Fur- nas: TM 40561—62; Namibe 7 km from Iona towards Oncocau, Iona Reserve: TM 40762; Assuncao: TM 40152; Saiona River, 25 km NW Cainde: TM 40976—77; Benguela Province: 35 km S Dombe Grande towards Lucira: TM 41246; Namibia: Kunene Region: Opuwo District: near Purros (18°46’S, 12°59E); TM 68465-—67; Hoanib River (19°18’S, 13°15’E): TM 64185: Hoanib River, 44 km E Mudorib River (19°18’S, 13°15°E): TM 56889; Epupa Falls (16°59’°S, 13°17’E): TM 38771—72, 71579; 6 km S Ohnborimbonga (1712Bb): TM 49084—85; MarienfluB, 40 mls S Kunene: TM 32532; near Otji- nende, Kaokoveld (1712Db): TM 49219; Epupa (1613Cc): TM 47775; N Okangwati on Epupa Falls Rd. (17°17'24’'S, 13°09°31°E): MCZ R185769; Paracamp, Sesfontein Bonn zoological Bulletin 57 (2): 257-266 (19°07'52’’S, 13°35'17’"E): MCZ_ R-184945-47; Paracamp, Sesfontein (19°07'55”S, 13°35’20”E): MCZ R184290; ca 2 km N of Sesfontein, Para Camp (19°07’28”S, 13°35’29”E): CAS 214736, 214754; ca 4 km N of Sesfontein, Para Campsite (19°07’56”S, 13° 35’18”E): CAS 223919-22. P. otaviensis: Namibia: Oshikoto Region: Tsumeb District: Farm Varianto (19°22°46"S, 17°44’27°E): MCZ R184867. P. parascutatus: Namibia: Kunene Region: Opuwo District: Mudorib River, 12 km from Hoanib River (19°23’S, 13°17’E): TM 68488—92; Okamungodona, 15 km W Orawanji (18°49’S, 13°39°E): TM 71519-21; Otunungwa, Kaokoveld: TM 32401—03; Kharu-gaiseb River (19°45’S, 13°25’E): TM 68517—18; Bottom of Van Zyl’s Pass (1712Da): TM 71497; 37 km N Sesfontein towards Kaoko Otavi (1813Dc): TM 48876-77; Otjiu, Kaokoveld: TM 32358; 32858—S9; 18 miles SW Orupem- be: TM 31494; Otjinungwa, Kaokoveld: TM 32860; 4 km NW Etenga towards Omborombongo (1712Bd): TM 49060; Nango- lo Flats (1712Ad): TM 24322; Otjiunongua (1712Ab): TM 32546; Sesfontein (19°07’S, 13°37°E): TM _ 79078; Ongongo/Kaoko: ZFMK 66434. P. scutatus: Angola: Namibe Province: Iona, Iona Reserve: TM 40751; Espinheira, Iona Reserve: TM 40615—18; 6 km S Rio Coroca, Iona: TM 40577; Namibia: Erongo Region: Omaruru District: Ugab River Bridge near Brandberg W. Mine: TM 36463; Ugab River (20°58’S, 14°12’E): TM 49708; Tsisab Gorge, Brandberg (2114Ba): TM 79286; Brandberg: SMF 58564; Sraussenhohle an der Jochmannswald, Brandberg: SMF 45658; Kunene Region: Khorixas District: Farm Palmwag (19°53’S, 13°53’E): TM 56865; Farm Paderborn (1914Dd): TM 17302; Kamanjab (1914Db): TM 17209, 17270, 36372; Damaraland (20°30’S, 13°49°E): TM 68754; Farm Huab (1914Db): TM 17338; Farm Palmfontein (part of Grootberg): TM 36465; Farm Blauwpoort (2014cb): TM 49419; Agab Spring (20°05’S, 13°50’E): TM 56936; Opuwo District: Epupa Falls (16°59’S, 13°17°E): TM 7135253; Otjiu, Kaokoveld: TM 32539. Received: 04.VII.2010 Accepted: 11.X.2010 ©ZFMK Bonn zoological Bulletin | Volume 57 Issu 2 ia pp. 267-274 Bonn, November 2010 A new Tarentola subspecies (Reptilia: Gekkonidae) endemic to Tunisia Ulrich Joger & Ismail Bshaenia Staatliches Naturhistorisches Museum, Pockelsstr. 10, D-38106 Braunschweig, Germany Abstract. Mitochondrial DNA sequences as well as morphological characters reveal that geckos of the genus Zarentola from Libya and central Tunisia are a monophyletic group which is different from both 7. mauritanica and T. deserti. Con- sequently, we elevate the former subspecies 7. mauritanica fascicularis to species rank. Together with 7) neglecta and T. mindiae, T. fascicularis constitutes the sister group of 7: deserti. Tarentola fascicularis comprises several evolutionary units, one of which we describe here as a subspecies endemic to south-central Tunisia. Key words. Zarentola, gecko, Tunisia, Libya, North Africa, taxonomy. INTRODUCTION The Mediterranean geckos of the genus Zarentola Gray, 1825 are classified in the nominative subgenus 7arento- la. The Canary Islands are inhabited by geckos of the sub- genera Jarentola (Eastern Canaries) and Makariogecko (western Canaries, Selvagens and Cape Verde Islands) whereas other subgenera inhabit sub-Saharan Africa and the Caribbean (Joger 1984a, b). To date, the nominative subgenus comprises the following species: Zarentola mau- ritanica (L., 1758), T. deserti Boulenger, 1891, 7. angus- timentalis Steindachner, 1891, 7. boehmei Joger, 1984. Al- though no subspecies have been described in the latter three species, the North African populations of 7) mauri- tanica have been assigned to a number of subspecies; in- cluding, 7 m. mauritanica, T. m. fascicularis (Daudin, 1802) from Egypt, 7 m. juliae Joger, 1984, and 7: m. pal- lida Geniez et al., 1999, both from Morocco. Zarentola angustimentalis (eastern Canary Islands) and 7: deserti (northern margin of the Sahara Desert) used to be classi- fied as subspecies of 7. mauritanica until they were ele- vated to species rank (Joger 1984b). In the case of 7. de- serti, sympatric records of 7: deserti and T: mauritanica in Tunisia and Algeria gave reason to assume that these were separate biological species. Tarentola deserti and T. mauritanica can be distinguished from each other in a number of characters (Table 1, Figs 1 and 2). In the field, 7. deserti is characterized often by its very large size, pale, often rosy body colour and a yel- lowish or ochre brown coloured iris, whereas typical 7. mauritanica is smaller and has a grey body and iris colouration. Bonn zoological Bulletin 57 (2): 267-274 In contrast, in south-central Tunisia (Bou Hedma Nation- al Park and areas to the north of the Chott al Djerid salt pan) populations of Zarentola were found that show a mix- ture of characters of both species (Table 1, Fig. 3). Their size is smaller than 7. deserti, but body and eye colour are close to 7: deserti (Joger & Bischoff 1989; Joger 2003). A preliminary study of morphological and electrophoret- ic characters (Willand 1997; Joger et al. 1998) showed that these geckos cluster morphologically with 7. mauritani- ca (yet not with any of its described subspecies), where- as their dorsal colour and pattern is close to 7 deserti and their blood plasma protein alleles are distinct and not shared by neither 7) mauritanica nor T. deserti. Previous molecular genetic studies (Carranza et al. 2002; Rato et al. 2010) of North African 7arentola were biased in that they concentrated on Moroccan populations but largely neglected Tunisian and Libyan populations. In this study, we use both morphological and molecular samples of Tunisian and Libyan Jarentola to determine the affini- ties and clarify the taxonomy of the enigmatic Zarentola of south-central Tunisia. MATERIALS AND METHODS Animals were collected during several trips to Morocco, Algeria, Tunisia (U.J.) and Libya (I.B.). Specimens from Egypt were kindly provided by Sherif Baha El Din and Adel Ibrahim. Blood samples were taken by heart punc- ture or from muscle tissue of dead animals and preserved ©ZFMK 268 Ulrich Joger & Ismail Bshaenia Fig. 1. Zarentola mauritanica (Tunisia). Operational taxonomic units (OTUs) were defined using a combination of mitochondrial DNA clades and geo- graphic proximity. Linear Discriminant Function Analy- ses (LDFA) were used to find variables that separate the groups. Principal Component Analyses (PCA) were ap- plied to see whether groups are distinguishable without previous definition of OTUs. Significance of character dif- ferences were tested with t-tests. DNA was extracted from the preserved samples using standard procedures. Universal primers were used to am- plify mitochondrial 12S rRNA (372 bp) and 16s rRNA (2 fragments of 448 and 604 bp). Sequences were determined using an automated sequencer, and aligned with CLUSTAL-W omitting gaps. Sequences from Genbank were added in some cases. A sequence of a paratype of the new subspecies was submitted to GenBank (IB47: Table 1. Distinguishing characters of North African Zarento/a (mean + standard deviation). Significant differences from T. sp. (Tunisia) are highlighted (in bold). Significance values are given for males (first value) and females separately (if different). *«* P<().001; ** P<0.01; * P<0.05; n.s. not significant. characters Timauritanica - Tunisia 7T’sp.- Tunisia T:sp.-complex-Libya T'deserti ssp.- Libya T.d.deserti - N-Africa N=22 N=15 N=139 N=73 N=24 Maximal head 83.5 86 79 Tell 103.3 + body length Lamellae under 11.0 + 1.0 10.3 + 0.8 10.9 + 0.6 n.s./*** 11.0 + 0.7 n.s./*** 1257 E See Ist toe Lamellae under 16.8 + 1.0 *** 15.3 + 1.0 15.8 + 1.0 15.9 + 0.8 n.s./** 18272 eS ees 4th toe Lamellae under 20.5 + 1.5 19.7+1.1 20.2 + 1.6 2053 eal 22:8 Eesha 5th toe Ventral scales 37 £33 34.3 + 2.7 36.6 + 3.2 40.0 + 2.7 *** 40.1 + 2.0 *** Dorsal tubercles 13.5 + 1.0 12.1+ 0.5 12.8 +1.4 13.4 + 0.9 **/*** 12.2+0.7 Gular scales 41.1+4.7 43.3 + 6.6 43.3+4.7 45.3+5.4 56.0 + 6.9 *** Interorbitals 14.9 + 1.2 ** 13.7 + 1.0 14.9 + 0.7 **« 14.6 + 0.9 ** 14.5 +0.7 Infralabials 8.0 + 0.7 7.9 + 0.7 7.9 + 0.5 8.0 + 0.5 8.8 + 0.6 *** Supralabials 7.8 + 0.6 PaO 7.9 +0.5 8.0 + 0.4 8.0 + 0.4 Relative 0.38 + 0.03 0.42 + 0.04 0.36 + 0.04 *** 0.36 + 0.04 *** 0.43 + 0.06 inter-orbital 0.39 + 0.02 0.41 + 0.03 0.37 + 0.04 n.s. 0.37 + 0.05 * 0.42 + 0.04 width Relative earto 0.84+40.03 0.91 + 0.03 0.66 + 0.07 *** 0.66 + 0.06 *** 0.89 + 0.04 mental distance 0.88 + 0.06 0.90 + 0.03 0.69 + 0.21 ** 0.68 + 0.09 *** 0.88 + 0.03 in 96% ethanol. Voucher specimens were preserved in 80% ethanol. Morphological data were taken as described by Joger (1984a). Measurements were taken to the nearest 0.1 mm and normalized as proportion to body length. Pholidotic counts were taken unaltered. Bonn zoological Bulletin 57 (2): 267-274 HQ437282). Phylogenetic trees were constructed using a Bayesian Markov Chain Monte Carlo inference applying an evolution model suggested by MODELTEST 3.7. Sta- tistical support for branches was indicated by posterior probability values (MrBayes). ©ZFMK A new Zarentola endemic to Tunisia 269 RESULTS Morphological comparisons CDF plots (Figs 4a, b) show that 7. sp. (Tunisia) are dif- ferent morphologically from 7 deserti as well as from un- described Zarentola from western Libya (‘7T. sp. com- plex’). When North African populations of 7) mauritani- ca are compared with 7. sp. (Tunisia), only males appear distinct, whereas females cluster with 7 mauritanica from Tunisia and Morocco (Fig. 5b). Fig. 2. Zarentola deserti (Biskra, Algeria). Significantly different characters which distinguish Tunisian 7arentola from Libyan and other North African populations are shown in Table 1. Molecular genetic affinities Mitochondrial gene sequences (12S rRNA, 16s rRNA) re- veal that 7. sp. (Tunisia) does not cluster with 7? mauri- tanica but with undescribed Libyan Tarentola (Fig. 8). These Libyan populations form several geographically re- stricted, monophyletic clades; the most western ones are sister to 7’ sp. (Tunisia) — yet with rather low statistical support. Zarentola deserti, T. mauritanica fascicularis (Egypt, East Libya), and even 7. neglecta appear more closely related to the western Libyan-Tunisian group of clades than 7. m. mauritanica. This 1s highly supported statistically (1.00 posterior probability). It is noteworthy that the populations near the (neo-) type locality of 7! m. Fig. 3. Tarentola sp. (Bou Hedma, Tunisia). Table 2. Uncorrected “P” distance between main clades, estimated of evolutionary divergence between sequences, based on pair- wise analysis of 1433 bp mtDNA sequences. Clade A Clade B Clade C Clade D Clade E Clade F Clade G Clade H Clade A Clade B 0.0921 Clade C 0.0972 0.0704 Clade D 0.1180 0.0977 0.0692 Clade E 0.1052 0.0853 0.0562 0.0705 Clade F 0.0974 0.0634 0.0630 0.0802 0.0645 Clade G 0.1145 0.0873 0.0662 0.0665 0.0694 0.0510 Clade H 0.1141 0.0886 0.0691 0.0711 0.0706 0.0578 0.0270 Outgroup 0.1234 0.1407 OMB Si 0.1545 0.1564 0.1422 0.1531 0.1490 Bonn zoological Bulletin 57 (2): 267-274 ©ZFMK Canonical discriminant function A & eu © N Ai eian ©3 a c Ce oe 9 °K af ew 0 KGa aA aa 2 ie Nacteavay s a Ww 4 e AL IGA 3 3 : .a ° ‘ of T T T SS) es 6 3 oO Function 1 Canonical discriminant function Function 2 Function 1 Fig. 4. CDF plots for Tarentola sp-Tunisia, Tarentola deserti- North Africa, Zarentola sp-complex-Libya (western Libyan 7. Jascicularis), and Tarentola deserti-Libya; males (above) and fe- males (below). fascicularis, in East Libyan Cyrenaica, belong to a sepa- rate clade (clade D). Genetic distances among Libyan and Tunisian Tarentola clades are provided in Table 2. Two of the mitochondrial clades occur sympatrically or Group Nr: “1 T. sp-complex-Libya " 2T. deserti-Libya ° 3ST. sp-Tunisia 04 T. deserti-North Africa C1 group center parapatrically: 7: deserti (Libya) and ‘7. sp. complex’ (clade H) at Itwellia (western Libya), clades C and D along the desert road Tobruk-Ajdabiya in Cyrenaica. Bonn zoological Bulletin 57 (2): 267-274 Ulrich Joger & Ismail Bshaenia Canonical discriminant function Group Nr: 41 T. sp-complex-Libya ¥ 27. deserti-Libya * 3 T. m. mauntanica-Morocco 4 * 47. m. mauritanica-Tunisia * 5T. sp-Tunisia * 6 T. m. meuritanica-Algeria se 97 T, deserti-North Africa . O group center 4 o * a Pia o- a o a Ps e) N AGA Sola Bp: < “a CT or ° i) » So aoe Ss 4 a4 vattt 4 £ anwar 4 CI s OED 3 ir Y » 245.5, y A +o . a 4 6 T T T UT af 6 3 ° 3 6 Function 1 Canonical discriminant function o o 0 o oo o o° ia o a a v Yah y ad 4” Ang “ Nene? . ¢ mote ve 4 5 of * *4 eRe r~ shige af A Fo a ° 3 % Set Y c hes Oat ae a by a c A aN x “an As ™ 64% 7 Spe mag 3 a ON + Sr | T T T 4 3 ° 3 6 Function 1 Fig. 5. CDF plots of different populations of North African Za- rentola, males (above) and females (below) plotted separately. DISCUSSION Parapatric or sympatric occurrence of mitochondrial clades could be interpreted in different ways: either different bi- ological species or co-existence of two mitochondrial dis- tinct populations in a mixed interbreeding organismal pop- ulation. In the case of 7! deserti in NW Libya, there is ev- idence of the former explanation, as the molecular differ- ences coincide with morphological differences. ©ZFMK A new TJarentola endemic to Tunisia Table 3. Variation of the Paratypes (part). Variable Mean Minimum Maximum Standard deviation Lamellae under 1st toe 10.27 9.00 12.00 0.80 Lamellae under 4 toe 15.27 14.00 17.00 0.96 Lamellae under 5‘ toe 19.67 18.00 21.00 heat Ventral scales 34.33 28.00 38.00 Did Supraorbital scales Sole) 5.00 6.00 0.46 Dorsal tubercles WATS 12.00 14.00 0.52 Gular scales 44.21 32.00 55.00 5.82 Interorbital scales S373 12.00 15.00 0.96 Head length 17.18 13.89 21.03 2.16 Head+body length 56.08 44.12 72.33 MOD Infralabialia 793 7.00 9.00 0.70 Supralabialia Hol) 6.00 9.00 0.70 Relative hindleg length 0.49 0.45 0.51 0.02 Relative head width 0.70 0.62 0.82 0.05 Relative head length 0.31 0.29 0.32 0.01 Relative foreleg length 0.36 0.32 0.40 0.02 Relative ear-snout length 0.90 0.84 0.95 0.03 When a conservative two-species concept (7) mauritani- ca —T. deserti) is applied, the mitochondrial tree unam- biguously affiliates all sequenced Libyan OTUs with 7: deserti, and not with 7) mauritanica. The mitochondrial genetic distance between the central Tunisian Tarentola, the Libyan clades and 7: deserti are lower than between the Tunisian clade and 7. m. mauritanica. This supports the view that despite some morphological similarity, the Tunisian and Libyan-Egyptian clades are not subspecies of 7. mauritanica. Assigning these OTUs to 7: deserti would, however, cre- ate a paraphyletic T. deserti, with T. neglecta and T. min- diae — which are without doubt separate species — with- in 7: deserti. The most parsimonious taxonomic solution with regard to the cladogram is to subsume clades C, D, E, G, and H under one separate species. Morphological data indicate that several of the mitochon- drially defined populations, in particular if they occur in desert areas (Sabha in South Libya, Algerian and Tunisian but not Libyan deserti, and also a subclade of clade D) can be distinguished by larger size and higher scale counts. These size-linked characters may be locally favoured by Bonn zoological Bulletin 57 (2): 267-274 environmentally triggered selection. On the other hand, morphological differences do not preclude genetic close- ness, and genetically distant clades may share morpholog- ical similarity. In conclusion, the genetically studied Zarentola from Libya and Egypt, as well as those from south central Tunisian, should be assigned to 7! fascicularis (a former subspecies of 7. mauritanica). Elevation of fascicularis to species rank is largely consistent with data of Rato et al. (2010), who distinguished two basal divisions in the sub- genus Zarentola. One of these branches lead to T. deser- ti and T. boehmei, another to 7: angustimentalis and T. mauritanica from Europe, Morocco, Algeria and north- ern Tunisia on one side, and to 7. (m.) fascicularis and Tarentola from Lampedusa and Conigli Islands on the oth- er side. Single individuals of ‘7. mauritanica’ from Alge- ria and of ‘7: deserti’ from Morocco were loosely connect- ed to the latter clade, but we do not know which taxa were really represented by those samples. The Tunisian sam- ples used by these authors clustered with 7? m. mauritan- ica, but they were exclusively from northwestern Tunisia. True 7. mauritanica exist in coastal areas of Tunisia and western Libya. OZFMK Die, Ulrich Joger & Ismail Bshaenia Males: 25 characters 13° 12 mare? Factor 2: 19,24% Factor 1: 24,69% Females: 25 characters Factor 2: 15,64 Factor 1: 20,77% Fig. 6. PCA of Zarentola sp. (Tunisia), 7. m. fascicularis (Li- bya), and different populations of 7 mauritanica. Males above, females below. In Tunisia, 7: deserti exists in the extreme south (south- southeast of the Chott al Djerid) and a new subspecies of 7: fascicularis in south central Tunisia between the Chott al Djerid and Djebel Bou Hedma. The description of this new subspecies is presented below. Description of a new subspecies of Tarentola fascicu- laris Tarentola fascicularis n. comb. Gecko fascicularis Daudin, 1802 Tarentola mauritanica mauritanica, Loveridge 1947 (partim, non Linnaeus 1768) Tarentola mauritanica fascicularis, Joger 1984 Terra typica (after designation of a neotype by Joger [1984]): Ain Teyanah, 20 km south of Benghazi, Libya. Bonn zoological Bulletin 57 (2): 267-274 Males: 25 characters Factor 2: 16,79% Factor 1: 24,15% Females: 25 characters Factor 2: 14,57% - 0 Factor 1: 20,28% Fig. 7. PCA of Zarentola deserti (Libyan populations), 7. sp. (western population of 7. fascicularis, Libya) and T. sp. (Tuni- sia). Males above, females below. Tarentola fascicularis wolfgangi ssp. n. Holotype. State Natural History Museum Braunschweig (SNHM-BS) N 41980, male, collected 19 August 1998 by Ulrich Joger (Fig. 9). Terra typica. Bou Hedma National Park, Tunisia (34.24°N, 9.23’E). Paratypes. 33 specimens; SNHM-BS 39920-39930, 41981, Bou Hedma; HLMD 2105-2109, 2265-2271, 2363- 2366, Bou Hedma; HLMD 1238-1240, Djebel Orbata/El Guettar; ZFMK 49525, 49526, Djebel Orbata: El Guettar. Description of holotype. Measurements (mm). Head + body 61.0, tail 71.4, head length 19.8, head width 14.7, head height 10.8, interorbital width 8.4, distance snout- ear 17.7; foreleg 23.9, hindleg 30.2, distance between fore- leg and hindleg 25.1, 4* toe length 5.1, toe width 2.0, di- ameter of eye 4.5. ©ZFMK A new Zarentola endemic to Tunisia 27 (BL147 T so Tarhunah Libya {BLISS Tgp Tajura Libye U0 1BL207 T go Misratah Libya nee 1BL185 T 90 Awellle Libya Clade H 2OMV10692 T sp ElPerkat Libya {BL072 T 3p Rass El Lia Libya {BL089 Ee Genen Pa lade G “BOSE t= ai Hedina Tunisia 18051 Tsp Bou Hedmea Tunisie 18047 T sp Bou Hedma pene 181220 Tsp Ras Lanu Libya 0.88) 1BL219 Tsp Ras Lanuf Libya 0.57; /BL245 T gp Taknis Libya oe Tsp Taknis el 1.00 184229 Tastm anne rr 1.00 1BL268 T sp Sidi Massod Litya 8EV8968 Tm fasicuctris Egypt Clade C 2CMV10638 T sp GaberAan Oasis Libya 18055 Tnegbcta E/ Qued Agena 1.00 BEVEOY T mindiae pee ila oak i ‘00 7014 T maureaaee Meke “ais {8030 T maurkanica Maliorca 1.00 18021 Tmaurtanica Mallorca 1.00r 18004 Tangustimentalis Lanzarnte ade esis 8003 T anqustimentatis Fuerteventura BEV9007 Tannulens annularis Egypt 18018 T boettgen Gran Canana 18019 T boettgen Gran Canaria 18010 T.d.delalandii Tenerife Clade B Fig. 8. 50% majority-rule consensus tree obtained from Baye- sian MCMC analysis, based on 1433 bp mtDNA sequences, de- picting the relationships among haplotypes. Tarentola delalandii designated as outgroup and Bayesian posterior probability va- lues are given near branches. Pholidosis. 36 longitudinal rows of ventral scales; 12 lon- gitudinal rows of dorsal tubercles, bearing strong central keel from which barely visible keels derive laterally; 13 lamellae under 1st toe, 15 lamellae under 4" toe, 20 lamel- lae under 5'h toe; 15 interorbital scales, 6 supraorbital scales; gular scales separated from mental by 3 scales, gu- lar scale count 43; 10 supralabials, 7/8 infralabials; ros- tral divided, touching nostril; nasal scales separated by one scale proximally and one scale distally. Colour (in ethanol) whitish, without any visible pattern. Fig. 9. Holotype of Zarentola fascicularis wolfgangi ssp. n. Bonn zoological Bulletin 57 (2): 267-274 Ww Variability of paratypes. Colour (in ethanol) light or medium grey dorsally, whitish ventrally. Most specimens bear following pattern: dark line on side of head from eye to above ear. Paired dark spots, followed posteriorly by unpaired whitish spot (without clear margins) distributed on mid-dorsum as follows: one in front of shoulder, one behind shoulder, two on back, one on pelvic region, one on base of tail, followed by 9-10 unpaired half-rings around dorsal part of tail. Scale count variation is shown in Table 3. Diagnosis. A small subspecies of 7: fascicularis; maxi- mum recorded bodythead length in males 72.3 mm, in fe- males 57.5 mm (up to more than 100 mm in male 7! de- serti, 81 mm in female 7: deserti; in eastern Libyan T. fas- cicularis, 97 mm can be attained in males of the Ras Lanuf population, yet only 79 mm in 7. fascicularis ssp. from northwestern Libya). Tail length usually clearly longer than body+head (index bodythead/tail 0.77-1.00; mean 0.84, as opposed to 0.98 in 7. deserti, 0.96 in T. f. fascicularis, and 0.96 in T. m. mauritanica). Snout (ear openings to mental) significant- ly longer than in T. fascicularis and T. deserti subspecies from Eastern Libya (about 90% of head length as opposed to 60-70%). Dorsal tubercles in 11—14 (most often 12) longitudinal rows, most often simply keeled (multiply keeled in 7. f fascicularis). 19-46 gular scales (45—59 gular scales in T. d. deserti). Different from all other Tunisian or Libyan Zar- entola (except T! neglecta group) by lower number of ven- tral scales (34.3 +/- 2.7) and lower number of lamellae un- derneath 1st and 4 toes (1st 10.3 +/- 0.8, 4th 15.3 +/- 1.0). 15—22 scale rows or lamellae underneath 5‘) toes (16-21 in 7. f. fascicularis, 2\—25 in T: deserti). Different from Tunisian 7. mauritanica by lower number of interorbital scales (13.7 +/- 1.0 versus 14.9 +/- 1.2); from Libyan 7. mauritanica by lower number of sublabials (7.9 +/- 0.7 versus 8.7 +/- 0.7); from western Libyan subspecies of 7. fascicularis by lower number of interorbital scales (13.7 +/- 1.0 versus 14.9 +/- 0.7). Rostral usually separated from nostril by small scales (in 7! f fascicularis rostral usual- ly in contact with nostril). Dorsal colour in life similar to 7. deserti: rosy or yellow- ish, with yellowish iris (grey in 7! m. mauritanica). Five dark transverse bands across back, often reduced to paired spots. Distribution. Endemic to Central Tunisia; known from Gafsa (Djebel Orbata) in the West to Bou Hedma in the East, south to Degache and Tozeur at northern banks of Chott al Djérid. ©OZFMK 274 Ulrich Joger & Ismail Bshaenia Fig. 10. Terra typica of Tarentola fascicularis wolfgangi ssp. n., Djebel Bou Hedma, Tunisia. Habitat. Bou Hedma National Park is famous for its relict subtropical savanna with Acacia tortilis raddiana as the dominating tree. The climate is semi-arid, with variable amounts of rainfall (annual mean about 250 mm) in au- tumn and winter. On the pediments of Djebel Bou Hed- ma (Fig. 10) as well as on the slopes of the mountain chains to the west and south of it, Zarentola fascicularis wolfgangi ssp. n. is found in rock crevices, on walls and underneath of road bridges; the geckos are active at night. Derivatio nominis. The species is dedicated to Wolfgang Bohme on the occasion of his retirement as the most suc- cessful German curator of herpetology after Robert Mertens. The senior author feels, however, also a strong affinity to the other Wolfgang, Wolfgang Bischoff, who retired this year, too. His field companionship in North Africa will be ever remembered. Acknowledgements. We thank Sherif Baha El] Din and Adel Ibrahim for providing Egyptian specimens, Wolfgang Bischoff for field companionship, Ulrich Willand for preliminary data, Miguel Vences and Susanne Hauswaldt for providing laborato- ry facilities. Bonn zoological Bulletin 57 (2): 267-274 REFERENCES Carranza S, Arnold E N, Mateo J A, Geniez M (2002) Relati- onships and evolution of the North African geckos, Geckonia and Tarentola. Mol. Phyl. Evol. 21: 244-256 Joger U (1984a) Morphologische und biochemisch-immunolo- gische Untersuchungen zur Systematik und Evolution der Gat- tung Zarentola (Reptilia: Gekkonidae). Zoologische Jahrbu- cher (Anatomie) 112: 137-256 Joger U (1984b) Taxonomische Revision der Gattung Zarento- la (Reptilia: Gekkonidae). Bonner zoologische Beitrage 35: 129-174 Joger U (2003) Reptiles and amphibians of southern Tunisia. Kaupia 12: 71-88 Joger U, Bischoff W (1989): Erste Ergebnisse einer herpetolo- gischen Forschungsreise nach Nordwest-A frika. Tier und Mu- seum (Bonn) 1: 99-106 Joger U, Amann T, Lenk P, Willand U (1998) Molekulare Merk- male und das phylogenetische Artkonzept. Zoologische Ab- handlungen, Staatliches Museum fiir Tierkunde Dresden, 50/Suppl. ,,100 Jahre Artkonzepte in der Zoologie™: 109-123. Rato V, Carranza S, Perera A, Carretero, M A & Harris D J (2010, in press): Conflicting patterns of nucleotide diversity between mtDNA and nDNA in the Moorish gecko, Tarentola mauri- tanica. Mol. Phyl. Evol. Willand U (1997) Revision der Untergattung Zarentola (Reptil- ia: Sauria: Gekkonidae). Unpublished diploma thesis, Univer- sity of Darmstadt Received: 05.X.2010 Accepted: 01.XI.2010 ©ZFMK Bonn zoological Bulletin | Volume 57 Issue 2 pp. 275-280 Bonn, November 2010 A new species of the genus 7ropiocolotes from Central Saudi Arabia (Reptilia: Sauria: Gekkonidae) Thomas M. Wilms!;*, Mohammed Shobrak? & Philipp Wagner? ' Zoologischer Garten Frankfurt, Bernhard-Grzimek-Allee 1, D-60316 Frankfurt am Main, Germany; E-Mail: thomas.wilms@stadt-frankfurt.de ? Biology Department, Science College, Taif University, P.O. Box 888, Taif, Saudi Arabia 3 Zoologisches Forschungsmuseum A. Koenig, Adenauerallee 160, D-53113 Bonn, Germany “ corresponding author Abstract. A new species of the genus 7ropiocolotes from central Saudi Arabia is described based on two specimens from the Ath-Thumamah region. The new species is a member of the subgenus Zropiocolotes and belongs to the clade includ- ing T. steudneri and T. nattereri. Key words. Zropiocolotes sp. n., Ath-Thumama, Saudi Arabia. INTRODUCTION The genus 7ropiocolotes Peters, 1880 comprises a group of small, nocturnal and ground dwelling geckos, rarely ex- ceeding 35 mm snout-vent length. Biogeographically the distribution of these geckos follows a Saharo-sindian pat- tern, ranging from Morocco and Mauritania in the west to western India (Sindaco & Jereméenco 2008; Agarwal 2009). About 13 nominal species are being distinguished within the genus 7ropiocolotes, but in addition there are published records of specimens which would possibly de- serve specific recognition and which are not yet formal- ly described (Arnold 1980; Kordges 1998; Anderson 1999; Baha el Din 2001, 2006; Sindaco & Jereméenco 2008). Beside this the species composition of the genus is under debate since the taxa of the eastern part of the distribu- tion area are assigned to Microgecko Nikolsky, 1907 and Asiocolotes Golubev, 1984 on a generic or subgeneric lev- el by some authors (Kluge 1983; Kuge 1991; Szcerbak & Golubev 1996; Sindaco & Jereméenco 2008) while oth- ers treat all of those taxa as belonging exclusively to the genus Tropiocolotes (Anderson 1961, 1999). We prefer to follow the more inclusive interpretation of 7ropiocolotes and use the name in the broader sense encompassing al- so the taxa of Iran, Afghanistan, Pakistan and India. There have been uncertainties concerning the taxonomy of some of the African and Arabian taxa within the genus, like 7: tripolitanus algericus Loveridge, 1947, T: t. apok- lomax Papenfuss, 1969, 7: steudneri (Peters, 1869) and T. nattereri Steindachner, 1901 (Baha el Din 1994, 2001; Werner 1998; Shifman et al. 1999). The main taxonomic issues were related to the validity of certain taxa (e.g., Bonn zoological Bulletin 57 (2): 275-280 T. t. apoklomax; Baha E] Din 2001), the taxonomic rank of certain taxa (e.g., 7) t. algericus which was assigned specific rank based on a proven sympatric occurrence with Tt. tripolitanus Peters, 1880; Baha El Din 2001), species delimitation (e.g., between T. nattereri and T: steudneri, Shifman et al. 1999; the type material of both species is untraceable and therefore neotypes should be designated and a thorough rediscription of both taxa prepared) and to the existence of hitherto unknown species which were discovered recently (7? nubicus Baha El Din 1999, T. bisharicus Baha El Din 2001). As already stated by Baha El Din (2001) the difficulty in finding and studying these diminutive animals, combined with their patchy geographical representation in scientif- ic collections has led to a less than satisfactory taxonom- ic evaluation to date. The specimens described as a new species in the present paper were collected in the Ath-Thumamah region in cen- tral Saudi Arabia, approximately 90 km northeast of Riyadh. According to Arnold (1986) the distribution of Tropiocolotes in Saudi Arabia is confined to north-west- ern Saudi Arabia, but already Tilbury (1988) recorded it from the Riyadh area. Thus the first specimen from Ath- Thumamah collected by Kordges was not the first pub- lished record of the genus in central Arabia (contra Ko- rdges 1998), but nevertheless the first record of the genus from Ath-Thumamah (contra Cunningham 2010, who list- ed Tropiocolotes as not yet confirmed for this area). OZFMK 276 Thomas M. Wilms et al. MATERIAL 106 specimens of the genus 7ropiocolotes from the col- lections of the Zoologisches Forschungsmuseum A. Koenig, Bonn (ZFMK), the Senckenberg Museum Frank- furt (SMF), the California Academy of Science (CAS) and the Natural History Museum Geneva (MHNG) belonging to 7! algericus (n=14), T. depressus (n=3), T. helenae (n=10), 7. nattereri (n= 14), T. persicus (n=12), T. scortec- ci (n=4), 7. steudneri (n= 43), T. tripolitanus (n=4) and the new taxon described herein (n=2) were examined. For the species not available in the present study (7. bishari- cus, T: latifi, T: levitoni, T. nubicus) morphological infor- mation were taken from Leviton & Anderson (1972), Szezerbak & Golubev (1996), Anderson (1999), Baha El Din (1999, 2001). The following characters were collected from 59 speci- mens from Algeria, Egypt, Israel, Jordan and Saudi Ara- bia (belonging to 7. nattereri, T. steudneri and the new tax- on described herein): snout-vent-length, tail length (only intact tails), number and size of postmental scales, num- ber of interorbitals (transverse scales across the interor- bital region at mid orbits, excluding palpebral folds), num- Fig. 1. Holotype of Tropiocolotes wolfgangboehmei sp. n. from Ath-Thumamah, Saudi Arabia (Fig. 1a: dorsal view, Fig. lb: ventral view, Scale: 1mm interline distance). Bonn zoological Bulletin 57 (2): 275-280 ber of upper and lower labials, number and characteris- tics of keels on subdigital lamellae, number and identity of scales bordering the nostril, number of scales around midbody. Beside this, data on colouration and pattern was collected. Additional data on morphological characters were taken from Baha el Din (1999, 2001) and Shifman et al. (1999). Measurements were taken with a digital cal- liper to the nearest 0.1 mm. Material examined Tropiocolotes algericus: Algeria: SMF 8167, Algerian Sahara; Mali: MHNG 2678.087, north of Bombax; Mo- rocco: MHNG 1553.065-067, Tarfaya; MHNG 993.027, Aouinet-Torkoz; SMF 73082-87, Goulimine; Western Sa- hara: MHNG 1545.076, El-Aioun. Tropiocolotes depres- sus: Pakistan: SMF 64490-92, east of Chiltan-Mountains, Quetta. Tropiocolotes helenae: tran: MHNG 2627.011- 16, MHNG 2641.100, MHNG 2646.056-058, Mehkuyeh. Tropiocolotes nattereri: Egypt: MHNG 2710.017-018, Wadi Feran; SMF 8165, NW Sinai; ZFMK 70653-59, Ras Mohammed; Israel: SMF 47112, Wadi el Hedhira, Cen- tral Negev; Jordan: ZFMK 64673, Aqaba; Saudi Ara- bia: CAS 148526, Hagl [29 18 N; 34.57 E]; CAS 148616, Jabal as Sinfa [27 57 N; 35 47 E]. Tropiocolotes persi- cus: Pakistan: SMF 63536-47, Hab Chauki. Tropio- colotes_ scortecci: Yemen: MHNG 2428.065, Al Mabraz, Wadi Zabid; MHNG 2428.065, MHNG 2553.041, Mafraq-Mocca; MHNG 2581042, Sayhut. Tropiocolotes steudneri: Algeria: CAS 138660-63, 3 km. East of Tamanrasset; ZFMK 19853, 15 km S Terhenanet; ZFMK 33839, 90km S In Salah; Egypt: CAS 156660, Maadi-Wa- di Gindali Rd. [29 59 N, 31 28 E]; MHNG 2710.019-020, Oasis Kharga; SMF 22119, Kosseir; ZFMK 2359, ZFMK 64633, Luxor; ZFMK 20537, Cairo, Mokatana Hils; ZFMK 64641, ZFMK 64643, 10 km NW Cairo; ZFMK 65477, Giza Abu Rawash; ZFMK 77765-67, between Beni Suef u. Korimat; Sudan: CAS 174014, Assalaya Pump Station 3; MHNG 1186.078-079, Tabo; ZFMK 33840-59, Wadi Half; ZFMK 38429, Erkowit. Tropiocolotes tripoli- tanus: Egypt: SMF 22472, Heliopolis; SMK 22473, Cairo; Tunisia: MHNG 1335.04, Tozeur; SMF 8166, Tunisian Sahara. Tropiocolotes sp. n.: Saudi Arabia: ZFMK 43668, ZFMK 87120, Ath-Thumama. Despite the overall similarity of the taxa involved and the generally low level of character displacement, which is typical for geckoes, it became clear, that the specimens from central Saudi Arabia differ in several characters from all known taxa in the genus 7ropiocolotes and will there- fore be described as a new species. OZFMK A new species of the genus 7ropiocolotes from Central Saudi Arabia DTG Fig. 2. SPECIES DESCRIPTION Tropiocolotes wolfgangboehmei sp. n. Type material: Holotype, ZFMK 43668, Ath-Thumama, Saudi Arabia, leg. T. Kordges, 1985; Paratype, ZFMK 87120, Ath-Thumama (25° 16’ N, 46° 37’ E), Saudi Ara- bia, leg. T. Wilms, 09.05.2001, 10:30 hrs Diagnosis. A small gecko with a maximum snout-vent- length of 29.4 mm. The species possesses all diagnostic characters of the genus Tropiocolotes (in the sense of Kluge 1967) including digits slightly angularly bent, not dilated, not fringed, not webbed, nor ornamented, covered below with a single series of transverse lamellae, pupil ver- tical, dorsal scales uniform, small, homogenous, imbricate to subimbricate, preanal and femoral pores usually absent. Tropiocolotes wolfgangboehmei sp. n. has two pairs of postmental shields and therefore differs from T. latifi (no postmentals), 7: helenae (one pair of postmentals) and 7. depressus (no postmentals or only one pair of very small postmentals). From 7 persicus it differs by having only four scales in contact with the nostril instead of five. It differs from T. algericus, T. tripolitanus, T. scorteccii, T. somalicus and T. bisharicus by its smooth dorsal sca- lation. 7. wolfgangboehmei sp. n. differs from 7: nattereri by possessing clearly bi- or tricarinated subdigital scales (versus smooth subdigital scales) and from T. steudneri and 7. nubicus by having two pairs of postmental shields of which the second is roughly a quarter of the size of the first (both pairs of roughly equal size in 7: steudneri and T. nubicus). Bonn zoological Bulletin 57 (2): 275-280 Paratype of Tropiocolotes wolfgangboehmei sp. n. from Ath-Thumamah (25° 16’ N, 46° 37’ E), Saudi Arabia in life. Description of the Holotype. An adult female with in- tact tail. Body depressed. Snout-vent-length (SVL) 29.4 mm, Tail length 32.8 mm. Head narrow, 9.3 mm long (about 31.6 % of SVL). Neck distinct. Right limb 10.8 mm long. 5‘ digit of left manus lacking claw, all other digits complete. Tail 1.12 times SVL, cylindrical tapering even- ly to its tip. Fig. 3. Paratype of Tropiocolotes wolfgangboehmei sp. 0. from Ath-Thumamah, Saudi Arabia (Fig. 3a: dorsal view, Fig. 3b: ventral view, Scale: 1 mm interline distance). ©ZFMK 278 Thomas M. Wilms et al. Fig. 4. Habitat and Paratype locality of Tropiocolotes wolf- gangboehmei sp. n. at Ath-Thumamah (25° 16’ N, 46° 37’ E), Saudi Arabia. Rostral 1.5 times as wide as high, divided partly by a me- dian cleft. Nostril bordered by rostral, first upper labial and two small postnasals, which are separated by two large internasals. The internasals are followed by one pair of subequal scales. Snout and upper surface of the head cov- ered by hexagonal scales which are juxtaposed. Loreal re- gion covered with slightly swollen scales, which are some- what smaller than the remaining scales on the head. 16 in- terorbitals, 10/10 upper labials, 8/8 lower labials. Occip- italregion covered by juxtaposed scales slightly smaller than the interorbitals, which become increasingly swollen in the neck. Mental slightly wider than rostral, pentago- nal in shape extending posteriorly not to the level of the suture between first and second lower labials. One pair of Tropiocolotes nattereri (SMF 47112), Wadi el Hedhi- ra, Central Negev, Israel (Fig. 5a: dorsal view, Fig. 5b: ventral view, Scale: 1mm interline distance). Fig. 5. Bonn zoological Bulletin 57 (2): 275-280 large postmentals, in contact with mental and the first two lower labials. Second pair of postmentals only about one fourth the size of the first postmentals, separated from each other by four granular scales. The second pair of postmen- tals is in contact with the second lower labials. Body scalation homogenous, scales imbricate and smooth. 58 scales around midbody. Chest widely opened by an in- cision. Dorsal sides of forelimbs covered with imbricate scales, scales of ventral sides juxtaposed and slightly swollen, somewhat smaller than scales on dorsal side of forelimbs. Dorsal and ventral sides of hind limbs covered with imbricate scales, which are almost equal in size. Pos- terior surface of thigh with smaller granular scales. Sub- digital lamellae strongly bi- or tricarinate. Lamellar for- mula (digit 1 to 5) for left manus: 9, 12, 14, 13, 11. Dorsal and ventral scales of the tail homogenous and im- bricate. Scales at tail base not carinate, but becoming in- creasingly so distally. Postanal sacs weakly developed with two enlarged tubercular scales on either side. A pair of slightly enlarged preanal scales present. Measurements (in mm, from preserved specimen): Snout- vent-length 29.4; tail length 32.8; head length 9.3; max- imum head width 5.1; maximum head height 2.7; orbit di- ameter 2.0; distance orbit — snouth 3.2; distance orbit — ear 2.3; ear diameter 0.6. Colouration of preserved specimen: The specimen 1s pre- served in 70% ethanol and has almost completely lost his coloration and pattern. Kordges (1998) depicted this spec- imen in black and white, and its pattern resembles the paratype almost exactly (in having six dark transverse bands on the back and twelve on the tail, as well as hav- ing exactly the same pattern of the head). Description of the Paratype. Paratype similar in most re- spects to holotype, except as noted. An adult male with intact tail, which was broken during preservation near the tail base. Body depressed. Snout-vent-length 27.3 mm, tail length 31.1 mm. Head narrow, head length 8.2 mm (about 30 % of SVL). Neck distinct. Right limb 9.3 mm long. All digits intact. Tail 1.13 times SVL, cylindrical taper- ing evenly to its tip. 15 interorbitals, 10/10 upper labials, 7/8 lower labials. Sec- ond pair of postmentals only about one fourth the size of the first postmentals, separated from each other by three granular scales. The second pair of postmentals is in full contact with the second lower labials, and almost reach- es the third labial on the left. Body scalation homogenous, scales imbricate and smooth. 59 scales around midbody, 66 scales between a well ©OZFMK A new species of the genus Tropiocolotes from Central Saudi Arabia 279 Fig. 6. Tropiocolotes steudneri (ZFMK 33850), Wadi Halfa, Sudan. (Fig. 6a: dorsal view, Fig. 6b: ventral view, Scale: 1mm interline distance). marked interruption between throat and chest and cloacal slit. Dorsal sides of forelimbs covered with imbricate scales, some of which show very slight carination, scales of ventral sides juxtaposed and slightly swollen, somewhat smaller than scales on dorsal side of forelimbs. Lamellar formula (digit 1 to 5) for left manus: 9, 11, 15, 13, 10. Measurements (in mm, from preserved specimen): Snout- vent-length 27.3; tail length 31.1; head length 8.2; max- imum head width 4.9; maximum head height 2.9; orbit di- ameter 1.7; distance orbit — snouth 2.9; distance orbit — ear 2.0; ear diameter 0.7. Colour in life: Head light brown with a broad dark brown band extending from the snout to just above the ear open- ing. A narrow yellow line extending axially from the ros- tral to the snout, up to the upper delimitation of the broad dark brown band. Palpebral fold yellow. Labials and un- derside of the head white. Dorsum light brown, with six broad, dark brown transverse bands. Colouration between those transverse bands yellowish brown. Ground colour of dorsal sides of limbs light brown. Hind limbs scattered with dark brown spots. Tail coloration light brown with ten dark brown transverse bands. Ventral side without any pattern, white. Bonn zoological Bulletin 57 (2): 275-280 Derivatio nominis. This species is named after Prof. Dr. Wolfgang Bohme in honour to his contributions to her- petology during his 39 years as curator of herpetology at the Zoological Research Museum A. Koenig, Bonn and as the academic mentor of two of the authors of the pres- ent paper. Habitat. The holotype was found near a small village at the border of the Ath-Thumamah area (Kordges 1998). The paratype was found under a stone in a small canyon within the Buwayb-Escarpment which is a cretaceous coral reef consisting of sedimentary rock, mainly lime- and sandstone. DISCUSSION Tropiocolotes wolfgangboehmei sp. n. is known only from central Saudi Arabia. Because of the close proximity of Ath-Thumamah to the city of Riyadh we consider the 7ro- piocolotes recorded by Tilbury (1988) likely to be con- specific with this new taxon. Based on the external mor- phology 7. wolfgangboehmei sp. n. is a member of the group consisting of 7) nattereri, T. steudneri and T: nubi- cus and is most probably the sister taxon of 7. nattereri. Biogeographically the distribution pattern of the Zropio- colotes of north-eastern Africa and Arabia is quite puz- zling, not the least because of the involvement of at least two taxa which are not yet formally recognized (Guibé 1966, Arnold 1980, Anderson 1999). Due to the clarifi- cation on the taxonomic identity of 7) nattereri and T. steudneri (Werner 1998, Shifman et al. 1999) and the de- scription in the present paper, it is clear, that actually three nominal 7ropiocolotes species are known to occur on the Fig. 7. Distribution of Arabian Tropiocolotes: & Tropiocolo- tes nattereri from Saudi Arabia [upper &: CAS 148526, Hagl (29° 18’ N; 34° 57’ E); lower A CAS 148616, Jabal as Sinfa (27° 57’ N; 35° 47° E)], @ Tropiocolotes wolfgangboehmei sp. n., @ Tropiocolotes spec. from Bandar-e-Lengeh (Anderson 1999), Iran; Black areas: approx. Distribution of Tropiocolotes scortec- ci, grey areas: approx. distribution of Tropiocolotes nattereri. ©ZFMK 280 Thomas M. Wilms et al. Arabian Peninsula (7. nattereri, T. wolfgangboehmei sp. n. and T. scortecci). T. nattereri is known from north-west- ern Saudi Arabia and the adjacent areas in Jordan, Israel and Egypt while 7: scortecci is an endemic species of southern Arabia (Oman and Yemen). 7? wolfgangboehmei sp. n. is the only species distributed in central Arabia some 800-1000 km away from the nearest Tropiocolotes local- ities in north-western Saudi Arabia, Oman/Yemen and Iran. In the past, several authors (Tilbury 1988, Schneider 1990, Baha El] Din 2006, Cunningham 2010) assigned the cen- tral Arabian Tropiocolotes to T: steudneri or T. nattereri. Two specimens from Bandar-e Lengeh on the coast of the Arabian/Persian Gulf in Iran were also tentatively assigned to T. steudneri (Guibé 1966, Anderson 1999). Based on the current distribution of Tropiocolotes it would zoogeo- graphically be extremely unlikely that the [ranian speci- mens belong to either 7. steudneri or T nattereri. Based on the morphological data given by Guibé (1966) and An- derson (1999) and the photograph given by Anderson (1999) we are sure that these animals do not belong to 7° wolfgangboehmei sp. n. but to a new, as yet undescribed taxon. Baha El Din (2001) suggested the investigation of the relationship of those Iranian specimens to two Tropi- ocolotes from eastern Dhofar, Oman, which were tenta- tively assigned to 7. scortecci by Arnold (1980) despite notable differences from T. scortecci specimens from fur- ther west in Dhofar and from the type locality of this tax- on in Hadramaut, Yemen. The taxonomy of the genus Tro- piocolotes, especially in Arabia, is still in need of a thor- ough revision, not only to clarify the species composition within the genus but also to gain more data on the distri- bution of the respective taxa and to establish a hypothe- sis on their phylogenetic relationships. Acknowledgements. We thank H.H. Prince Bandar ibn Saud (Director General, Saudi Wildlife Commission, Riyadh) for the support and continuous interest in our herpetological studies in Saudi Arabia. The study would not have been possible without the help of many colleagues of which we would like to thank H.E. Prof. Dr. A. H. Abuzinada (former Director General, NCW- CD, Riyadh), Prof. Dr. I. Nader (Former Director of KK WRC, Thumama), and Dr. I. Galal (Riyadh). For the loan of important material we thank Prof. Dr. W. Bohme (ZFMK, Bonn), Dr. A. Schmitz (MHNG, Geneva), J. Vindum (CAS, San Francisco) and Dr. G. Kohler (SMF, Frankfurt). REFERENCES Agarwal I. (2009) First record of Tropiocolotes persicus euphor- biacola Minton, Anderson and Anderson 1970 (Sauria: Gekkonidae) from the Republic of India, with notes on its habitat and natural history. Russian Journal of Herpetology 16 (2): 83-87 Anderson SC (1961) A note on the synonymy of Microgecko Nikolsky with Zropiocolotes Peters. Wassmann Journal of Bi- ology 19 (2): 287-289 Bonn zoological Bulletin 57 (2): 275-280 Anderson SC (1999) The lizards of Iran. Society for the Study of Amphibians and Reptiles, Ithaca, 442 pp. Arnold EN (1980): The reptiles and amphibians of Dhofar, Southern Arabia. Journal of Oman Studies, Special Report 2: 273-332 Arnold EN (1986) A key and annotated check list to the lizards and amphisbaenians of Arabia. Fauna of Saudi Arabia 8: 385-435 Baha el Din S (1994) A contribution to the herpetology of Sinai. British Herpetological Society Bulletin 48: 18—27 Baha el Din S (1999) A new species of Tropiocolotes (Reptilia: Gekkonidae) from Egypt. Zoology in the Middle East 19: 17-26 Baha el Din S (2001) A synopsis of African and south Arabian geckos of the genus Tropiocolotes (Reptilia: Gekkonidae) with a description of a new species from Egypt. Zoology in the Middle East 22: 45—56 Baha el Din S (2006): A Guide to the Reptiles and Amphibians of Egypt. The American University in Cairo Press, Cairo, 359 Pp. Cunningham PL (2010) Checklist of terrestrial reptiles in three protected areas in the Kingdom of Saudi Arabia. Herpetolo- gical Review 41 (1): 25-28 Guibé J (1966) Reptiles et amphibiens récoltes par la Mission Franco-Iranienne. Bulletin du Muséum Natinal d’ Histoire Na- turelle 38 (2): 97-98 Kluge AG (1967) Higher taxonomic categories of gekkonid lizards and their evolution. Bulletin of the American Muse- um of Natural History 135 (1): 1-59 Kluge AG (1983) Cladistic relationships among gekkonid lizards. Copeia 1983 (2): 465-475 Kluge AG (1991) Checklist of gekkonid lizards. Smithsonian Herpetological Information Service (85): 1-35 Kordges T (1998) Die Reptilienfauna des Thumama Nature Park bei Riyadh, Saudi Arabien. Faunistische Abhandlungen Staatliches Museum ftir Tierkunde Dresden, Band 21/Suppl. Contribution to a “Herpetologia Arabica” Nr. 7: 67—83 Leviton A, Anderson SC (1972) Description of a new species of Tropiocolotes (Reptilia: Gekkonidae) with a revised key to the genus. Occasional Papers of the California Academy of Sciences 96:1—7 Schneider B (1990) Verbreitung, Unterartgliederung, Okologie und Schuppenmorphologie der Gekkoniden—Gattung Tropi- ocolotes aus dem Bereich der Sahara. Mitteilungen Pollichia 77: 409-419 Shifman S, Shacham B, Werner Y (1999): Tropiocolotes nattereri (Reptilia: Gekkonidae): comments on validity, variation and distribution. Zoology in the Middle East 17: 51-66 Sindaco R, Jereméenco K (2008) The Reptiles of the Western Palearctic. Monografie della Societas Herpetologica Italica 1, Latina, 579 pp. Szcerback N, Golubev M (1996) Gecko fauna of the USSR and contiguous regions. Society for the Study of Amphibians and Reptiles, St. Louis, [IX + 233 pp. Tilbury C (1988) An annotated checklist of some of the com- moner reptiles occurring around Riyadh, Kingdom of Saudi Arabia. African Journal of Herpetology 34: 25-34 Werner Y (1998) The desert herpetofauna in and near Israel: a personal review of advances (1986-1997), with new data (Am- phibia; Reptilia). Faunistische Abhandlungen Staatliches Museum fiir Tierkunde Dresden, Band 21/Suppl. Contributi- on to a “Herpetologia Arabica” Nr. 14: 149-161 Received: 15.IX.2010 Accepted: 10.X.2010 ©ZFMK Bonn zoological Bulletin Volume 57 Issue 2 pp. 281-288 Bonn, November 2010 Description of a new subspecies of Kinyongia uthmoelleri (Miller, 1938) (Squamata: Chamaeleonidae) with notes on its captive propagation Nicola Lutzmann!, Jan Stipala?, Ralph Lademann}, Patrick Krause+, Thomas Wilms’ & Andreas Schmitz® 'Seitzstrasse 19, D-69120 Heidelberg, Germany, langstrasse@web.de, phone: +4962217298625 2Centre for Ecology and Conservation, University of Exeter, Tremough Campus, Penryn, Cornwall, TR10 9EZ, England, janstipala@hotmail.com 3Muhlgasse 30, D-71034 Boblingen, Germany, ralph.lademann@web.de 4Lisztstrasse 3, D-53115 Bonn, drpkrause@web.de; 5Zoologischer Garten Frankfurt, Bernhard-Grzimek-Allee 1, D-60316 Frankfurt am Main, Germany, thomas.wilms@stadt-frankfurt.de 6Department of Herpetology and Ichthyology, Muséum d’histoire naturelle, | route de Malagnou, CH-1208 Geneva, Switzerland, andreas.schmitz@ville-ge.ch Abstract. A new population of Kinyvongia uthmoelleri was found in the South Pare Mountains in Tanzania in 2000 by J. Mariaux of the Natural History Museum of Geneva (MHNG). The morphology of this population corresponds well with that of other previously known populations of K. uthmoelleri from Mt. Hanang and the Ngorongoro crater highlands. Specimens from South Pare and Ngorongoro are morphologically very similar and show some distinctive characters which are divergent from the holotype of K. uthmoelleri and other specimens from Mt. Hanang: smaller size, smooth squama- tion on head and body, smooth head crests, clearly bi-forked parietal crest (only in males), parietal crest composed of only a single row of scales, a relatively narrower and longer head and no sexual dimorphism in the tail length. K. uth- moelleri is known from only few museum specimens but these morphological differences and geographic isolation jus- tify describing the Ngorongoro and South Pare populations as a new subspecific taxon: Kinyongia uthmoelleri artytor noy. ssp. The new subspecies has been successfully kept and bred in captivity by one author, and a short description is given of its captive maintenance. Key words. Kinyongia uthmoelleri, new subspecies, South Pare Mountains, captive propagation. INTRODUCTION Despite several recently described Kinyongia taxa from East Africa (Menegon et al. 2009; Necas 2009; Necas et al. 2009) the diversity within this genus has not yet been completely uncovered. Several publications covering ma- terial from the Eastern Arc Range have contributed to knowledge on the systematics and taxonomy of these chameleons (Mariaux et al. 2008; Tilbury et al. 2006). The type material of the taxon described in the present paper was collected in 2000 and was at that time deposited un- der the name “Bradypodion tavetanum” in the Muséum histoire naturelle (MHNG) in Geneva. Kinyongia uthmoelleri was described by Miiller (1938) as Chamaeleo uthmélleri on the basis of a single specimen from Mt. Hanang. This specimen was collected at 2300 m asl in montane forest. In Loveridge’s (1957) check list of East African reptiles and amphibians he designated wth- moelleri as a subspecies of Ch. fischeri, a two-horned Bonn zoological Bulletin 57 (2): 281-288 species. Mertens (1966) followed this classification, de- spite the fact that he treated it as a full species in an ear- lier publication after discovering the second specimen known to science in the Staatliches Museum fir Naturkunde Stuttgart (SMNS) (Mertens 1955). On the ba- sis of lung and hemipenial morphology Klaver & BOhme (1986) recognized uthmoelleri as a full species and includ- ed it in the genus Bradypodion. Bo6hme & Klaver (1990) discovered a third specimen, the first recorded female of this species, in the Royal Museum for Central Africa in Tervuren (MRAC). The above mentioned second and the third specimens were collected from the locality of Old- eani in the Ngorongoro crater highlands, a massif sever- al hundred kilometres north of the type locality on Mt. Hanang. Price (1996) also mentions statements from lo- cal people about locations between Babati and Singida (a road that passes close to Mt. Hanang) and 72 km north- east of Mt. Hanang but up til now the presence of K. uth- ©ZFMK 282 Nicola Lutzmann et al. Head view of the holotype of K. u. uthmoelleri (pho- Fig. 1. to: G. Vogel). moelleri at these locations have not been confirmed. Re- cently the taxon uthmoelleri was placed with all other east African Bradypodion in a new genus, Kinyongia (Tilbury et al. 2006). In the last 15 years only two authors have pub- lished details on the captive husbandry and breeding of K. uthmoelleri, specimens collected from the Ngorongoro crater highlands (Price 1996; Necas & Nagy 2009). Around the year 2000, specimens of “Bradypodion uth- moelleri” appeared in the international pet trade. These animals were very small in overall size, more slender and with smoother scalation than K. uthmoelleri specimens from Mt. Hanang. Even after six years of keeping some of these specimens in captivity these distinct characters have not changed and so ontogenetic change in these char- acters can be ruled out. Unfortunately, the geographic ori- gin of these specimens was not known until four similar specimens were discovered in the collection of the Muséum d’histoire naturelle in Geneva in 2004, which suggests they originate from the same locality, the South Pare Mountains, and belong to the new subspecies de- scribed in this paper. MATERIAL AND METHODS In total 20 specimens of K. uthmoelleri of both subspecies with a confirmed collection locality were located in mu- seum collections and investigated: 8 from Mt. Hanang (5 males [ZSM 1/1948 (Holotype), ZFMK 74955, ZFMK 82188 and ZFMK 82189], 3 females [ZFMK74953, ZFMK 74954 and ZFMK 82191] and one subadult [ZFMK 82190]), 8 from the Ngorongoro crater highland area [1 male (SMNS 324), 2 females (ZFMK 58664 and ZFMK 58665), 1 subadult (MRAC R.G. 21852), 4 em- bryos (ZFMK 58666-69] and 4 from the South Pare Mountains [2 males (MHNG 2612.65 and MHNG 2612.66), 1 female (MHNG 2612.67), 1 juvenile (MHNG 2612.64)]. It seems probable that the embryos in the Bonn zoological Bulletin 57 (2): 281-288 ZFMK collection are the unhatched specimens reported by Price (1996). Head-body length (HBL), tail length (TL), total length (ToL), head length (HL) and head width (HW) were meas- ured in all specimens except in the embryos. The data of MRAC R.G. 21852 were taken from Bohme & Klaver (1990). The ratio of HL to HW and the percentages of HL to HBL, TL to ToL and TL to HBL were calculated. In addition, we recorded head crest morphology following Necas (1994), and the morphology and pattern of body scalation. RESULTS All measurements and investigated morphological char- acters of the specimens are listed in Tables 1-3. The mor- phological traits which differentiate the male specimens of Mt. Hanang from those of the South Pare Mountains and Ngorongoro crater highlands are: higher measure- ments, a relatively broader and shorter head, rougher (more convex) scalation on the head and body, canthus parietalis (cp) not bi-forked anteriorly but fan-shaped an- teriorly and the cp composed of two rows of scales (Fig. Fig. 2. gel). Type material of K. u. artytor ssp. n. (photo: G. Vo- ©ZFMK New subspecies of Kinyongia uthmoelleri Table 1. Morphological measurements of K. wthmoelleri in mm. TL remark specimen locality sex HBL ToL HW HL ZFMK 74955 Mt. Hanang m 90.1 Py PARE [Sal 26.6 ZFMK 82188 Mt. Hanang m 85.2 119.6 204.6 15.8 26.0 ZFMK 82189 Mt. Hanang m 92.8 WAST PASS) 16.4 30.0 ZFMK 82190 Mt. Hanang m 69.9 93.1 163.0 12.0 22.7 subadult ZSM 1/1948 Mt. Hanang m 93.0 134.0 227.0 16.0 32.0 holotype of K. u. uthmoelleri SMNS 324 Ngorongoro area m 83.0 116.0 199.0 13.0 31.0 MHNG 2612.64 South Pare Mountains m 40.0 46.0 86.0 6.5 13.5 juvenile; paratype of K. u. artytor ssp. n. MHNG 2612.65 South Pare Mountains m 80.0 100.0 = 180.0 13.0 31.0 holotype of K. u. artytor ssp. n. MHNG 2612.66 South Pare Mountains m 67.0 86.0 153.0 10.0 24.0 paratype of K. u. artytor ssp. n. ZFMK 74953 Mt. Hanang f 86.1 95.3 181.4 13.2 20.2 ZFMK 74954 Mt. Hanang f 82.0 91.5 173.5 13%5 24.1 ZFMK 82191 Mt. Hanang if 78.5 82.9 161.4 12.9 ZS ZFMK 58664 Ngorongoro area f 78.6 95.1 Sid) 11.4 21.1 ZFMK 58665 Ngorongoro area ie 76.1 92.2 168.3 1222 21.0 MRAC R.G.21852 Ngorongoro area f 54.0 61.0 115.0 8.0 19.0 — subadult MHNG 2612.067 South Pare Mountains f 70.0 81.0 151.0 10.0 21.0 paratype of K. u. artytor ssp. n. 1). The females show the same differences between both populations except that the females from the Mt. Hanang population show also a fan-shaped cp anteriorly, instead of no furcation at all in the females from the South Pare Mountains and Ngorongoro highlands. Additionally, the Mt. Hanang specimens are sexually dimorphic in tail length relative to body length (males having relatively longer tails than females), whereas relative tail length be- tween the sexes of specimens from the South Pare Moun- tains and Ngorongoro highlands specimens is more or less the same. Based on these key characters that differentiate the two groups, we describe the populations from the South Pare Mountains and the Ngorongoro crater high- lands as a new subspecific taxon. Fig. 3. Holotype of K. u. artytor ssp. n. (photo: N. Lutzmann). Bonn zoological Bulletin 57 (2): 281-288 Kinyongia uthmoelleri artytor ssp. n. We chose the syntopic specimens collected by J. Mariaux & C. Vaucher in the South Pare Mountains during their journey in 2000 as the type specimens (Fig. 2). Diagnosis. A small chameleon, which differs from the nominate form on Mt. Hanang in the following charac- ters: less convex scalation on body and head, smooth head crests, parietal crest distinctly bi-forked anteriorly, the ridge of the parietal crest contains only one scale row, a higher ratio of HL to HW and HL to HBL (relatively longer and narrower heads), smaller total length [153.0-199.0 mm in males (204.6—227.0 mm in K. u. uth- moelleri) and 151.0—-173.7 mm in females (161.4—181.4 mm in K. u. uthmoelleri)| and no sexual dimorphism in the relative tail length. Description of the Holotype (Figs 3-5). MNHG 2612.65, adult male, 1840 m asl, South Pare Mountains, North Tan- zania, leg. J. Mariaux & C. Vaucher, 29. 09. 2000. HBL 80.0 mm, TL 100.0 mm, ToL 180.0 mm, HL 31.0 mm, HW 13.0 mm, the belly is cut and the intestine removed, both hemipenes are partly everted, length of lower jaw 21.0 mm, distance from front edge of eye to nostril 9.8 mm, distance from nostril to snout tip 5.4 mm, distance from lower jaw to the tip of casque 7.5 mm, head width between eyes 6.5 mm, canthus temporalis from eye to an- gle 7.7 mm, canthus parietalis (cp) is bi-forked anterior- ly (Fig. 5), distance from bifurcation of cp to the top of ©ZFMK 284 Nicola Lutzmann et al. Table 2. Ratios of morphological measurements of K. uthmoelleri. specimen location sex HL/HW HLas% HBL TLas % ToL TL as % HBL ZFMK 74955 Mt. Hanang m 1.76 29.52 57.68 136.29 ZFMK 82188 Mt. Hanang m 1.65 30.52 58.46 140.38 ZFMK 82189 Mt. Hanang m 1.83 32.33 S53 135.45 ZFMK 82190 Mt. Hanang m 1.90 32.47 57.12 133.19 ZSM 1/1948 Mt. Hanang m 2.00 34.41 59.03 144.09 SMNS 324 Ngorongoro area m 2.38 37.35 58.29 139.76 MHNG 2612.064 South Pare Mountains m 2.08 33.75 53.49 115.00 MHNG 2612.065 South Pare Mountains m 2.38 38.75 55.56 125.00 MHNG 2612.066 South Pare Mountains m 2.40 35.82 56.21 128.36 ZFMK 74953 Mt. Hanang f 1.53 23.46 52.54 110.69 ZFMK 74954 Mt. Hanang f 1.79 29:39 52.74 111.59 ZFMK 82191 Mt. Hanang f 1.65 27.13 51.36 105.61 ZFMK 58664 Ngorongoro area f 1.85 26.84 54.75 120.99 ZFMK 58665 Ngorongoro area fe a2 27.60 54.78 121.16 MRAC R.G.21852 | Ngorongoro area f 2.38 35.19 53.04 112.96 MHNG 2612.067 South Pare Mountains f P| 30.00 53.64 Seyi casque 13.1 mm, length of bifurcation of cp 4.4 mm, max- imum width of bifurcation of cp 4.3 mm, one conical scale in the neck smaller than 2.0 mm, no ventral or tail crests, collection and field number (TZ-141) are tied around the left hind leg. The scales on the head, the head crests and the body are flat. Only the ridge of the cp is pronounced though not denticulate. Fig. 6 shows the colouration of the holotype in life. Variation on the paratypes (MNHG 2612.64, 2612.66- 67). All measurements of the paratypes and the other spec- imens of K. wu. artytor ssp. n. are shown in Tables 1-2. MNHG 2612.64 is a juvenile male, the belly is cut and the intestines are removed, the colouration after preser- vation is very dark, collection and field number (TZ-138) is tied around the right hind leg. MNHG 2612.66 is an adult male and fits quite well with the description of the holotype: belly is cut but the intestines are still present, 2 conical scales in the neck, collection and field number Fig. 4. Portrait of the holotype of K. u. (photo: N. Lutzmann). artytor ssp. Nn. Bonn zoological Bulletin 57 (2): 281-288 (TZ-143) are tied around the right hind leg. The original colouration is better preserved: head is greyish, red radi- ations on the eyes, which continue darker on the head sides, the interstitial skin is red around throat and neck, the lateral stripe is greyish on dark background, the tail is greyish. MNHG 2612.67 (Fig. 7) is an adult female with a flat casque, cut belly without intestines and one conical scale in the neck. Collection and field number (TZ-144) are tied around the left hind leg, the colouration after preservation is very dark with only some greyish flat scales on the head and body. Distribution. K. wu. artytor ssp. n. is know only from the South Pare Mountains and the Ngorongoro crater high- lands (Fig. 8). Etymology. The subspecies name “‘artytor” 1s the latinised substantive of the Greek verb “aptuetv" (artyein), which can be translated as “to prepare / to make ready requiring skills”. We name this new subspecies in honour and trib- ute to Prof. Dr. Wolfgang Bohme and his skills to prepare dozens of students on their way to scientific careers, which was also the case for four of the authors of this publica- tion. Captive maintenance. All specimens were kept individ- ually in full gauze terrariums indoor and outdoor in the same cages in order to minimize the stress of relocation. The size of the terrartums were for females 50x50x80 cm and for males 45x50x70 cm (length x width x height). All specimens were kept outdoor from spring to autumn, if the temperatures did not fall consistently below 10 °C at night time. The highest recorded temperature was 35 °C at noon, the lowest 5 °C at night time. The cages were ex- posed to the sun in the morning and fell into shade around ©ZFMK New subspecies of Kinyongia uthmoelleri Table 3. Morphological characters of K. uthmoelleri. No oO Nn head scalation No. scale rows specimen location SeX body scalation —_bi-forked cp on the ridge of cp ZFMK 74955 Mt. Hanang m rough rough no 2 ZFMK 82188 Mt. Hanang m rough rough no 2 ZFMK 82189 Mt. Hanang m rough rough no 2 ZFMK 82190 Mt. Hanang m rough rough no 1-2 ZSM 1/1948 Mt. Hanang m rough rough no 2 SMNS 324 Ngorongoro area m flat flat yes l MHNG 2612.065 South Pare Mountains m flat flat yes ] MHNG 2612.066 South Pare Mountains m flat flat yes ZFMK 74953 Mt. Hanang tg rough rough no 7 ZFMK 74954 Mt. Hanang f rough rough no 2 ZFMK 82191 Mt. Hanang f rough rough no 2 ZFMK 58664 Ngorongoro area f flat flat = | ZFMK 58665 Ngorongoro area f flat flat = 1 MRAC R.G.21852 Ngorongoro area f flat flat ? ? MHNG 2612.067 South Pare Mountains =f flat flat - l noon. In spring and autumn the cages were sprinkled with water four times per day (in midsummer 6 times) for up to four minutes in the hottest time of the day. During the winter the terrariums were illuminated with common ter- rarium-tubes (T5 with 35 W) 13 hours per day. A halo- gen spot was activated for 45 minutes three times per day for basking, so that the ambient temperature stayed be- tween 22 and 24 °C at day time and between 6 and 16 °C at night time. The terrariums were completely sprinkled with water in the morning and evening. The diet consist- ed of small arthropods, mainly self-bred crickets, grasshoppers, flies, cockroaches etc. Every second feed- ing the food was enriched with vitamins and minerals. On- ly pregnant females were additionally given small pieces of cuttlebone. To trigger mating behaviour, the males were transferred into the cages of the females. Immediately, the males started head bobbing and displayed bright colours. Fig.5. Head view of the holotype of K. wu. artytor ssp. n. (photo: N. Lutzmann). Bonn zoological Builetin 57 (2): 281-288 In all cases the females displayed a colouration of green- ish-white with small black dots, whereon the males stopped courtship. Matings have not be observed until now, but after several days the females started gaining weight and became visibly rounder. The males were sub- sequently removed, because it seemed that the females on- ly lay their eggs if there were no males in their vicinity. Older females laid their clutches without test excavations, younger females with test excavations at a depth of 5 to 7 cm into the terrartum substrate. The clutches consisted of 7 to 12 eggs. The dimensions of the eggs were approx- imately 8.0x4.0 mm. The eggs were incubated in com- pletely closed, small plastic boxes in wet vermiculite. Af- ter approximately 115 days at 19-21 °C during the day and 15-18 °C at night, the temperatures were increased to 22 °C during the day and 20 °C at night. At this time the humidity of the vermiculite was also increased to sim- ulate the beginning of a rainy season. Hatching started af- ter 147 to 161 days. After the hatchlings opened the egg shells, they occasionally paused for up to 3 days to resorb the yolk. The young chameleons were kept individually in smaller cages 25x25x40 cm under the same conditions as the adults. It should be taken into account that the tem- perature changes should not be as pronounced for the ju- veniles as for the adults, because it seems that they are un- able to thermoregulate effectively. The maximum record- ed lifespan in captivity for this species is six years (Fig. 9). DISCUSSION The genus Kinyongia contains currently 17 species, all of which are restricted to moist montane forests in the East and Central African highlands. Recently several new ©ZFMK 286 Nicola Lutzmann et al. Fig. 6. species have been described from montane forests in Kenya and Tanzania (Menegon et al. 2009, Necas 2009, Necas et al. 2009) and several subspecies have also been raised to species status based on genetic divergence and detailed morphological studies (Mariaux et al. 2008). No doubt more species remain to be discovered in the still poorly surveyed mountain ranges across East Africa. The discovery of K. uthmoelleri in the South Pare Mountains also shows that species’ distribution ranges are not well documented and it is quite likely that K. uwthmoelleri also occurs on other massifs in-between these now known pop- ulations, such as Mt. Kilimanjaro and Mt. Meru (Fig. 8). K. uthmoelleri has a similar distribution to the Trioceros sternfeldi species complex, including the recently de- scribed 7. hanangensis (Krause & B6hme 2010). Although the phylogeography of all 7: sternfeldi populations has not been investigated, the Mt. Hanang population has been identified as a divergent sister clade to the Mt. Meru/ Kil- imanjaro populations. A similar pattern is found in K. uth- moelleri, the Mt. Hanang populations morphologically di- Fig. 7. (photo: J. Mariaux). Female paratype of K. u. artytor ssp. n. in life Bonn zoological Bulletin 57 (2): 281-288 Holotype of K. u. artytor ssp. n. in life (photo: J. Mariaux). vergent from the Ngorongoro/ Pare populations, suggest- ing that despite its geographically intermediate position, Mt. Hanang populations have been isolated for a longer period of time. Volcanic activity in the North of Tanza- nia, which created these massifs, persisted from Oligocene (37 myr ago) to the Quaternary. Subsequent colonisation and population fragmentation of chameleon populations on these massifs has resulted in their diver- sification into a number of morphologically similar but clear divergent (sub-) species. The rarity of some species of the genus Kinyongia in mu- seum collections is explainable because they inhabit the rainforest canopy and their cryptic morphology and be- haviour (Necas & Nagy 2009). Unfortunately Kinyongia uthmoelleri is one of the rarest chameleons of East Africa in museum collections, although Price (1996) mentioned, that this species is common in the Ngorongoro crater high- lands. But during eight days of fieldwork he also found only five specimens. In total, there are only 20 specimens in museum collections in Europe from three different lo- calities and more intensive fieldwork is required to bring to light if this reflects the real situation of population den- sities, distribution and ecology of this species in the wild. Further collections will also confirm if the morphologi- cal variation recorded here, from the relatively few spec- imens available, is consistent within and between the three populations. Nevertheless, there are pronounced morphological differ- ences between the Ngorongoro and South Pare specimens and the specimens from the type locality on Mt. Hanang (Miller 1938). These are sufficiently distinct to justify their description as a new taxon. This is similar to the sit- uation where K. boehmei was originally described as a subspecies of K. tavetana (Lutzmann & Necas 2002) and later elevated in to full species rank based on genetic di- vergence from all other two-horned chameleons (Mariaux OZFMK New subspecies of Kinvongia uthmoelleri 287 Cg Ngorongoro crater ad highlands #® a Mt. Hanang 160 Kilometers Fig. 8. and Ngorongoro crater highlands)]. et al. 2008). Molecular studies have revealed numerous cryptic species among East African chameleons (Matthee et al. 2004; Tilbury & Mariaux 2006; Mariaux et al. 2008, Krause & Bohme 2010) and follow-up studies using mo- lecular data should provide a better insight into the evo- lutionary relationships and genetic divergence that exists between the three isolated populations of K. uthmoelleri, some of which may justify species status. Acknowledgements. We specially want to thank Jean Mariaux, Geneva, for providing us the with type material, information on the type locality and pictures of this new taxon. Thanks also to the colleagues providing acces to collection and material: Wolf- gang Bohme and Ulla Bott (ZFMK), Frank Glaw (ZSM), An- dreas Schliiter and Axel Kwet (SMNS). REFERENCES Bohme W, Klaver C (1990) Zur Kenntnis von Bradypodion uth- moelleri (Miller, 1938). Salamandra 26 (4): 260-266 Klaver C, Bohme W (1986) Phylogeny and classification of the Chamaeleonidae (Sauria) with special reference to hemipenis morphology. Bonn. Zool. Monogr. 22: 5—60 Bonn zoological Bulletin 57 (2): 281-288 TANZANIA Mt. Kilimanjaro are Mountains Elevation (metres ak | 0- 1000 [_] 1000 - 1500 [=>] 1500 - 2000 (5)} 2000 - 2500 (i) 2500 - 3000 ls) 3000 - 6000 SE Collecting localities of K. uthmoelleri |K. u. uthmoelleri (Mt. Hanang); K. u. artytor ssp. nov. (South Pare Mountains Krause P, Bohme W (2010) A new chameleon of the 7rioceros bitaeniatus complex from Mt. Hanang, Tanzania, East Africa (Squamata: Chamaelonidae). Bonn. Zool. B. 57(1): 19-29 Loveridge A (1957) Check list of the reptiles and amphibians of East Africa (Uganda, Kenya, Tanganyika, Zanzibar). Bull. Mus. Comp. Zool. 117: 153-362 Lutzmann N, Necas P (2002) Zum Status von Bradypodion tave- tanum (Steindachner,1891) aus den Taita Hills, Kenia, mit Beschreibung einer neuen Unterart (Reptilia: Sauria: Chamaeleonidae). Salamandra 38 (1): 5-14 Matthee C, Tilbury CR, Townsend T (2004) A phylogenetic re- view of the African leaf chameleons: genus Rhampholeon (Chamaeleonidae): the role of vicariance and climate change in speciation. Proc. Roy. Soc. 271: 1967-1975 Mariaux J, Tilbury CR (2006) The pygmy chameleons of the Eastern Arc Range (Tanzania): Evolutionary relationships and the description of three new species of Rhampholeon (Sauria: Chamaeleonidae). Herpetological Journal 16: 315—331 Mariaux J, Lutzmann N, Stipala J (2008) The two-horned Chameleons of East Africa. Zool. J. Linn. Soc. 152: 367-391 Menegon M, Tolley K, Jones T, Rovero F, Marshall AR, Tilbury CR (2009) A new species of chameleon (Sauria: Chamaeleonidae: Kinyongia) from the Magombera forest and the Udzungwa Mountains National Park, Tanzania. African Journal of Herpetology 58 (2): 59-70 ©OZFMK 288 Nicola Lutzmann et al. 7 Fig. 9. Mertens R (1955) Amphibien und Reptilien aus Ostafrika. Jh. Ver. Vaterl. Naturk. Wiirtt. 110: 47-61 Mertens R (1966) Liste des rezenten Amphibien und Reptilien: Chamaeleonidae. Das Tierreich, Berlin Miiller L (1938) Uber die von den Herren L. Uthmiller und L. Bohmann im britischen Mandatsgebiet “Tanganyika Territo- ry” gesammelten Chamialeons. Zoologischer Anzeiger 122: 20-23 Necas P (1994) Bemerkungen zur Chamaleon-Sammlung des Naturhistorischen Museums in Wien, mit vorlaufiger Beschrei- bung eines neuen Chamaleons aus Kenia (Squamata: Chamaeleonidae). Herpetozoa 7 (3/4): 95-108 Necas P (2009) Ein neues Chamaleon der Gattung Kinyongia (Tilbury, Tolley & Branch 2006) aus den Poroto-Bergen, Stid- Tansania (Reptilia: Sauria: Chamaeleonidae). Sauria 31 (2): 41-48 Bonn zoological Bulletin 57 (2): 281-288 A six year old K. u. artytor ssp. n. in captivity (photo: R. Lademann). Necas P, Sindaco R, Koreny L, Kopecna J, Malonza PK, Mod- ry D (2009) Kinyongia asheorum sp. n., a new montane chameleon from the Nyiro Range, northern Kenya (Squama- ta: Chamaeleonidae). Zootaxa 2028: 41—50 Necas P, Nagy P (2009) Ein Beitrag zur Kenntnis von Uthmdllers Chamialeon, Kinyongia uthmoelleri (Miller, 1938). Sauria 31 (1): 15-21 Price T (1996) Zur weiteren Kenntnis von Bradypodion uthmoel- leri (Miiller, 1938). Salamandra 32 (3): 199-202 Tilbury CR, Tolley KA, Branch WR (2006) A review of the sys- tematics of the genus Bradypodion (Sauria: Chamaeleonidae), with the description of two new genera. Zootaxa 1363: 23-38 Received: 30.VII.2010 Accepted: 15.X.2010 ©ZFMK Bonn zoological Bulletin | Volume 57 Issue 2 pp. 289-296 | Bonn, November 2010 A new species of the genus Lycodon (Boie, 1826) from Yunnan Province, China (Serpentes: Colubridae) Gernot Vogel! & Patrick David? ‘Society for Southeast Asian Herpetology, Im Sand 3, D-69115 Heidelberg, Germany; E-mail: Gernot. Vogel@t-online.de _ 2Muséum national d’ Histoire naturelle, Département Systématique et Evolution, Reptiles & Amphibiens, UMR 7205 OSEB, Case postale 30, 57 rue Cuvier, F-75231 Paris Cedex 05, France; E-mail: pdavid@mnhn. fr Abstract. A new species of the genus Lycodon is described from Yunnan Province, People’s Republic of China. It dif- fers from the superficially similar Lycodon fasciatus by the fact that the loreal is not entering orbit, in colouration, and lower numbers of subcaudals and infralabials. From the Lycodon ruhstrati group it differs by the colouration of the ven- ter and the dorsal bands. This new species is only known the Chinese province of Yunnan. Keywords. Oriental Region, China, Colubrinae, Lycodon fasciatus, taxonomy, Lycodon synaptor sp. n. INTRODUCTION Snakes of the species rich genus Lycodon Bole, 1826 re- ceived considerable attention in most regions of Asia. Six new species were described form the Philippines (Ota & Ross 1994; Lanza 1999; Gaulke 2002) and new species were discovered in Cambodia (Daltry & Wiister 2002), In- dia (Mukherjee & Bhupathy 2007) and Myanmar (Slowin- ski et al., 2001). The taxonomy of the Chinese part of the genus remained unattended until recently. Pope (1935) list- ed five species, namely Lycodon capucinus Bote, 1827, Lycodon fasciatus (Anderson, 1879), Lycodon laoensis Ginther, 1864, Lycodon ruhstrati (Fischer, 1886) and Ly- codon subcinctus Bote, 1827. This arrangement has not changed for the next 75 years. Vogel et al. (2010) reviewed the Lycodon ruhstrati complex and described Lycodon ruhstrati abditus as a new subspecies from China, and revalidated Lycodon futsingensis (Pope, 1928). Detailed examination of banded specimens of the genus led us to the conclusion, that the diversity is much higher in this region and that several species await description. In the course of our ongoing review of the Lycodon fas- ciatus complex, we came upon two specimens of the genus Lycodon from Yunnan, China, which seemed to be differ- ent from L. fasciatus. A detailed examination showed clear morphological differences which lead us to describe them as new species. Bonn zoological Bulletin 57 (2): 289-296 MATERIAL & METHODS This revision is based on a total of 67 preserved specimens of Lycodon fasciatus auctorum examined for their exter- nal morphological characters and on several photographed specimens. They are listed in the Appendix I. Compara- tive material of the L. ruhstrati complex is listed under Vogel et al. (2010). A total of 53 morphological characters were recorded for each specimen. The characters and their abbreviations are listed in Table 1. Not all of these characters have been used for this study, but all of them were compared. Measurements, except body and tail lengths, were taken with a slide-caliper to the nearest 0.1 mm; all body meas- urements were made to the nearest millimetre. The num- ber of ventral scales was counted according to Dowling (1951). Half ventrals were not counted except they were present on both sides (divided ventrals). The terminal scute is not included in the number of subcaudals. The dorsal scale row counts are given at one head length behind head, at midbody (1.e., at the level of the ventral plate correspon- ding to a half of the total number of ventrals), and at one head length before vent. We considered being sublabials those shields that were completely below a supralabial. Values for paired head characters are given in left / right order. ©ZFMK 290 Gernot Vogel & Patrick David Table 1. List of morphological characters used in this study and their abbreviation. Abbreviation Characters N° Morphometry l SVL 2 TaL 3 TL 4 Rel TL Anatomy 5 TEETH Scalation 6 DSR i ASR 8 MSR 9 PSR 10 Keel 11 VEN 12 PreVEN 13. VEN not 14. VEN keel [S56 16 ANA 17. Lor-l 18 ~ Lor-r 19. Lo touch-l 20 Lo touch-r 21 SL-l 22 SL-r 23 SL/Eye-l 24 SL/Eye-r 25. Larg SL-l 26 _Larg SLrl 29 Vel 28 IL-r 29 ~IL-tot 30“ IL/Ist child 31 = PreOc-l 32 PreOc-r 33 PostOc-l 34 ~=PostOc-r 35. ATem-1 36 =©6©ATem-r 37 =PTem-| 38 PTem-r 39 ~=ParaR 40 ‘Paras 41 Parab Pattern 42 BODCOL 43 Bands 44 Tail bands 45 Tail venter 46 Bellycol 47 Bellyspeck 48 First band 49 Broad base 50. _ Broad vert 51 Edged 52 Coul throat Ve throat Snout-vent length (mm) Tail length (mm) Total length (mm) Relative tail length TaL/TL Number of upper maxill. teeth (one side) Dorsal scale rows Dorsal scale rows at neck Dorsal scale rows at midbody Dorsal scale rows before vent Number of keeled dorsal rows Ventral plates Number of preventrals Ventrals notched or not Ventrals keeled Subcaudal plates Anal plate: 1: single — 2: divided Number of loreal scale (0 or 1) at left Number of loreal scale (0 or 1) at right Loreal scale touches eye at left Loreal scale touches eye at right Number of supralabials at left Number of supralabials at right Numbers of the SL entering orbit at left Largest SL left Largest SL right Number of infralabials at left Number of infralabials at right Total number of infralabials Number of IL in contact with anterior chin shield Number of preoculars at left Number of preoculars at right Number of postoculars at left Number of postoculars at right Number of anterior temporals at left Number of anterior temporals at right Number of posterior temporals at left Number of posterior temporals at right Temporal row containing paraparietals Plates surrounding paraparietals, see Inger & Marx (1965) Scales between the paraparietals Body colour!: grey; 2: brown or ochre Number of bands on body Number of bands on tail Colouration of tail venter Colouration of belly Speckling of belly Number of VEN before the first band starts, counted left side Number of VEN that are covered at the base of the first band Numbers of vertebral scales that are covered by the first band Dorsal bands with light margins Colour of the throat Dark VEN on the throat before the first band Bonn zoological Bulletin 57 (2): 289-296 Numbers of the SL entering orbit at right The white or light bands on the body and tail were count- ed on one side. Hardly visible or incomplete bands were counted as one, bands that were fused were counted as two. The collar on the neck was not counted and bands covering the anal shield were added to the bands of the body. Museum abbreviations BMNH: The Natural History Museum, London, UK. — BNHS: Bombay Natural History Society, Mumbai, India. — CAS: California Academy of Sciences, San Francisco, USA. — CIB: Chengdu Institute of Biology, Chengdu, People’s Republic of China. — FMNH: Field Museum of Natural History, Chicago, USA. — KIZ: Kunming Insti- tute of Zoology, Kunming, People’s Republic of China. — MNHN: Muséum national d’Histoire naturelle, Paris, France. — NMW: Naturhistorisches Museum Wien, Aus- tria. — ZFMK: Zoologisches Forschungsmuseum Alexander Koenig, Bonn, Germany. — ZMB: Zoologis- ches Museum fiir Naturkunde der Humboldt-Universitat zu Berlin, Berlin, Germany. — ZSM: Zoologische Staatssammlung, Miinchen, Germany. RESULTS Lycodon synaptor sp. n. Holotype. BMNH 1905.1.30.63 adult female (tail dissect- ed), from “Tongchuan, Yunnan”, today Dongchuan, 100 km north of Kunming, Yunnan Province, People’s Repub- lic of China (Figs 1—3). Collected by the J. Graham Ex- pedition, unknown date. Be 12a Ee D Fig. 1. Dorsal view of preserved holotype of Lycodon synap- tor sp.n., BMNH 1905.1.30.63 from Dongchuan, 100 km north of Kunming, Yunnan Province, People’s Republic of China. Pho- tograph by Gernot Vogel. ©ZFMK New Lycodon from Yunnan 29] Fig. 2. Ventral view of preserved holotype of Lycodon syn- aptor sp. n., BMNH 1905.1.30.63 from Dongchuan, 100 km north of Kunming, Yunnan Province, People’s Republic of Chi- na. Photograph by Gernot Vogel. Paratype. MNHN 1905.0283, adult female (tail dissect- ed), from “Tongchuan Fu, Chine”, at present Dongchuan, Yunnan Province, People’s Republic of China. Collected by W. F. H. Rosenberg on 21s‘ July 1905. Diagnosis. A species of the genus Lycodon characterized by: (1) loreal scale not entering orbit; (2) 15—17 dorsal scale rows at the forepart of the body and 17 dorsal scale rows at midbody; (3) upper and vertebral dorsal rows (6-7) keeled; (4) 201—203 ventrals in females, males un- known; (5) 68-69 Sc in females, males unknown; (6) a relative tail length of about 0.189—0.192 in females, males unknown; (7) 8 supralabials with SL 4—6 touching the or- bit (7) 30-31 narrow white bands on a dark body; (8) width of the first band vertebral 0.5—1.0 scales, on the base 3 ventrals; and (9) the belly with discreet bands through- out. Fig. 3. Lateral view of the right side of the head of preserved holotype of Lycodon synaptor sp. n., BMNH 1905.1.30.63 from Dongchuan, 100 km north of Kunming, Yunnan Province, People’s Republic of China. Photograph by Gernot Vogel. Bonn zoological Bulletin 57 (2): 289-296 The new species can be recognized by the combination of the loreal scale not entering orbit (entering in L. fas- ciatus sensu stricto), its narrow dorsal bands, with the first band starting at ventral S—9 (more irregular in L. fascia- tus [Fig. 4] and species of the L. ruhstrati group, where they usually start later) and the dark throat, which usual- ly is light in other species of the L. fasciatus group and the L. ruhstrati group. Most other characters match with Lycodon fasciatus. Detailed comparisons with other species of the genus Ly- codon appear below in the Discussion. Fig. 4. Dorsal view of Lycodon fasciatus. CIB 9804, from Ruili City, Yunnan. Note the irregular bands. Photograph by Ger- not Vogel. Etymology. This species is indirectly named in honour of Dr. Wolfgang Bohme. It was always a publicized aim of Wolfgang Bohme to unite professional and amateur her- petologists. We, both authors have always been amateur herpetologists, so it is a delight for us to dignify his ef- forts towards this aim. The specific name synaptor, a noun in apposition, stems from the Greek word “ovuvonpts” meaning “a connection”. In this case, this noun underlines the connection of these two kinds of herpetologists. We suggest the following common names: Boehme 5 wolf Snake (English), Bohmes Wolfszahnnatter (German). Description of the holotype. Habitus. Body elongate, somewhat laterally compressed; head flattened anterior- ly, well distinct from the neck; snout depressed and elon- gate; nostril oval, large, in the middle of the nasal. Eye moderate, with a vertically elliptic pupil. SVL 374 mm; TaL 89 mm; TL 463 mm. ©ZFMK 292 Gernot Vogel & Patrick David Fig. 5. Ventral view of Lycodon fasciatus. CIB 9804, from Ruili City, Yunnan. Note he whitish colouration of the anterior part and te speckling of the posterior part. Photograph by Ger- not Vogel. Dentition. A total of 10 maxillary teeth, with the follow- ing formula: 4 small anterior teeth + 2 strongly enlarged teeth + a wide gap + 2 small teeth + a small gap + 2 strong- ly enlarged, posterior teeth. Body scalation. 201 VEN (+ 2 preventrals), 68 SC, all paired. Anal single. Dorsal scales in 17—17—15 rows, the 7 upper rows feebly keeled. Vertebral row not enlarged. No apical pit detected. Head scalation. Rostral, triangular, hardly visible from above; nasal vertically divided by a furrow below and above the nostril; two small, trapezoidal internasals, wide- ly in contact each with the other and with prefrontals; two large prefrontals, longer and wider than internasals; a rather small ogive-shaped frontal, about 1.3 times longer than wide and about 0.8 time as long as the suture between the parietals; 2 large parietals, each edged with three large scales, 2 upper temporals and a larger paraparietal poste- riorly; | / 1 wide, triangular supraocular; 1 / 1 small pre- ocular, located above the posterior part of loreal; 2 / 2 pos- toculars, about the same size; | / 1 subrectangular loreal, elongateand narrow, not entering orbit, in contact with SL 2 and 3, the large preocular, the prefrontal (long contact) and the posterior part of nasal; 8 / 8 SL, of which the first four are higher than long, SL 1—2 in contact with nasal, SL 3-5 entering orbit, 6 and 7 SL largest; 2+2 / 2+2 temporals, lower anterior temporal a bit broader than up- per one, posterior temporals smaller; 8 / 8 infralabials, IL 1-4 in contact with the first pair of sublinguals; anterior and posterior pair of sublinguals of about same length, but anterior pair wider. Coloration in preservation. Body and tail dark blackish- brown, with 31 crossbands on body and 9 on tail, narrow and cream, not speckled; these crossbands, about | dor- sal scale long, widen at their ventrolateral limit, up to about 5-7 dorsal scale long; the first crossband, beginning at the level of VEN 9; the second crossband is 8 scale rows be- hind the first one; 9 cream crossbands on the tail, also not speckled. The head is uniformly blackish-brown, a broad nuchal col- lar extends from the 6‘ and 7‘ supralabial across the low- er posterior temporal across the posterior half of the pari- entals. The underside of the head ist dark in the anterior half and cream in the posterior one; the throat is cream, with a dark clowdy spot on the preventrals and the first ventral. The venter is dark, with faint cream bands rather regular, 2 ventrals wide and with 34 ventrals in between. With- in these bands some ventrals are dark on one half and cream on the other, especially in the posterior part of the body. The under surface of the tail is banded as the ven- ter with the cream bands about 1.5 SC wide. Fig. 6. Comparison of lateral head scalation of Lycodon synaptor sp. n. (BM 1905.1.30.63 Holotype) and one L. fasciatus (BNHS 1223) where the Lo touches the eye. This is the case only in 6.3% of all cases seen by us. Please note that in L. synaptor sp. n. the Lo is well separate from the eye by the preocular scale, whereas it is tapering and narrow in the L. fasciatus, where it is more or less inserted between 2 sales and falls short from the eye. Drawings by Dick Visser. Bonn zoological Bulletin 57 (2): 289-296 ©ZFMK New Lycodon from Yunnan 293 Table 2. Pholidosis of the two type type specimens of Lycodon synaptor sp. n. BMNH_ 1905.1.30.63 Characters MNHN_ 1905.0283 holotype paratype Sex Female Female SVL 374 395 TaL 87 92 Reine 0.192 0.189 ASR 17 15 MSR 17 Ig) VEN 201 203 SC 68 69 Lo enters eye no no Dorsal bands 31 30 Tail bands 9 9 First band at VEN no 9 5) Broad base [VEN] fi gS} Broad vertebral [Dorsals] 2 l Variation. The paratype, MNHN 1905.0283, agrees in most respects with the description of the holotype with the throat being dark instead of light. The maxilla are miss- ing in the paratype. A comparison of the most important morphological characters is summarized in Table 2. Distribution. China. Lycodon synaptor sp. n. is current- ly only known from the region of Dongchuan, Dongchuan County, in the province of Yunnan, China. Biology. There is no information available on the biolo- gy of this species. However, the region of Dongchuan is highly mountainous. Dongchuan is located between high mountains of the ranges known as Gongwang Shan and Liangwang Shan. In the area, the highest point is 4.344 meters high, and lowest is 695 meters asl. DISCUSSION Lycodon synaptor sp. n. is superficially similar to L. fas- ciatus but differs from the whole L. fasciatus group (in- cluding L. butleri) by the loreal, which does not enter or- bit in L. synaptor. We have examined six specimens (out of 35) of Lycodon fasciatus sensu stricto in which the lo- real does not enter orbit (eight occurrence, three times on both sides [4.7 %]), but the morphology of the anterior region of the eye is different. In specimens of L. fascia- tus in which the loreal does not enter orbit, the posterior region of this narrow scale is very tapering (Fig. 6). Its apex is more or less inserted between the preocular and the 4th SL and falls short from the eye. In contrast, in L. Bonn zoological Bulletin 57 (2): 289-296 synaptor, the loreal scale is well separated from the orbit by a broad preocular. The tail is a bit shorter in L. synap- tor sp. n. (0.189—0.192 vs. 0.190—0.219 in 29 females of L. fasciatus). There are also differences in the shape of the bands and the colouration of the belly (compare Figs 2 and 5). L. synaptor sp. n. has eight infralabials, whereas only one specimen out of 60 specimens of L. fasciatus had eight infralabials on both sides and rarely that character is seen on one side (5 %) in L. fasciatus. L. synaptor sp. n. has less subcaudals than L. fasciatus (68—69, x = 68.5, versus 70-88, x = 79.9 with one outlier having 67 subcaudals). For a complete comparison of scale counts see Table 3. L. synaptor sp. n. differs from the other Chinese and In- dochinese species as follows: from L. subcinctus by the fact that in L. synaptor sp. n. both a loreal and a preocu- lar are present. From L. laoensis, L. zawi and L. capuci- nus it differs by the anal shield which 1s single in L. synap- tor sp. n. but divided in the latter two species. Furthermore the colouration is much different. From the L. ruhstrati group it differs by the colouration of the bands (small, reg- ular, completely light in L. synaptor sp. n., getting broad- er posteriorly, irregular and partly speckled with brown in the L. ruhstrati complex), by the number of infralabi- als (eight in L. synaptor, nine to ten in the L. ruhstrati group) and by the colour of the belly, which is speckled or uniform light in the L. ruhstrati group, but clearly band- ed in L. synaptor sp. n. From L. paucifasciatus Rendahl, 1943, occurring in Vietnam, L. synaptor sp. n. differs by the number of anterior dorsal scale rows (15-17 in L. synaptor sp. n. and 19 in L. paucifasciatus). ©ZFMK 294 Gernot Vogel & Patrick David Table 2. Important characters in the Lycodon fasciatus / ruhstrati groups. Lycodon fasciatus synaptor sp.n. ruhstrati futsingensis cardamomensis n females 35 2 23 6 l TL, females N=29 N=2 N=22 N=5 N=1 679 487 876 WB 545 Rel TL, females 0.190-0.219 0.189-0.192 0.207—0237 0.205—0.217 0.25 N=29 N=2 N=22 N=11 N=1 VEN, females 180-219 201-203 217-229 198-208 223 N=35 N=2 N=23 N=13 N=1 SC, females (67) 70-88 68-69 90-108 78-85 92 N=29 N=2 N=21 N=5 N=1 IL both sexes 9-10 (8)* 8 10 (9.11) 9-10 (11) 10 N=120 N=2 N=86 N=44 N=4 Bands 19-43 30-31 19-46 19-33 12-13 N=60 N=2 N=45 N=22 N=4 Tail bands 7-21 9 10-23 9-18 6 N=62 N=2 N=43 N=21 N=4 First band 5-18 5-9 8-17 13-23 unknown N=51 N=2 N=34 N=18 Broad base 3-12 3 5-9 (12) 5-8.5 unknown N=51 N=2 N=34 N=18 Lo enters orbit Yes (rarely not**) No No (rarely yes***) — No No N=126 N=4 N=86 N=44 N=4 Belly banded Yes Yes No No No N=63 N=2 N=25° N=22 N= “8 in 6 cases (5 %), in one specimen from Shillong on both sides (1.6 %) “not entering in 8 occurrences (6.3 %), three times on both sides (4.7 %) (see above) “in 6 specimens, all from Fujian the Lo touched the eye. FORK banded in juveniles only Werner (1922) described Dinodon yunnanensis from Yun- nanfu, now Kunming, Yunnan Province. This species was synonymised with Lycodon fasciatus by Pope (1935: 188), but according to our unpublished data, this taxon might be valid. In any way this name is available, so we com- pared Lycodon synaptor sp. n. with Dinodon yunnanen- sis Werner, 1922 for which we re-examined the holotype (NMW 23417; adult female). Lycodon synaptor sp. n. dif- Bonn zoological Bulletin 57 (2): 289-296 fers from D. yunnanensis mainly by the loreal which is touching the eye in D. yunnanensis, but also by the num- ber of ventrals (201—203 against 193) and the number of bands on the body (30-31 vs. 23) and the number of in- fralabials (eight vs. nine in D. yunnanensis). While preparing a review for the Lycodon fasciatus group, we found quite a lot of obviously independent lineages, ©ZFMK New Lycodon from Yunnan 295 which seem to constitute distinct species. Some of them are restricted to small areas. A discussion of these lineag- es will follow in the main review (Vogel & David in prep.). Lycodon synaptor sp. n. differs so much from other mem- bers of the group that we decided to describe it separate- ly. The closest localities we got from Lycodon fasciatus are from the vicinity of Kunming in Yunnan province, which lies about 100 km south of Dongchuan. This latter place (previously Tongchuan or Tongchuan Fu) is the type locality for several reptile species, as Cuora yunnanensis (Boulenger 1906), Nanorana yunnanensis (Anderson, 1879), Odorrana grahami (Boulenger 1917) and Bombi- na maxima (Boulenger 1905). It is a relatively small city that had about 300.000 inhabitants in 2006. Revised Key for the genus Lycodon in China According to our data, there are several unnamed species of the genus Lycodon living in China. So this should be regarded as preliminary key. 1. Both a loreal and a preocular scales present ................006+ 2 Either loreal or preocular absent ................0... L. subcinctus Jao ANTE CRAIGS cocosligenascase08s ants arose sc cICGICE ae TE Coer aE EERE 3 ANTE SITIES. scoedectngseaaboReSERE DEES E ae PREEEL EERE SEER 4 3. Anterior chinshields not more than 1.5 times longer than posterior ones; loreal in extensive contact with internasal, when adult no crossbands on body ................- L. capucinus Anterior chinshields 2 to 3 times longer than posterior ones; loreal not, or barely in contact with internasal (very rarely a strong contact), when adult yellow crossbands on forepart Olt OOGH?. accoonsidoscosteonee noseee Lecce eee MEER cae L. laoensis G\, TRIG? TORIC IEC scconcsasaccsbaccesco sco udentaaqsosacencecacode oguceciasscnosaere sae 5 IBIS Gt lO eTOVET EC | sce condnscecschoseoodacoae cers aondonoossspacbooosoriaaaeocHnesE 6 5. Lo not touching the eye, 8 lower labials .. L. synaptor sp. n. Lo usually touching the eye, usually 9-10 lower labials 20000600000600¢00 00 00H IEE DEE ECE: CEE CEE CUR ee ESCs L. fasciatus GaDorsallmows| keeled «.......2:2--s-ccs-ceeecneso-00e L. ruhstrati abditus Dorsalirows'SMOOth 2.2. ...c-c.cceccsecsencsestereswees L. futsingensis Bonn zoological Bulletin 57 (2): 289-296 Acknowledgements. The first author is indebted to Annemarie Ohler and Alain Dubois (MNHN) for the grants to work in the Paris collection. We are grateful to Silke Schweiger, NMW for sending pictures of the type of Dinodon yunnanensis. Ke Jiang and Jian Luo helped a lot with further information on Chinese Lycodon. Montri Sumontha provided scalecounts. We also thank Colin J. McCarthy (BMNH), Varad Giri (BNHS), Robert C. Drewes and Jens V. Vidum (CAS), Wang Yuezhao, Zeng Xiao- mao and Ermi Zhao (CIB), Alan Resetar (FMNH), Dingqui Rao (KIZ), Ivan Ineich and Annemarie Ohler (MNHN), Franz Tiede- mann and Richard Gemel (NMW), Wolfgang B6hme (ZFMK), Mark-Oliver Rédel and Frank Tillack (ZMB), Frank Glaw and Dieter Fuchs (ZSM) for the possibility to examine specimens de- posited in the collection of their respective institutions. Dick Vis- sers made the drawings for us. Many thanks for that. REFERENCES Daltry JC & Wiister W (2002) A new species of Wolf Snake (Ser- pentes: Colubridae: Lycodon) from the Cardamon Mountains, southwestern Cambodia. Herpetologica 58: 498-504 Dowling HG (1951) A proposed standard system of counting ventrals in snakes. British Journal of Herpetology 1: 97-99 Gaulke M (2002) A new species of Lycodon from Panay Island, Philippines (Reptilia, Serpentes, Colubridae). Spixiana 25: 85-92 Lanza B (1999) A new species of Lycodon from the Philippines, with a key to the genus (Reptilia Serpentes Colubridae). Trop- ical Zoology 12: 89-104 Mukherjee D & Bhupathy S (2007) A new species of Wolf Snake (Serpentes: Colubridae: Lycodon) from Anaikatti Hills, West- ern Ghats, Tamil Nadu, India. Russian Journal of Herpetol- ogy 14: 21-26 Ota H & Ross CA (1994) Four new species of Lycodon (Ser- pentes: Colubridae) from the Northern Philippines. Copeia 1994: 159-174 Pope CH (1935) The reptiles of China. Turtles, crocodilians, snakes, lizards. Natural History of central Asia, X. American Museum of Natural History, New York Slowinski JB, Pawar SS, Win H, Thin T, Gyi SW, Oo SL, Hla T (2001) A new Lycodon (Serpentes: Colubridae) from north- east India and Myanmar (Burma). Proceedings of the Califor- nia Academy of Sciences 52: 397-405 Vogel G, David P, Pauwels OSG, Sumontha M, Norval G, Hen- drix R, Thanh VN, Ziegler T (2009) A revision of Lycodon ruhstrati (Fischer, 1886) auctorum (Squamata: Colubridae), with the description of a new species from Thailand and a new subspecies from the Asian mainland. Tropical Zoology 22: 131-182 Werner F (1922) Neue Reptilien aus Stid-China, gesammelt von Dr. H. Handel-Mazzetti. Anzeiger der Akademie der Wis- senschaften in Wien 59: 220-222 Received: 24. VIII.2010 Accepted: 10.X.2010 OZFMK 296 Gernot Vogel & Patrick David APPENDIX I Additional comparative material of Lycodon fasciatus India. Shillong, Assam, BMNH 92.1.25.1; BMNH 1908.6.23.6; BMNH 1908.6.23.8; BMNH 1907.12.16.28-29; BMNH 94.10.4.2; BNHS 1219-20, 1229. Laos. Xieng-Khouang, Laos, MNHN 1928.69. Myanmar. Maymyo, Burma, BMNH 1925.12.22.15—16; Mogok, Burma, BMNH_ 1900.9.20.5—7; BMNH 1901.4.26.1Mogkok, Burma, BNHS 1221 “Burma”, BMNH 1908.6.23.9-11 ; Burma, BNHS 1218; Toungyi, Shan State, Myanmar, BMNH 91.11.26.31; Bhamo, Bur- ma, BMNH 1925.4.2.28; Kachin Hills, Burma, BMNH 1925.9.17.10—11; South Shan State, BMNH 1908.6.23.14; Burma-Siam Border, BMNH 1937.2.1.12; Rangoon, Bur- ma, BMNH_ 1940.3.3.3; Maymyo, Burma, BNHS 1222-24; Thandung Hills, BNHS 1228. Bonn zoological Bulletin 57 (2): 289-296 People’s Republic of China. Western China CAS 55147; Yunnan, MNHN 1919.148; “Yunnan Fu” (holotype of Dinodon yunnanensis), NMW 23417; Kuantun, ZSM 75/1938; Kunming, Yunnan, BMNH 1930.11.16.4; Ruili City, Yunnan, CIB 9804; RuiLi, Yunnan,, CIB 9805; XiShuangBanNa, Yunnan, CIB 9806, CIB 9808, CIB 9809; LongChuan GongWa, Yunnan, KIZ 74 I 0035; LongChuan HuSa, Yunnan, KIZ 74 I 0145; Tengchong County, Yunnan KIZ 74 II 0262; Menglian, Yunnan, KIZ 75 1473; TengChong TuanTian, Yunnan, KIZ 74 I 0263; Kunming, Yunnan, KIZ 73009; Kunming, Yunnan, KIZ 77004; Kunming city, Yunnan, KIZ 83007; Yunnan, KIZ 83017; FMNH 15148; Tibet; MNHN 1921.0465 Tibet; Yunnan,; ZMB 65453 Thailand. Chiang Mai, Thailand, FMNH 178369; CAS 172715, Southern Thailand ? FMNH 178368, Nan province Thailand FMNH 270716. Vietnam. Phong Nha, Vietnam, ZFMK 86448—S0 (Gen- Bank: EU999214-215); ZFMK 80665; Ziegler unreg. ©ZFMK Bonn zoological Bulletin | Volume 57 Issue 2 pp. 297-306 Bonn, November 2010 A crowned devil: new species of Cerastes Laurenti, 1768 (Ophidia, Viperidae) from Tunisia, with two nomenclatural comments Philipp Wagner!* & Thomas M. Wilms? ' Zoologisches Forschungsmuseum Alexander Koenig, Adenauerallee 160, 53113 Bonn, Germany; philipp.wagner.zfmk@uni-bonn.de ? Zoologischer Garten Frankfurt, Bernhard-Grizmek-Allee 1, 60316 Frankfurt a. Main, Germany; * corresponding author Abstract. A distinctive new species of the viperid genus Cerastes is described form Tunisia. It is closely related to Cerastes vipera but easily distinguishable from this invariably hornless species by having tufts of erected supraocular scales form- ing little crowns above the eyes. These crown-like tufts consist of several vertically erect, blunt scales which differ dras- tically from the supraocular horns of C. cerastes or C. gasperettii that consist of one long, pointed scale only. Although the new species is based on only one single specimen, further specimens had originally been available but were subse- quently lost in private terraria. The taxonomic status of the nomen ‘“Cerastes cerastes karlhartli” is discussed and the name is found to be unavailable (nomen nudum). Also the authorship of “Cerastes cornutus” is discussed and ascribed to Boulenger. Key words. Cerastes cerastes, Cerastes vipera, Cerastes sp. n., Cerastes c. karlhartli, Cerastes cornutus, horned viper, North Africa, Tunisia. INTRODUCTION The genus Cerastes Laurenti, 1768 includes only five taxa (three species and two subspecies), which are distributed in northern Africa and on the Arabian Peninsula. All species are stout-bodied and, as desert snakes, are char- acterized by many xeromorphic physiological and mor- phological adaptations. The most impressive adaptations are the strongly keeled serrated lateral body scales, a char- acter they share with their proposed sister taxon Echis (Joger & Courage 1999, Pook et al. 2009), but not with the also sand living snakes of the genus Bitis. The largest member of the genus is Cerastes cerastes (Lin- naeus, 1758) with a maximum body size of 80 cm and an average size of 35 to 60 cm. The distribution range of the nominotypic form includes all Saharan countries with a southernmost distribution in Sudan (Phelps 2010) and the northernmost in central Tunisia (Schleich et al. 1996). Eastwards it reaches the Sinai, Israel and Jordan (Phelps 2010). Cerastes c. cerastes occurs in sandy and rocky deserts and around well vegetated oases but not in wind- blown dunes (Phelps 2010). Cerastes c. hoofienii Wern- er & Sivan 1999, the second subspecies occurs in the ex- treme southwestern edge of the Arabian Peninsula in Yemen and Saudi Arabia. Bonn zoological Bulletin 57 (2): 297-306 The second North African species is C. vipera (Linnaeus, 1758). Its distribution range is very similar to C. cerastes but more restricted to the Saharan Desert and reaches east- ward to the Sinai and Israel as it only occurs in dune sys- tems. Therefore, according to Phelps (2010) both species were never recognized as locally syntopic, but Joger (2003) found both species occurring parapatrically at the edge of the Grand Erg Oriental in Tunisia. Cerastes vipera is the smallest viper of the genus and grows up to a max- imum size of under 50 cm with an average about 35 cm. The remaining species is Cerastes gasperettii Leviton & Anderson, 1967 with its subspecies gasperettii and C. g. mendelssohni Werner & Sivan, 1999. It is distributed on the Arabian Peninsula and eastwards to Iraq and Iran, over- lapping with other Cerastes species in its distribution on- ly in the southern Sinai and the northwestern edge of the Arabian Peninsula. The common name ‘Horned Vipers’ is misleading as not all species and not all populations possess supraocular horns. In C. cerastes and C. g. gasperettii specimens usu- ally bear horns but several populations are hornless. If present, the horns are formed by a long, sulcated, single ©ZFMK 298 Philipp Wagner & Thomas M. Wilms spike above each eye, usually surrounded by a ring of elongate spinose but non-sulcated scales. This polymor- phism is also known in other viperid snakes as the supraocular horns of Bitis caudalis and the supranasal horns of Bitis gabonica are absent in some specimens (FitzSimons 1962, Branch 1988). However, C. g. mendelssohni and C. vipera are strictly hornless as op- posed to the other taxa. Gasperetti (1988) described the characters of the genus Cerastes as (a) the eyes are small to moderate and sepa- rated from the labial scales by several rows of small scales; (b) body scales with club or anchor shaped keels, not ex- tending to the extremity of the scales; (c) lateral body scales smaller than vertebral scales; (d) anal scale entire; and (e) ventral scales with an obtuse keel on each side. For many decades, only two species, C. cerastes and C. vipera, were recognized but Werner (1987) and Werner et al. (1991) elevated C. gasperettii to full species status, which was later accepted by many authors (e.g. Schatti & Gasperetti 1994, Phelps 2010). Following Baha el Din (2006) the two African species are easy to distinguish. In C. vipera, supraocular horns are never present, there are less than 14 interorbitals and counts of ventral scales are below 120, whereas in C. cerastes there are more than 14 interorbitals and more than 130 ventrals. Schleich et al. (1996) distinguished the two snakes mainly by the position of the eye (lateral in cerastes and directed upwards in vipera), by the presence or ab- sence of a supraocular horn and by the number of interor- bital scales (15—21 in cerastes and 9-13 in vipera). Ge- niez et al. (2004) distinguished both species by their lon- gitudinal rows of dorsal scales at midbody (27-35 in cerastes; 23—27 in vipera), but also mentioned the eyes on the top of the head in C. vipera. In the collection of the Zoologisches Forschungsmuseum Alexander Koenig (ZFMK) a specimen of Cerastes is present, which is generally similar to C. vipera but clear- ly distinct in possessing supraocular crown-like scale tufts instead of horns of a solitary scale. Because of this strik- ing character, as such tufts or horns are absent in C. vipera, the specimen was examined and compared with other Cerastes specimens of the ZFMK collection and with rel- evant literature. MATERIAL & METHODS This description is based on the comparison of 75 pre- served Cerastes specimens and three vouchers of other snakes of the ZFMK collection (see below) and the rele- vant literature (Jooris & Fourmy 1996; Schleich et al. 1996; Geniez et al. 2004; Baha el Din 2006; Phelps 2010). Bonn zoological Bulletin 57 (2): 297-306 Measurements were taken with a digital-calliper to the nearest 0.1 mm. The number of ventral scales was count- ed excluding the anal scale. The number of subcaudals in- cluded the terminal scale. The dorsal scale row count is given as (a) fore body: one head length behind head, (b) midbody at the level of the ventral plate corresponding to a half of the total number of ventrals), and (c) hind body one head length before vent. For SEM images a Hitachi S-2460N was used to compare the scale morphology of different snake species. Dorsal body scales from about the middle of the dorsum were used from the following specimens: Cerastes sp. n. (ZFMK 58054, Tunisia), Bitis peringueyi (Boulenger, 1888) (ZFMK 44887: Namibia, Swakopmund) and Bitis schneideri (Boettger, 1886) (ZFMK 88450: Namibia, without locality). These were compared with SEM-pho- tographs from C. cerastes and C. vipera published by Joger & Courage (1999). Material examined. Cerastes cerastes: ALGERIA: ZFMK 7649-7650, Colomb-Béchar; ZFMK 18082, 60km west of Toug- gourt; ZFMK_ 18083-084, 20km north of Bou-Saada; ZFMK 18085, Hoggar Mts., Guelta Afiale; ZFMK 23000, south of Temassinin, Flatters; ZFMK 23001, Bordj-Saada; ZFMK 23002-005, south of Ouargla; ZFMK 38248, 20km south of Djanet. EGyet: ZFMK 22996, Isna (=Esne); ZFMK 22997, vicinities of Cairo; ZFMK 50295, Aswan desert; ZFMK 50296, Faijum desert; ZFMK 50299-300, Nada el Wahda desert; ZFMK 32488, 50297-298, without locality. LiByA: ZFMK 63668, Wa- di Matendus; Mauretania: ZFMK 17593, Chami. Morocco: ZFMK 65218, Draa Valley. NIGER: ZFMK 20258, between Ar- lit and Agadez, 120km south of Arlit; ZFMK 36629, 40km north- east of Wadi Gougaram. SUDAN: ZFMK 32462, 100km south- west of Burget Tuyur depression; ZFMK 32463, Dafur, Teiga Mts., west of Eisa; ZFMK 32464, Darfur, Djebel Rahib; ZFMK 33697, Nubian desert, 130km southeast of Wadi Halfa; ZFMK 33698-700, Wadi Halfa; ZFMK 38410, 80km north of Port Su- dan. TunistA: ZFMK 18081, 10km west of Tozeur, Oasis Stil; ZFMK 22998-999, Tunisian desert; ZFMK 29047, El Hamma du Djerid near Tozeur; ZFMK 29809-812, 29046, Tozeur; ZFMK 47020-024, between Tozeur and Nefta; ZFMK 49858, Oasis Nef- ta. Cerastes gasperettii: IRAQ: ZFMK 18843-844, vicinities of Basrah; ZFMK 19414, Basrah. KINGDOM OF JORDAN: ZFMK 44340, Wadi Araba, Fidan. KINGDOM OF SAUDI ARABIA: ZFMK 43659, 100km northeast of Riyadh. UNITED ARAB EMIRATES: ZFMK 52419, Al-Mundam. Cerastes vipera: ALGERIA: ZFMK 22984, El Alia; ZFMK 41176, Ain Sefra. EGypt: ZFMK 22989- 994, 50339, vicinities of Cairo; ZFMK 22995, Sinai, Wadi Ar- ish; ZFMK 50301-302, El Wasta, Abwid desert. LipyA: ZFMK 32489, Tripolis. MAURETANIA: ZFMK 17594, Chami. TUNISIA: ZFMK 22985-988, Tunesian Sahara, without locality. WESTERN SAHARA: ZFMK_ 83340, Laayoune Plage; Bitis caudalis: NAMIBIA: ZFMK 65212, Swakopmund; Bitis peringueyi: NAMIBIA: ZFMK 88453, without locality. Bitis schneideri: NAMIBIA: ZFMK 88450, without locality. ©ZFMK A new species of Cerastes from Tunisia F500 um—4 Ghee NA WT 1mm | PERING, + 200 pm 4 Fig. 1. _ SEM images of dorsal body scales of viperid snakes. 1= Cerastes sp. n. from central Tunisia (ZFMK 58054); 2= Cerastes cerastes published by Joger & Courage 1999; 3= Cerastes vipera published by Joger & Courage 1999; 4= Bitis peringueyi, 4A published by Joger & Courage 1999, 4B & 4C from Namib- ia (ZFMK 44887), and 4A= published by Joger & Courage 1999; 5= Bitis schneideri from Namibia (ZFMK 88450). A= dorsal scale, complete; 1B= dorsal scale, verrucate, secondary structure; 2B= dorsal scale, verrucate, secondary structure; 3B= dorsal scale, liniar, tertiary structure; 4B= dorsal scale, verrucate to cristate, secondary structure; 5B= dorsal scale, cristate, sec- ondary structure; 1C= dorsal scale, secondary structure in detail; 2C= dorsal scale, secondary structure in detail; 3C= dorsal scale, secondary structure in detail; 4C= dorsal scale, secondary structure in detail; 5C= dorsal scale, secondary structure in detail. Bonn zoological Bulletin 57 (2): 297-306 ©ZFMK 300 Philipp Wagner & Thomas M. Wilms RESULTS & DESCRIPTION The comparison of the fine structure of a dorsal, non-rat- tling scale of the single specimen with images of lateral, rattling scales of Cerastes and lateral scales of Bitis (see fig. 1) shows similarities between the single specimen and Cerastes. On the other hand, differences of the typical scale fine structure of Bitis are distinct enough to recog- nize the single specimen as a non-Bitis species. The struc- ture typical in Bitis specimens is obvious in Bitis schnei- deri (fig. 1.5). They possess a structure of slender, elon- gated bulges, which are very distinct from Cerastes. How- ever, Bitis peringueyi, (fig. 1.4) also a sand burrowing snake, is the only species of Bitis showing a similar scale structure to Cerastes species, but obvious from figure 2, this species is very distinct from the new species of Cerastes. However, all Cerastes species are similar in their verru- cate secondary structure; cell borders are well visible. These borders are invisible in the cristate or verrucate sec- ondary structure of Bitis species. The comparison of the voucher with C. vipera and C. cerastes results ina morphological similarity to C. vipera. Both are similar in body size, shape of the nostril and head scalation (see fig. 2, tab. 1). However, in other aspects they are clearly distinct: the specimen has lower scale counts as in C. vipera in its morphological variation of the en- tire distribution in northern Africa. The specimen possess- es supraocular horns, which are absent in C. vipera and horns, encompassing several scales, are also not known in C. cerastes. Therefore, we regard this specimen as a new species of Cerastes: Cerastes boehmei sp. n. Holotype. ZFMK 58054. Female specimen from Tunisia, SW Remada, east of Djebel National Park, close to the road midway between Beni Kadeche (Bani Kheddache) and Ksar el Hallouf, leg. T. Holtmann, 1991. Diagnosis. This new species of Cerastes 1s characterized by: (a) head depressed, eyes on the lateral part of the head but slightly directed upwards; (b) supraocular coronets (crowns) present, consist of several sulcated, medium sized scales, instead of the supraocular horn formed by a single sulcated long scale in C. cerastes or C. gasperettii; (c) nostril slit-shaped; (d) low number of interorbital scales; (e) 19-26-16 dorsal scale rows around fore-, mid- and hind body. Differential diagnosis. The new species differs from (a) C. vipera in possessing supraocular coronets, a low num- Bonn zoological Bulletin 57 (2): 297-306 ber of interocular scales (7 instead of 9-13 fide Schleich et al. [1996], but 6—13 fide Jooris & Fourmy [1996]), a lower number of circumocular scales (11 instead of 19-29 fide Jooris & Fourmy (1996), a lower number of supral- abial/infralabial scales (11—12/12-11 instead of 20-33/19-27 fide Jooris & Fourmy [1996]) and a lower number of subcaudal scales (25 instead of 33—57 fide Jooris & Fourmy [1996]). Counts of dorsal scale rows around midbody are ambiguous and depending on the method of counting (see fig. 3). They differ from C. vipera (21 instead of 23—27 fide Phelps [2010]) or lie with 26 scale rows within this range; from (b) C. cerastes in a low- er number of interocular scales (7 instead of 15—21, fide Schleich et al. [1996]), in its smaller size, in possessing a slit-shaped nostril, in possessing supraocular coronets each formed by more than one elongated scale; and final- ly from (c) C. gasperettii in possessing supraocular coro- nets each formed by more than one elongate scale and in possessing a slit-shaped nostril. From all recognized synonyms of C. vipera (mainly C. vipera inornatus Werner, 1929 and C. richiei Gray, 1842) the new species differs in possessing supraocular coronets, whereas the synonymised taxa are lacking horns or equiv- alent structures. From the recognized synonyms of C. cerastes (mainly C. c. mutila Doumergue, 1901) the new species differs in possessing supraocular coronets, as all synonymised taxa are lacking horns or equal structures. Following Boulenger (1896) Cerastes cornutus Boulenger, 1896 (see also nomenclatural comment as part of the discussion), regard- ed as a synonym of C. cerastes following e.g. Schleich et al. 1996, is also with either horn-bearing or hornless in- dividuals, but differs from C. boehmei sp. n. in a higher number of interorbital scales (15 to 21), in possessing supraocular horns made up of a single scale, a higher num- ber of scale rows around midbody (27-35) and a higher number of ventral (130-165) scales. Description of holotype (fig. 4). Habitus. Body elongate and slender, somewhat compressed and oval in profile; head flattened, triangular and well distinct from neck; Eye small to moderate, with vertically elliptical pupil, on up- per lateral side, but nearly on top of head; nostril slit- shaped, slightly longer than first supralabial scale. Measurements (in mm): Total length: 218.5; head length: 16; head width: 9.4; head height: 5.5; snout-vent length: 195; tail length: 25.5. Scalation of head: Rostral broader than high, semicircu- lar, slightly visible from above; menthal scale only in con- tact and smaller than first infralabial scale, followed by two large chinshields; nasal scale divided by large scale ©ZFMK A new species of Cerastes from Tunisia 301 Fig. 2. | Comparison of different viperid snakes from Africa. 1= Cerastes boehmei sp. n.: ZFMK 58054, Tunisia; 2= Cerastes vipera: ZFMK 22985, Tunisia, without locality; 3= Cerastes vipera: ZFMK 83340, Western Sahara, Laayoune Plage; 4= Cerastes cerastes: ZFMK 63668: Libya, Wadi Matendus; 5= Cerastes cerastes: ZFMK 65218, Morocco, Draa Valley; 6= Bitis caudalis: ZFMK 65212: Namibia, Swakopmund; 7= Bitis peringueyvi: ZFMK 88453: Namibia, without locality. A= head in profile; B= head from above; C= head from below. Bonn zoological Bulletin 57 (2): 297-306 ©ZFMK 302 Philipp Wagner & Thomas M. Wilms Table 1. Comparison of the three Cerastes species occurring in northern Africa. C. boehmei sp. n. C. vipera* C. cerastes** Interorbital scales Vl Ventral scales 110 Subcaudal scales 2S Position of the eye lateral Supraocular horn present Scale rows around midbody 21 (26)*** Circumocular scales 11 less than 14 (6-13) more than 14 (14-21) below 120 more than 130 33-57 18-42 directed upwards lateral absent present/absent 23-37 27-35 19-29 - *=fide Jooris & Fourmy 1996, Schleich et al. 1996, Baha el Din 2006, Phelps 2010. **= fide Schleich et al. 1996, Baha el Din 2006, Phelps 2010. ***= see fig. 3 bearing nostril at its upper fringe, with smaller overlay- ing scale; five, more or less trapezoidal internasal scales, the two outer scales twice as large as three inner scales, all keeled; no enlarged prefrontal scales; occipital tuber- cle absent; supraocular coronets present, consist of elon- gate, sulcate scales, four on left, five on right side; 11 cir- cumocular scales on each side; interorbital scales 7 at mi- dlevel of supraocular coronets; loreals 3 on each side; supralabial scales: 11 on left, 12 on right side, only first in contact with nasal scale, three scales between supral- abial scales and eye (including ocular scale); infralabial scales: 12 on left, 11 on right side. Scalation of body: Ventral scales: 110; subcaudal scales: 25; number of scale rows around fore-body: 19, mid-body: 21 or 26 (see fig. 3), hind-body: 16; vertebral row not en- larged, in 107 scales on body. Colouration in preservative. After 20 years of preserva- tion in ethanol, the specimen has more or less uniform sandy colouration. Head uniform, upper side down to height of supralabial scales sandy, underside up to height of infralabial scales dirty white; body sand-coloured, with irregular pattern of slightly darker blotches; upper fore part of tail banded dark sand-coloured, underside dirty white, from mid-tail to tip uniform dark, nearly black on both sides; belly dirty white. Colouration in life. Similar to preserved specimen: uni- form yellowish sand-coloured with shades of some slightly darker blotches. Head and forepart of body uni- form yellowish-sandy, without darker pattern (see fig. 5). Etymology. This new species is named, in deep respect, after our ‘scientific father’ Prof. Dr Wolfgang Bohme, deputy director and head of the Herpetology section at the Zoologisches Forschungsmuseum Alexander Koenig in Bonn, for his contributions to African herpetology for the Bonn zoological Bulletin 57 (2): 297-306 past four decades and for the time he invested in his young students. With his encouraging lectures, discussions, ex- cursions and fieldtrips he had a significant influence on the authors leading to their scientific current dedication with herpetological systematics, ecology and zoogeogra- phy. Distribution. So far only known from the type locality, but an adult male was caught by a local snake hunter near Beni Kadeche (T. Holtmann pers. comm.). The new species appears currently to be endemic to Tunisia and is probably widespread in the area of Bani Kheddache. Biology. Nearly no information is available on the biol- ogy of this species. In captivity the adult female gave birth to living young which reflects the close relationship to C. vipera. In respect to colouration, a sandy habitat can be assumed. Comments. Although the species is described here based on a single voucher, more specimens were known but be- came apparently lost. This specimen was one of three ju- veniles caught together with an adult female at the type locality. The adult female had five juveniles in captivity. Additionally, an adult male was caught in the area of the type locality by a local hunter. All of these specimens have shown the described character of the unique supraocular horns. DISCUSSION Though described from a single specimen only, the valid- ity of C. boehmei sp. n. is beyond doubt. As becomes ob- vious from the comparison of fine structure of body scales, the new species must be clearly assigned to the genus Cerastes, being distinct from burrowing Bitis species from southern Africa. However, the fine structure is similar to ©ZFMK A new species of Cerastes from Tunisia 303 Fig. 3. both, C. cerastes and C. vipera but many other characters (e.g. shape of nostril, position of the eye, pholidosis, re- productive biology) show that the new species is more closely related to C. vipera than to C. cerastes. Accept- ing the results of the morphological analysis of C. vipera published by Jooris & Fourmy (1996) the new species has lower counts in circumocular, subcaudal, supralabial, in- fralabial scales and of ventral and interocular scales are on the lower limit of morphological variation in C. vipera. Therefore, C. boehmei sp. n. is clearly distinct from C. vipera also in pholidosis. Nevertheless, in C. cerastes both hornless and horned individuals are known but (a) the morphology of the “crowns” of C. boehmei sp. n. is clear- ly distinct to all other known horn structures in Cerastes and (b) the fact that supraocular horns or similar structures are completely unknown in C. vipera strengthens the va- lidity of the new taxon as new and full species. Also Jooris & Fourmy (1996) who analysed 246 specimens compar- ing pholidosis in relation to a directed distribution did not mention any individuals with horn-like structures. Also none of the known synonyms of C. vipera possesses horns or similar structures. Nevertheless, supraocular horns as spontaneously mutation are extremely implausible. Only one case is documented where a specimen of Macrovipera lebetina possessed a solitary horn only on one side of the head (Bohme & Wied! 1994). Nevertheless, in C. cerastes and C. gasperettii hornless and horn-bearing individuals are known and a taxonom- ic differentiation is only known from C. gasperettii where the subspecies mendelsohni is hornless. Therefore, it can be also assumed that the supraocular scale tufts are sim- ply a so far unknown variation within of C. vipera. But first of all, C. vipera is a well known species and e.g. Jooris & Fourmy (1996) have analysed a high number of vouchers and no single specimen is known which possess Bonn zoological Bulletin 57 (2): 297-306 Two different methods to count scale rows of dorsal scales around the midbody region of the holotype of Cerastes boehmei sp. n., ZFMK 58054. supraocular tufts and second these supraocular tufts are strongly abnormal and very distinct to the supraocular horns of cerastes and gasperettii who possess similar supraocular horns to each other. The function of supraocular horns remains unknown. There were many speculations on the function of the horns in Crotalus cerastes Hallowell, 1854 from America. Klauber (1956) mentioned that they serve as radiators of heat or shaders for the eyes, whereas Cowles (1953) re- garded them simply as a whim of evolution and Cohen & Myres (1970) suggest that they have the function of an eyelid protecting the snake’s eye while passing through burrows. They supported this hypothesis with an ecolog- ical comparison between C. cerastes and C. vipera: the former 1s only known to bury itself partially and frequent rodent burrows whereas C. vipera is only known to bury itself fully in sand and is not reported from rodent bur- rows. However, another, not yet discussed, function could be a sexual recognition between the two snakes. In many reptile groups (e.g. Chamaeleonidae, Agamidae) body or- naments are known for identification during mating time. Although only males possess in most cases ornaments, it should be verified if only those population of C. cerastes (as this species does not strictly possess horns) bear horns, which occur directly syntopically with the hornless C. vipera. Currently the new species is only known from central Tunisia and a restricted distribution can be assumed. A similar case is found in Pseudocerastes urarachnoides Bostanchi, Anderson, Kami & Papenfuss, 2006 which was described from a small area and a further study (Fathinia et al. 2009) found a third locality relatively close the lo- calities of the types only. ©ZFMK 304 Philipp Wagner & Thomas M. Wilms Fig. 4. Holotype of Cerastes boehmei sp. n.: ZFMK 58054 from SW Remada, Tunisia. Bonn zoological Bulletin 57 (2): 297-306 ©ZFMK A new species of Cerastes from Tunisia 305 Fig. 5. The SEM analyses of dorsal scales show that Bitis peringueyi is different to other Bitis species in its second- ary structure (Beyerlein 1993, and fig. 1). This structure is more or less a verrucate structure and not comparable with the cristate structure of e.g. Bitis schneideri, but sim- ilar to the Cerastes species. However, it is distinct to Cerastes as the imprints of borders between Clear layer cells, which are present in Cerastes, are not visible. Both, the Cerastes species and B. peringueyi are moreover sim- ilar in their habitats as B. peringueyi 1s one of the sand- burrowing Bitis species living in windblown sands of the Namib desert. Other burrowing Bitis like e.g. B. schnei- deri (see fig. 1) or B. caudalis (imaged in an unpublished thesis, see Beyerlein 1993) show the typical cristate sec- ondary structur of Bitis. B. schneideri occurs in stable veg- etated sand dunes and not like B. peringueyi and Cerastes in windblown sands. Therefore, this scale structure could be interpreted as an adaptation for this special habitat of windblown sands. Nomenclatural comment to the nomen ‘Cerastes c. karlhartli’. Sochurek (1974) ‘described’ a subspecies of C. cerastes which he called *C. cerastes karlhartli’. How- ever, the description was done in his privately published so-called ‘Herpetologische Blatter’, which according to art. 8.1 of the International Code of zoological Nomen- clature (ICZN 1999) does not constitute a publication and therefore the description is not valid. Later, Sochurek (1979) used the name again in a summary of North African snakes but failed to provide a diagnosis, description or fig- Bonn zoological Bulletin 57 (2): 297-306 Living holotype of Cerastes boehmei sp. n. in captivity. ure and moreover did not designate a holotype. He only mentioned the distribution and a type locality. Later, Tiede- mann & Haupl (1980) accepted the name as a valid sub- species in their herpetological type catalogue of the Nat- ural History Museum in Vienna. Werner & Sivan (1992) placed the ‘subspecies’ in the synonymy of C. cerastes, whereas Golay et al. (1993) placed it in the synonymy of C. gasperettii. Werner et al. (1999) did not mention the name, whereas McDiarmid et al. (1999) and Baha el Din (2006) followed Werner & Sivan (1992). However, all these authors gave neither a diagnosis nor a figure of a specimen. Therefore the name ‘Cerastes cerastes karl- hartli must be recognized as a nomen nudum. Nomenclatural comment to the nomen Cerastes cornu- tus Boulenger, 1896. The name is used for the first time in Forskal (1775, IX), and he is often mentioned as the author of this taxon name (e.g. Schleich et al. 1996, Ba- ha el Din 2006). But Petrus Forskal died during the Dan- ish Arabia expedition, and Carsten Niebuhr published Forskal’s results after his death. Nevertheless, the name is part of a summary about different species which Forskal wanted to describe, but finally never managed to do be- fore his untimely death. Additionally, this nomen is not accompanied by either a description or a drawing. There- fore, Boulenger (1896) who was the first to use the name together with a description must be recognised as the au- thor of Cerastes cornutus, despite the fact that Boulenger himself mentions Forskal as the author of this species. ©ZFMK 306 Philipp Wagner & Thomas M. Wilms Acknowledgements. We are grateful to Thorsten Holtmann (Oberhausen) for providing information about the species and the image of the holotype alive. We are very thankful to Melanie Strauch (Bonn) who provided SEM specimens of the holotype and Bitis schneideri. Wolfgang Bohme (Bonn), Andreas Schmitz (Geneve) and Bradley Sinclair (Ottawa) gave very important comments to the manuscript and we are thankful for fruitful dis- cussions with them. REFERENCES Baha el Din S (2006) A guide to the reptiles and amphibians of Egypt. The American University in Cairo Press, Cairo, 359 pp. Beyerlein P (1993) Systematische Relevanz epidermaler Ultra- strukturen bei modernen Schlangen (Colubroidea). 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Fauna of Saudi Arabia 14: 348-423 Schleich HH, Kastle W, Kabisch K (1996) Amphibians and Rep- tiles of North Africa. Koeltz, Koenigstein, 627 pp. Schmidt KP (1939) Reptiles and amphibians from Southwest- ern Asia. Publ. Field Museum of natural History, zoological Series 24: 49-92 Sochurek E (1974) Cerastes cerastes karlhartli n. spp. Herpe- tologische Blatter 1: 3 Sochurek E (1979) Die Schlangen Nordafrikas. Mitteilungen der Zoologischen Gesellschaft Braunau 3: 219-226 Tiedemann F, Haupl M (1980) Typenkatalog der Herpetologi- schen Sammlung. Reptilia. Kataloge der wissenschaftlichen Sammlungen des Naturhistorischen Museums in Wien (Selbst- verlag Naturhistorisches Museum Wien) 4 (Vertebrata 2): 1— 79 Werner YL (1987) Ecological zoogeography and Saharo-Arabi- an, Saharo and Arabia reptiles in the sand deserts of southern Israel. In: Proceedings of the Symposium on the Fauna and Zoogeography of the Middle East, Mainz, 1985. Krupp F, Schnider W, Kinzelbach R (eds.), Beihefte zum TAVO A 28, 272-295 Werner YL, Sivan N (1992) Systematics and zoogeography of Cerastes (Ophidia: Viperidae) in the Levant: 2, Taxonomy, ecology, and zoogeography. The Snake 24: 34-49 Werner YL, Sivan N, Kushnir V, Motro U (1999) A statistical approach to variation in Cerastes (Ophidia: Viperidae), with the description of two endemic subspecies. Kaupia 8: 83-97 Werner YL, Verdier A, Roseman D, Sivan N (1991) Systemat- ics and zoogeography of Cerastes (Ophidia: Viperidae) in the Levant: 1, Distinguishing Arabian from African Cerastes cerastes. The Snake 23: 90-100 Received: 10.V.2010 Accepted: 29.VI.2010 OZFMK Bonn zoological Bulletin | Volume 57 Issue 2 | pp. 307-328 Bonn, November 2010 The taxonomic history of the Linnean genus Lacerta (Squamata: Sauria: Lacertidae) in the mirror of book-illustration Josef Friedrich Schmidtler Oberfohringer Str. 35, D-81925 Miinchen, josef@schmidtler.eu Abstract. The taxonomic history of the Linnean genus Lacerta illustrates the general taxonomic history in herpetology and can be visualized by the history of book illustration. There is a cohesive pattern in lumping Lacerta (Linnaeus, 1758; comprising lizards, crocodiles and salamanders; expanding to almost 100 species in Shaw, 1802) and splitting (Lauren- ti, 1768; comprising among others his new genus Seps, a part of Linnaeus’ Lacerta), since the creation of binominal nomen- clature by Linnaeus, and proceeding above all to the controversy of Boulenger and Méhely after 1900. These wavelike advances through the centuries are also characterized by a slow consolidation of the higher systematic categories (class- order-family-subfamily, etc.) and by a gradual reduction of the term Lacerta to almost the species level. This develop- ment ended now in an enormous generic and specific splitting within the family Lacertidae (Arnold et al. 2007), main- ly based upon mitochondrial DNA research. The remaining “true” Lacerta comprises at present only half a dozen species, all of them close relatives of the type species, Lacerta agilis. There is an historical interdepency between verbal descriptions and illustrations in the taxonomic advances of the genus Lacerta. The first illustrations of lizards (in the 16'® century) are in equal measure characterized by the lack of system- atic insight and the lack of technical options. Copper engravings (handcoloured) were used a little later. Since the end of the 18th century, new techniques accompanied and immensely facilitated a better recognisability of taxa: wood engrav- ings — lithographs — chromolithographs — photos — modern digital colour photographs. The better understanding of the diagnostic scale structures called for their schematic depiction, and a schema of the dorsal drawing pattern was estab- lished. Diagrams for identification keys and/or of the phylogenetic relationships have become an indispensable part of modern taxonomic work. On the contrary, the genetic revolution of the last 20 years caused a great loss in importance of morphological characters, whereas top-quality digitalized coloured photos have shifted their importance mainly to pop- ular publications on ecology, ethology, field herpetology and terraristic studies. Keywords. Genus Lacerta; history; interdepency text/illustrations. 1. INTRODUCTION: SOME COMMENTS ON THE ZOOLOGICAL TERM GENUS The history of the Linnean genus Lacerta is likewise a his- tory of the term “genus”. As Mayr et al. stated in 1953, the genus is a collective taxonomic unit consisting of a number of similar or related species. It is distinguished from all other higher categories by being recognized in the scientific name. The nomenclature proposed in Lin- naeus’ Systema Naturae (1758; animals) is binominal, con- sisting of two names, each with its own function. The func- tions which Linnaeus visualized for the components of the scientific name are diametrally opposite. The specific name signifies singularity and distinctness; the generic name calls attention to the existence of a group of simi- lar or related species — it relieves the memory (Mayr et al. 1953: 48). Bonn zoological Bulletin 57 (2): 307-328 Even before Linnaeus there was a recognition of the cat- egories genus and species. So, Plato definitely recognized two categories, the genus (“genos”’) and the species (“e1- dos”), and so did his pupil Aristotle. The naturalists of the pre-Linnean era were not consistent in the Latin names they gave to plants and animals. These names ranged all the way from uninominals (a generic name only), and bi- nominals (a generic and a simple trivial) to polynominals (a generic name with several trivial epithets). The reason for this confusion was that they tried to combine two dif- ferent functions in the name: naming (in the restricted sense of the word) and describing (Mayr et al. 1953: 202; see the legends in the images of Gessner and Aldrovandi Figs 2 and 3 hoc loco). ©ZFMK 308 Josef Friedrich Schmidtler An objective criterion for the generic rank does not exist equivalent to the biological species concept (“reproduc- tive isolation”) in species systematics (see Mayr 1984: 141, 219; Jahn 2004: 237, 397; Joger 1996) as a criteri- on. It is therefore impossible to give an objective defini- tion of the genus. So Mayr et al. (1953: 48) came to the following conclusion: “A genus is a systematic category including one species or a group of species of presumably common origin, which is separated from other similar units by a decided gap”. They suggest for practical rea- sons that the size of the gap be in inverse ratio to the size of the unit; the latter qualification should prevent the recognition of unjustified monotypical genera. The general view on the definition of the category genus has not changed much since then, contrary to the differ- ent species concepts (e.g. Joger 1996). Even Mayr et al. (1953) had attenuated their clause appearing so strict (“An objective criterion does not exist ....”; p. 48) when dis- cussing the presence of an “ecological niche” (p. 50) be- tween genera. Later on Dubois (1988) and recently Dubois & Bour (2010) have extensively discussed the demand of “hybridizability” as a criterion for the definition of gen- era and subgenera. Additionally, the genetic revolution in taxonomy since the 1990s has decidedly consolidated the phylogenetic trees. So Speybroeck et al. (2010), in the in- troduction to their recent species list of the European her- petofauna, come to the decision: “As a distinct genus, we tend to recognize monophyletic clades that are genetical- ly as divergent as other widely accepted genera in the same group. This is usually the approach employed by authors of scientific papers....”. As a conclusion one might assert, that it was molecular biology which gave rise to a still con- tinuing revolution in herpetological taxonomy, and — above all — to an enormous generic splitting, be it of the old Linnean genera Testudo, Rana, Coluber, or Lacerta, the latter being discussed here. 2. A BRIEF HISTORY OF THE LINNEAN GENUS LACERTA 2.1 The “lumper” Linnaeus (1758 / 1766) and his fol- lowers The history of the genus Lacerta reflects also a history of zoological terms and categories, which can be dealt here only with its basic intentions. The word “Lacerta” (or the male gender “Lacertus”’) is of Latin origin. One of its three meanings is the linguistically derived English term “lizard”. In this sense it is traceable in the Historia Nat- uralis of Plinius or in some works of the classic Latin po- ets Ovidius and Virgilius (Scheller, 1796). Since the era of renaissance this term was renewed by natural scientists in both the male and female gender (see Figs 1, 2, 4). Thus Bonn zoological Bulletin 57 (2): 307-328 the term Lacerta / Lacertus had a long history before Lin- naeus began to use it in the different editons of his “Sys- tema Naturae” since 1735. Linnaeus (or “Linné” after nobilitation), in his famous 10th edition of 1758, divided the class “Amphibia” into three orders: I. Reptiles, H. Serpentes, HI. Nantes. The “Rep- tiles” comprise the four genera Testudo, Draco, Lacerta, Rana; the Serpentes comprise the six genera Crotalus, Boa, Coluber, Anguis, Amphibaena, Caecilia. The Nantes comprise six genera, all of them being transferred later on into the class Pisces. The Linnean Rana, parts of Lacer- ta and Caecilia constitute the current class Amphibia whereas the other genera in Linné’s orders Reptiles and Serpentes are comprised in the present-day — polyphylet- ic — Reptilia. Linnaeus’ (1758/1766) large genus Lacerta is an aggregation of 43/49 species, e.g. comprising the cur- rent Lacertidae (type species of Linnaeus’ Lacerta is Lac- erta agilis by later designation in Fitzinger 1843: 20), many other Reptile orders (like the crocodylia) and fam- ilies, and even amphibians (e.g., salamanders; see Fig. 1). His genus Lacerta is encompassed by the diagnosis “Cor- pus, tetrapodum, caudatum, nudum” (body with four legs, caudate, “naked”; the latter characterization being com- pletely incomprehensible, since his genus Rana is also characterized to be “naked”!). It seems that Linnaeus did not misjudge completely the heterogenity of his genus Lacerta. He tried to resolve the problem by species groups, characterized by short diagnoses and different stars. So his Lacerta agilis is within a group characterized by “** Cau- da verticillata” (Tail round) and the group with the fire salamander, “Lacerta salamandra”’,, 1s characterized by aK Palmis tetradactylis; Corpore alepidoto nudo” (fore legs with four toes; body without scutes, naked). Nevertheless the newt “Lacerta vulgaris” (number 25; now: Lissotriton vulgaris) is grouped together with geck- os and skinks. Gmelin (1789) was formally a follower of Linnaeus, but he undermined his concept in the so called 13 edition of Linné’s “Systema Naturae” where he accumulated the number of Lacerta species up to 77. Gmelin introduced eleven species groups within Lacerta, characterized by short diagnoses and mostly (but not always!) naming them (nominative plural of a main species being included, like “Salamandrae” with five stars ( ), comprising the Linnean Lacerta salamandra, No. 47) — or his ““Ameivae s. Sepes” (“‘s.” = sive”; English: “or’”’) comprising the Lin- nean Lacerta agilis — or the ““Lacerti” (with nine stars) cov- ering current tailed amphibians and reptiles, except lac- ertids, like the newt “Lacerta vulgaris” (now: Lissotriton vulgaris which is therefore not a part of his “Salaman- drae”’!). It is clearly noticeable that Gmelin, following Lin- naeus, avoided dividing the genus Lacerta formally, un- like Laurenti (1768) had executed. The non-scientific rea- ©OZFMK Taxonomic history of the genus Lacerta sons may have been similar as described below in the dis- cussion on Shaw. Donndorf (1798) followed Gmelin (1789). He used the German terms “Geschlecht und Gattung” instead of “Gat- tung und Art” (genus and species; “Vorrede” p. 5). In his genus Lacerta Gmelin’s system with eleven species groups, characterized by eleven stars, is comprised; he added however 14 newer species (“neuere Gattungen”) within these species groups and nine species of undeter- mined species groups. Shaw’s (1802) General Zoology (vol. II, part II], Amphib- ia) is the last of the great encyclopaedias around 1800 which formally retains the generic name Lacerta in the broad Linnean sense. Its number of species has increased up to 86. Like the preceding encyclopaedias Shaw divid- ed Lacerta into nine “sections or sets” giving them Eng- lish names. He admitted however: “The above divisions Linnaeus (1758) Laurenti (1768) Wagler (1830) Dumeéril & Bibron (1839) Boulenger (1920) Arnold (1973) Arnold & al. (2007) o Pa ia") me) =) Oo Oo & 2 <= foe € <¢ Testudines Crocodilia Fig. 1. 309 neither are, nor can be, perfectly precise...” His “4. Lizards proper “ comprised also the current day Lacer- tidae, among them the ’Green lizard” “Lacerta agilis” tak- ing first place. Smith & David (1999: 12, 13), when dis- cussing the taxonomic situation then, drew the conclusion: “The rudimentary level of understanding of herpetologi- cal classification in Shaw’s time is admirably exemplified by his treatment of the Division Lacertae, containing on- ly two genera — Draco and Lacerta — that are extremely disparate in diversity. Nevertheless, Shaw was much more aware of the diversity and affinities of members of his genus Lacerta than is apparent in the assignment to one genus, inasmuch as he recognized nine distinct groups. To us it seems strange that such diversity was not reflected taxonomically when the relatively minor specialization of Draco received such emphasis. However, although Shaw boldly named new species or changed names, he reflect- ed the trepidation widely shared at that time among his colleagues in splitting Linnean genera. Change then, as other Squamata other Lacertini Gallotiinae Eremiadini Zootoca Podarcis Serpentes Anguidae Lacertidae Overview of the gradual reduction of the Linnean genus Lacerta (Laurenti’s genus Seps respectively) from selected mo- nographs: Linnaeus (1758) — Laurenti (1768) — Wagler (1830) — Dumeéril & Bibron (1839) — Boulenger (1920) — Arnold (1973) - Arnold et al. (2007: fig. 20); displayed upon a current phylogenetic tree (strongly simplified; from Dawkins, 2008, figs. p. 366, 422, 462; Arnold et al. 2007 figs.). The symbols Z or S make clear a quotation of “Lacerta” (or “Seps” by Laurenti) within a gi- ven current systematical unit; X (no species from this group being described then). Bonn zoological Bulletin 57 (2): 307-328 ©ZFMK 310 Josef Friedrich Schmidtler 362 Det Kohe von den Eydeken foll auch viel Krafft haben fiir etlicheRrancEheiten dev Augen. Lcbendig in Del geFocht/ follein gare Gefiche machen. Dahero Becherus: Gefnert Thierbuch Die Eydcy lebendigin Ocl man fochen thut/ Es macht cin weif Geficht/ ift vor die Rehte gut. rob SEL SSB- GOST SB SSS SOB BES 0F- U) BEST ES BES OB~ SEBS OR ESF 263 Von der grinen Eyoere. Lacertus viridis. Grtiner Heydor/Egochs/Sltachs/ oder Eder. : Fig. 2. “Lacertus viridis” from Gessner’s (1671) last edition, the so called “Frankfurter Ausgabe”. Wood cut.- The eye-catching bars across the tail indicate the whorls typical for a Lacertid tail. The text consists of medical and cosmetic advices, the latter in the form of a verse. The different insights of a painter and a mere engraver as well as the different qualities of a water colour com- pared with an engraving are shown by a comparison to an Aldrovandi image (Fig. 3). now, came slowly. Shaw worked in a surprisingly substan- tial intellectual milieu of peers who would look critical- ly at any change from established authority”. A compara- ble thoughtfulness seems to have been widespread in sc1- ence — at least then (compare the situation of N.M. Op- pel after his studies in Paris, being surrounded by “natu- ral philosophers” in Munich since 1811; Schmidtler 2009: 509; Figs 16, 17 hoc loco). The “Histoire Naturelle des Quadrupédes Ovipares et des Serpens” by Lacepede (1788/89) is a milestone in the his- tory of natural science. It highlights the beginning of the acceptance of Linnaeus’s binominal system also in France, then leading in natural science. Up to that time the well known scientific controversies of Linnaeus (1708-1779) and Buffon (1708-1788) had prevented to a large extent the application of Latin binominal names in the French zoological and botanical literature, especial- ly in the dozens of volumes of Buffon’s “Histoire Na- turelle” having appeared since 1750. The acceptance of Linnaeus’ binominal system by Lacepéde appears admit- tedly in a rather hesitant and concealed manner. It turns Bonn zoological Bulletin 57 (2): 307-328 up only in the gigantic Latin “Synopsis methodica Quadrupedum oviparorum” beside the French “Table méthodique des Quadrupedes ovipares” (see the elaborate description in David et al. 2002: Fig. 2). Here Lacepéde accepted two classes. His first class (“Quadrupedes ovipari caudati”) comprises two genera, 7estudo and Lacertus (“Corpus absque testa’). The latter, with 56 species, is di- vided into eight divisions (“divisio”) which are each de- scribed shortly, but not named. As Spix (1811: 342) point- ed out there is however a contradiction between La- cepéde’s (1788) zoological findings in the text and the con- struction in the “Table méthodique” when stating that the salamanders are nearly related with the frogs. He accept- ed two current lacertid species in his third division (Lac- ertus cinereus and Lacertus viridis; both highly collective groupings, comprising among others the current genera Lacerta / Timon and Podarcis / Zootoca respectively; see also Schmidtler & Béhme, in prep.). The six species of salamanders are, contrary to Linnaeus and Gmelin, con- centrated in one division (“VIII. Divisio”). It may be not- ed that the name with the male gender “Lacertus” La- cepéde, 1788 is regarded to be an unjustified emendation ©ZFMK Taxonomic history of the genus Lacerta 311 — Fig. 3. Podarcis siculus ssp. From the collection of Aldrovan- di’s natural history images (16'" century), see also Ceregato & Alessandrini (2007: fig. 462 upper part) and Delfino & Cerega- to (2008). Water colour (Tempera). — The shapes and colours of lizards are excellently painted but the head shields are scarcely indicated. The naming of tail anomalies (especially Lacerta “‘bi- ceps”!!!) reveals the lack of a species concept and the lack of a preset nomenclatural terminology. See chapter 3.2. of Lacerta Linnaeus, 1758 (see David et al. 2002: 24). La- cepede’s volume | and volume 2 (on the snakes) were re- jected in general as a non-binominal work. This opinion remained heavily controversial (see David et al. 2002: 22; Dubois & Bour 2010). Anyway, one year later many species were adopted by Bonnaterre (1789) who has there- fore become the correct author of many of Lacepéde’s names not being available. On Bonnaterre’s Lacerta see more in chapter 2.2. As obvious from the title, Bechstein’s encyclopedia (1800-1802), “Herrn De la Cepede’s Naturgeschichte der Amphibien” is first of all a translation from Lacepéde’s (1788 / 89) Histoire Naturelle, but comprising many ad- ditions. Bechstein used German terms. In his “Methodis- che Ubersicht der eyerlegenden vierfiiBigen Thiere” he translated Lacepede’s system with the terms Classe, Gat- tung (genus) and Art (species), inserting the term “Fam- ilie”, apparently in the sense of a species group below his “Gattung” in some genera, like the “Eidechsen” (lizards). He neglected Linnaeus’ Latin binominal teminology to a large extent. His terminological system concerning the cat- egory “Art” (species) is inconsistent and confusing. In his “Zweyte Gattung, dritte Familie” (vol. II, like in Lacepéede 1788) some current Lacertidae (“L. cinereus, die graue Ei- dechse” and “L. viridis, die griine Eidechse’”’) are com- prised. He gave an excellent picture of the “Graue Ei- dechse / L. cinereus” (vol. I, Taf. 1; depicting the male and the female of a present-day Lacerta agilis), and demonstrating thereby that this important taxon was then differently understood in different European countries (see Fig. 4. Lacerta agilis (male). Left figure: Draft of the right figure; drawer Résel von Rosenhof (Cod. Icon. 48, Bayerische Staats- bibliothek Miinchen; before 1758). Water colour. — Right figure: Frontispiece in Résel v. Rosenhof (1758). Hand coloured engra- ving.- The changes in scientific insight by the famous drawer and natural scientist Résel mirror as well his personal “metamor- phosis” as the changes in general views in differentiating a “salamander” (left: with its nude skin!) from a lizard (right: with its li- felike scaly skin and pileus scutes; but: occipital and interparietal scutes are still lacking!). Both, salamanders and lizards, became in the same year parts of the Linnean genus Lacerta. Bonn zoological Bulletin 57 (2): 307-328 ©ZFMK 312 Josef Friedrich Schmidtler also Schmidtler 2004; Schmidtler & Bohme in prep.). In his “Anhang” (additions; vol. I, 297-325) however, Bech- stein on the one hand accepted the modern binominal ter- minology of Laurenti (1768) and Schneider (1799; e.g. Stellio phylluros, p. 307) or used Gmelin’s (1789: 1060) Latin species group terms, (p. 311; like Stelliones = “Spiegeleidechsen”). 2.2. Early generic splitting after Linnaeus Laurenti (1768) was the first to subdivide the Linnean gen- era of amphibians and reptiles (Testudines excluded) in a comprehensive work, especially Linnaeus’ large genera Lacerta and Coluber. Laurenti totally suppressed the name Lacerta, but established instead of 11 new genera within his order II ““Gradientia” (see Kuzmin 2005: 246), among them “Seps’’ comprising also the current species of Cen- tral European Lacerta and Podarcis and some of their syn- onyms (see chapter 2.4; see Fig. 1). After Steyneger’s (1936) type species designation (Seps caerulescens = La- certa agilis; see also Dubois 2010; and Fig. 6 hoc loco) Laurenti’s Seps became a junior synonym of Lacerta. Seps Laurenti comprised after all only four current families, all within the Squamata (Lacertidae, Scincidae, Teiidae, P1668. } il Wi) i| | I tll AN AAW Fig. 5. Gymnophthalmidae) and appears therefore much more re- stricted than the Linnean Lacerta. Laurenti’s splitting had still an earlier forerunner in Garsault’s (1764) long forgotten and just rediscovered work “Les Figures des Plantes et Animaux” here concern- ing in particular the French herpetofauna around Paris in ten plates (see Welter-Schultes et al. 2008, 2009; Dubois & Bour 2010; Fig. 5 hoc loco). Garsault (1764) used the species names Lacertus terrestris (now: Podarcis muralis (Laurenti); nomen conservandum), Lacertus viridis (now: Lacerta bilineata Daudin; nomen conservandum), the genus names Scincus, Salamandra (with the French name “salamandre” behind; depicting Salamandra salamandra terrestris Bonnaterre). A certain systematic unstableness is however unmistakable when depicting the crested newt (today 7riturus cristatus (Laurenti), nomen conservan- dum) under the Latin nomen Lacertus aquatilis, but simul- taneously under its French name “Salamandre d’eau”. Valmont de Bomare in the second issue of his “Diction- naire d’ Histoire Naturelle” (1767/68) added for the first time Latin names to the French names. There appear like- wise considerable systematic inconsequences: On the one hand, under the key word and generic name “Lézard / Lac- | | | ' b j | =) jidse id j j 1} | | EY Mi i TORING “Lacertus terrestris” (= Podarcis muralis; western subspecies) from Garsault (1764), a forerunner of Laurenti (1768), having been rediscovered in the last years (see Dubois & Bour 2010). Copper engraving. — The pattern of the upper head scutel- lation (right figure) is not yet perfect; the frontal and postfrontals are not executed within the central part of the pileus. Bonn zoological Bulletin 57 (2): 307-328 ©ZFMK Taxonomic history of the genus Lacerta 313 Tas: I. , Fig. 6. Seps caerulescens (= Lacerta agilis; Fig. ID), Seps muralis (= Podarcis muralis; Fig. 1V), Seps argus = a juvenile La- certa agilis; Fig. V). From Laurenti (1768: Tab. I, upper part). Copper engraving. — Laurenti’s (1768: Tab.I, fig.II]) Seps caerules- cens has accomplished perfection for the first time in the history of a lacertid engraving: The arrangement and shape of all of the pileus scutes are accurate. In equal measure the dorsal pattern is very representative for the species. This figure is all the more out- standing, as Laurenti himself was obviously not yet aware of the enormous impact of head scutellation in species recognition. So it was the exactness of the drawer and the engraver who were solely responsible for the quality of the figure. Developments like these demonstrate the prospective relevance of naturalistic figures in book illustration for the scientific progress in reptile syste- matics about 1800. Otherwise, the quality of the smaller figures IV and V is considerably lower and does scarcely contribute to species recognition. ertus” (Vol. III, p. 548, 1768), all the “amphibian species” in Linnaeus’ (1758) sense are understood. On the other hand, under the keyword Salamandre / Salamandra, Val- mont de Bomare (vol. V, p. 441, 1767) described only “re- al” salamanders, and explicitely the two species Salaman- dra terrestris (currently: Salamandra salamandra ter- restris Bonnaterre) and Salamandra aquatica (apparent- ly a collective species comprising some current species of water newts, especially 7riturus cristatus). This diction- ary was however suppressed by the ICZN (Anonymus 1925; Dubois & Bour 2010), its nomenclature not being always binominal for species. Although Bonnaterre (1789; see also 2.1), often misun- derstood as a mere copyist of Lacepede, took over many details from Lacepede (1788) in his “Tableau Ency- clopédique”’, he did not follow him (nor indirectly Lin- naeus) in his generic lumping. On the contrary, he wide- ly accepted the generic splitting by Laurenti. His first class (“Reptilia ecaudata”) comprised the three genera of Lau- renti: Rana, Hyla, Bufo, whereas his second class (Rep- tilia caudata) with seven genera approached Laurenti’s sec- ond order (less genera indeed), excluding the snakes as well, but comprising the turtles (Zestudo). Bonnaterre’s Lacerta comprised 52 species, among them still some of Laurenti’s new “lizard” genera, like Basiliscus, Iguana, Ameiva, Stellio. Bonnaterre suppressed Laurenti’s gener- Bonn zoological Bulletin 57 (2): 307-328 ic name Seps and the current Lacertidae are comprised in his genus Lacerta. He also doubted the validity of some of Laurenti’s new species (e.g. Seps caerulescens, now Lacerta agilis). Latreille in Sonnini & Latreille (1801) in gross terms ac- cepted the generic systematics of Laurenti (1768). Some of Laurenti’s species of the new genus Seps were includ- ed in the “Ie genre Lézard, Lacerta’. Latreille anticipat- ed many of Daudin’s (1801-1803; see chapter 3.3) de- scriptions and took the opportunity to thank him for his communications (1801, vol. 1, p. 215; “M. Daudin...a eu Vamitié de me communiquer, par extrait, les descriptions qu’il a faites de plusieurs reptiles de la famille des lézards... Il me sera doux, en le citant, de lui payer a la fois le tribute de mon estime et celui de l’amitié”’). Harp- er (1940) named this procedure a certain sort of piracy. In early regional faunas, which do not cover the whole family of lacertids, the acceptance of Linnaeus’ lumping or Laurenti’s splitting was heterogenous. Being one of the first authors, Schrank (1784 and 1798) completely adopt- ed Laurenti’s genera (e.g. “Salamandra atra” Laurent, “Seps viridis” Laurenti), whereas Razoumowsky (1789) or Wolf in Sturm (1799, 1802, 1805) were using Linnaeus’ terminology system (e.g. “Lacerta agilis” Linnaeus or Lacerta paradoxa s. helvetica” (n.sp.; now the newt Lis- ©ZFMK 314 Josef Friedrich Schmidtler THE GREEN LIZARD. Tux colours of this species are subject to variety, becoming pale at certain seasons of the year, and more particularly after the death of the animal. The upper parts of the body are of a beautiful green, more or less variegated with yellow, grey, brown, and even sometimes with red. In warm regions it grows to a larger size than in more temperate countries, being sometimes found thirty inches in length. The inhabitants of Africa eat the flesh of this animal. Fig. 7. “Green lizard” (= Lacerta viridis); probably the first lizard in the renewed wood cut technique by Bewick (1809 — 1816; “wood engraving”; cf. Dance, 1989, Schmidtler, 2007). As usual then, engravings of the “abhorrent” reptiles (so Linna- eus 1758:194) were significantly of a lower quality than the birds or mammals (cf. the images in Bewick 1791). Nevertheless this green lizard is recognizable here. It was a great advantage of this printing technique that the images could be printed together with the text upon the same page (unlike copper engravings or litho- graphs - these upon separate plates). So, later on, wood engra- vings proved to be adequate for the popular small English “chap books”, or the German “Naturgeschichten”. This kind of letter press was also often used for schematic figures in a text page. sotriton helveticus) in the former; “Lacerta atra” (Lau- renti), and “Lacerta agilis” Linnaeus in the latter. In con- trast Koch in Sturm (1828) made use of Laurenti’s gener- ic names (“‘Seps stellatus” Schrank, “Triton alpestris” Lau- renti) in the same “Deutschlands Fauna”. 2.3. An enormous increase of knowledge since 1800 Since about 1800 the knowledge in natural science in- creased immensely and many new species were described. Laurenti’s (1768) system of splitting the Linnean genera began to win recognition. Nonetheless, Laurenti’s total re- placement of the generic name Lacerta, e.g. by Seps, was usually not accepted. Bonn zoological Bulletin 57 (2): 307-328 Some months after the issue of Sonnini & Latreille’s (1801) encyclopaedia Daudin started his “Histoire Na- turelle des Reptiles” (“An X” = 1801; see Harper 1940 for the exact data). His “Second ordre. Les Reptiles Sauriens” approximately conforms with the genus Lacer- ta lumped by Linnaeus and Lacepéde (1789), but the sala- manders were transferred to his fourth order “Les Rep- tiles Batraciens” comprising also the frogs. His genus Lac- erta is one of 16 genera within these “sauriens”, most of them being current lacertids except the Ameivas. His generic systematics resembles Laurenti’s (1768) splitting system in general. One of the decisive differences was its essential feature in the formal persistence of a large genus Lacerta, whereas Laurenti’s generic name Seps was made use of for only some two- or four-legged saurians. Daudin’s greatest progress may be the redivision of his newly split genus Lacerta: It comprised 32 species sub- divided into seven unnamed “sections”. These sections presage the present lacertid genera in some very ambigu- ous outlines. For example, his second section “Lézards verds” contains Lacerta ocellata (now Timon lepidus) and Lacerta viridis (now: within Lacerta s. str.) as well, where- as his fourth section “Lézards tachetés” contains “Lacer- ta lepida”’ (a young Timon lepidus) and his new Lacerta maculata (a very cryptic name in some respects). Espe- cially with Daudin the level of knowledge began to in- crease immensely. This growth did not only include fur- ther generic taxa but also an inflation of species names by naming “real” new species, also individual or local variations, both sexes or juveniles. Replacement names took the upperhand more and more. The names for the one current species Lacerta agilis (three Seps species names in Laurenti (1768); see Schmidtler 2004 and Kuzmin 2005) were augmented by Daudin to three more names (male, female, young) in his fifth section “Lézards gris”. This fifth section comprised also his “lézard gris” with the Latin name “Lacerta agilis” (currently Podarcis muralis). Until very recently Oppel (1811, see Fig. 16) was held as the author of the family Lacertidae (“Lacertini”) (now: Lacertidae Batsch, 1788; cf. e.g. Speybroeck & al. 2010). Based upon Dumeéril (1806) he moved ahead the system- atics in the higher categories and made them clear by trees as a forerunner of evolutionary ideas (Schmidtler 2009). Merrem (1820) was the first to publish a schematic im- age and a terminology of the lacertid head scutellation (chapter 3, Fig.12). His genus Lacerta comprised 27 species, some of them being new. His systematics is based in general upon Daudin (1801-1803), Oppel (1811) and Cuvier (1819). He introduced some new terms and taxa in the higher categories. So his genus Lacerta is part of the “stirpes” A. Ascalabotae, the “tribus” 1. Gradientia, the III. order Squamata and the class 1. Pholidota (the 2. class is named Batrachia). ©ZFMK Taxonomic history of the genus Lacerta 315 P,Z.S 1908). Pl. LXVI. Fig. 8. Lacerta chlorogaster (= Darevskia chlorogaster) from Boulenger (1909: Pl. LVI). Below one chromolithograph (i.e. an image printed successively with differently coloured lithograph plates), and above two (pen-) lithographs (scutellation of pileus and surroundings of the anal region). The highly informative combination of naturalistic and schematic figures upon one lithogra- phic plate turned up at first in the middle of the 19t" century. The “Neue Classification der Reptilien nach ihren natiir- lichen Verwandtschaften” was Fitzinger’s (1826) first im- portant work (see Mertens, 1973). His “XI. Familia. Lac- ertoidea. Lacertoiden” comprises three genera, among them Lacerta with 17 species. It was apparently the first time in a systematic listing that neither this family nor the genus Lacerta comprehended any taxa now being ranked outside the present-day Lacertidae. It was the age of the great systematic monographs and shortly afterwards Wagler (1830) published his “Natiirlich- es System der Amphibien mit vorausgehender Classifica- tion der Saugethiere und Végel”. Wagler’s monograph is especially distinguished by comprehensive and progres- sive morphological and anatomical descriptions and con- siderations (pp. 211-344). His genus Lacerta only com- prised lizards belonging to the current genera Lacerta (s.str.) and Timon. His “Familia III. L. autarchoglossae” comprehended the Linnean taxa Lacerta and Tachydro- mus, as well as the new lacertid genera Zootoca, Podar- cis, Aspistis, Psammuros (the latter two are still synonyms of Psammodromus Fitzinger), apart from some genera be- Bonn zoological Bulletin 57 (2): 307-328 longing to other current families. Zootoca and Podarcis were regarded mostly as synonyms subsequently, but were revalidated more than 150 years later. All in all Wagler’s systematics of the genus Lacerta appears rather modern (Fig. 1). The “Histoire Naturelle des Reptiles” in eight large vol- umes by Duméril & Bibron (1834-1854) represents a new kind of herpetological monograph, compared with Daudin’s (1801-1803) work. Especially because of the im- mense growth of knowledge the different species chap- ters increased, comprising different sub-chapters (e.g. in Lacerta vivipara: “caractéres, synonymie, déscription” with “patrie et moeurs” in seven pages). His species chap- ters on Lacerta were based on many detailed new works, including also relatively new disciplines (e.g. reproduc- tion biology) by Milne Edwards (1829), Duges (1835), Cocteau (1835) and Tschudi (1837). Dumeril & Bibron (1839) were lumpers, compared with Wagler (1830). Their genus Lacerta comprised 16 species (some of them new), subdivided into three species groups. Their genus Lacer- ta is currently ranked in 14 genera, some of them having ©ZFMK 316 Josef Friedrich Schmidtler a. Pact AZ » Be oe | eee >. ry. Qa ry la. ie i Fig. 9. Ventral sides of “Lacerta muralis vars. lilfordi / serpa / brueggemanni”’ (i.e. now three different species of Podarcis), from Boulenger (1905: pl. XXII). — Hand coloured photos (The combination of hand colouring and photographs was very unusu- al then in natural science). The spotting and the colouration are of systematic importance in these “varieties”. The diagnostic fea- tures, especially the sutures of the shields, are not presentable simultaneously in the same figure (See Fig. 8!). — In 1853 the new technique of photography had been received with enthusiasm after a publication on reptiles (varans and a crocodile) and other ani- mals (“Even the best painter would not have the patience and ability to make visible all the details and structures. ..”; see Niekisch 2010). been described before Duméril & Bibron. The present-day Lacertidae corresponds to Duméril & Bibron’s subfami- ly “Coelodontes” comprising nine genera. Duméril & Bibron (1839: 1-19; 181-189) published a substantial his- torical outline of their family “Lacertiens ou Autosaures” and their genus Lacerta, respectively. It is worth mentioning the chapters on “Erpétologie” or “Lézards” in different French natural science dictionaries, which are now more or less forgotten. They mirror im- posingly the general advances in herpetology between 1800 and 1850 and in Lacerta in particular: See Bosc d’ Antic (1817: 521-528), Cloquet (1819/1823), Bory de Saint- Vincent (1826/1828), Cocteau (1835) and Meunier in Guérin (1836). Contrary to Duméril & Bibron (1839), Fitzinger (1843) proved to be a splitter. Within his class Reptilia he includ- ed the categories “Series”, “Ordo”, “Tribus” and ‘“‘Famil- Bonn zoological Bulletin 57 (2): 307-328 ia”. The present-day Lacertidae were divided into three families: Lacertae, Tachyscelides and Eremiae. His first family Lacertae comprised four genera (Scelarcis, Podar- cis, Chrysolamprus, Lacerta), most of them being subdi- vided into subgenera. As Mertens (1973: V) stated, Fitzinger’s (1843) work is of tremendous significance for the study of amphibians and reptiles, not so much of the nearly one hundred new generic and subgeneric names proposed, but because he always cited generic type species. In the case of Lacerta this tremendous signifi- cance is manifested in Fitzinger’s (1843: 20) determina- tion of “Lacerta agilis. Linné” as the type species of Lac- erta. The excellent coloured engraving by Wolf (1799) may have been here the decisive motive. Like Kaup (1836; see also Fig. 17), Fitzinger (1843: 12) was also a follow- er of the so called, unusual “Naturphilosophie” (natural philosophy), then distributed above all among German speaking natural scientists. ©ZFMK Die Griinen vom Kaiserstuhl e Kalserstuhl ist Y cults irmstes Weinbaugeblet und vul kanischen Ursprungs. Die dortigen LéBberge sildlicher Lagen erwirmen sich im Hoch: sommer bis auf 40 °C und darilber. Aber Gas macht ein vorziigliches Welnbauge- Diet noch nicht aus — erst die Durch schnittstemperaturen sind es, die Spitzen: weine, aber auch seltene Tiere und Pflan zen gedeihen lassen. Unsere Smaragdeidechsen Klimatisch bevorzugte Inmitten kihlerer Gegenden — der Kalserstuhl liegt zwi Taxonomic history of the genus Lacerta 317 Text und Fotos von Walther Rohdich Norden geschoben: flr Orchideen, In Sekten und Reptilien. Damit kommen im Kalserstuhl Arten vor, die wir erst wieder, und da sehr zahlreich, im Mittelmeer raum antreffen. Dem uns hier am mels: ten interessierenden Tier wollen wir uns Ugste Eldechse, die (Lacerta dilineata), Si Deutschland, aber Schwesterart Lace! einigen Stellen inselartig vor. Die Sma ragdeidechse ist die Wirmebedtirftgste unserer Eldechsen. AuGer am Kaiserstuhl Ein Maikifer ist cin fetter Happen lebt sie noch In der und im Donaugel 4) sowie an} fa), Das Vorkommen Mark Brandenburg ten Jahrzehn: bau der Welnber hen. Der Bestand Kurz elnig zu lhrer Biologie: Die Smaragdeldechse ist dle gréfte Eldechse Mitteleuropas, mit einer Linge von 40 cm. Sle Ist durchgehend leuchtend grin gefdrbt, mit ve; enen Anordnungen dunkler h nungselemente, wie Punk: far dieses prachtyolle Mannchen im schen den ruppigen Gebirgen Schwarz Hochra eid. wald und Vogesen — sind meistens Oasen fiir ungewéhniiche Tiere und Pflanzen. So auch hier: Hauptlebensriume, die weiter siidlich lie gen, haben Aus MWufer nach te und Flecken. Die Mannchen sind in Fig. 10. Male of Lacerta bilineata from the journal “Reptilia” (Nr. 20, 2004: 68): A perfect layout and a perfect digital colour photo have been combined here. The beautiful and viewable pho- tos serve firstly for keepers (demonstrating feeding habits, etc.) and field herpetologists (identification in the field / shop) in po- pular articles. In the light of modern genetic methods the images have lost their traditional predominance in underlining systema- tic descriptions. 2.4 The era (ca. 1880-1920) of George Albert Boulenger The end of the 19th century was initially characterised by new questions and topics as is displayed by Eimer’s (1881) indication of «darwinism» in the caption of his article. There infrageneric and infraspecific, geographical re- searches came to the fore. I should like to emphasize the basic advances, such as his formation of terminologies in the dorsal pattern (Fig. 14). We may remember here his long-standing controversy with Bedriaga concerning the origin of colouration in insular lizards (see Muller 1994). In this Darwinian sense Bedriaga (1886) tried to explain the phylogenetic relations and origins of lacertid taxa (the genus Lacerta with the five subgenera Lacerta, Algy- roides, Tropidosaura, Zerzumia, Bettaia) by detailed dis- cussions. His subgenus Lacerta, however, contained still Bonn zoological Bulletin 57 (2): 307-328 species of all current subfamilies and tribes. With respect to book illustration (see chapter 3) it may be regretted that his descriptions were corroborated by a single lithograph- ic plate only. The reasons here — as ever — may have been economical ones. Simultaneously the time of Boulenger’s great comprehen- sive catalogues in herpetology commenced. In the intro- duction of his Catalogue of the Lizards (three volumes) Boulenger (1887) displayed the immense increase in the numbers of lacertid species known and characterized: Dumeéril & Bibron (1839), 43 species — Gray (1845), 57 species — Boulenger (1887), 97 species. Boulenger’s Lac- ertidae comprised 17 genera and his genus Lacerta com- prised 21 species, among them species of the whole Eurasian and African range, i.e. species within the current subfamilies Gallotiinae and Lacertinae (some species of the current tribe Eremiadini not excluded; see Figs 1 and 19). The chapter «11 Lacerta muralis» (1887: 28) with many «varieties» underlines his very typical species con- cept. Boulenger’s vol. II contains a set of excellent lith- ographs, among them the new species L. parva (now Parvilacerta) and L. yayakari (now Omanosaura). Meéhely (1909) carried out intensive studies on morphol- ogy and osteology of European and Caucasian lacertids. Aside from the further development of the terminology of scutellation, osteology (Fig. 13), and pattern of muralis- like lacertids, he described the genus Apathya and the «I. Gruppe: Archaeolacertae» of his genus Lacerta, compris- ing species of the current genera Anatololacerta, Phoeni- colacerta, Hellenolacerta, Dinarolacerta, Iberolacerta and Darevskia. His victorious species concept («species» in- stead of Boulenger’s L. muralis-varieties) displays his fa- mous controversy with Boulenger. Schreiber (1912) adopted the view of Méhely and his species concept within the European-Caucasian lacertids. Within the current genus Podarcis he accepted as the first in a large monograph not less than eight species, most of them, especially Lacerta muralis and Lacerta serpa (= Po- darcis siculus), comprising many varieties and subvari- eties. Boulenger (1920 / 1921), covered the lacertids in their whole Eurasian and African range. Irrespective of the ac- ceptance of six «sections» within Lacerta he insisted up- on his system of the one muralis-like species, following his catalogue (Boulenger, 1887) and later papers (1905 and 1913 especially; see Fig. 19). His Lacerta muralis (belong- ing to his subgenus Podarcis Wagler) covers not less than 31 (!) «varieties». Most of them are presently species or subspecies or invalid forms within the genus Podarcis, but there are also taxa of the current genera Archaeolacerta, Iberolacerta and Darevskia to be found. It seems now that ©OZFMK 318 Tab. IZ. iS Tee 87 At Ait i Josef Friedrich Schmidtler TABOR es: AMPHIBIA nonmilla fiftens: 1, AMPHISBANA 87. annulis circularibus trur: cum I—7o 3 caudamque 1—15 cins gentibus. a, Caput. b. Anus, 2. COLUBER 89 feuta abdomen tegentia i—8o; fquame caudam fubtus tegen: tes I—I7. a. Caput. b. Anus: c. Apex caude. 9. ANGUIS 88. fquame abdomen tegentes 1-1203 > 1 ? {quame caudam tegentes 1—17. a. Caput. b. Anus. 4. CROTALOPHORUS 9%. feuta abdominis I—903 feuta caude1—1 33 crepitaculi articuli I—5; c. Crepitaculum. a. Caput. b. Anus. z, DRACO 92. pedes quatuor; cauda; Ale due cum radiis cartilagineis alarum. aby Qs Ps Fig. 11. Some reptiles and their scale countings depicted by Linnaeus (1756: Tab. III; 94 edition. Copper engraving). The scale countings refer mostly to the ventrals and subcaudals in snakes and were given especially in the text on the genus Coluber. In the diagnoses of the genus Lacerta no scutellation features were used then. This is apparently the first attempt of a schematic delinea- tion and description of body shields in herpetology. The same features were used also, without depictions, in the text of the 10t and 12th editions (1758 / 1766). his system was a relatively superficial morphological one, because he accepted also some (morphologically) conspic- uously different species besides his «L. muralis», like Lac- erta taurica (now within Podarcis), or Lacerta chloro- gaster (now within Darevskia). This was one of Boulenger’s rare mistakes in which, soon later, the her- petologists of this time did not follow his exceptional au- thority. Mertens & Miller (1928) adopted Boulenger’s (1920) Eu- ropean subgenera (Archaeolacerta, Podarcis, Zootoca, Lacerta), but they did not diverge in substance from the species concept of Méhely (1909) and Schreiber (1912). They were the first to accept geographical subspecies (see Wermuth in B6hme 1981), 1.e. a trinominal nomenclature in European herpetology (e.g. «Lacerta agilis exigua Eich- wald»). In addition they carried out some changes being nomenclaturally necessary (e.g. Lacerta lepida Daudin, 1802 - instead of the preoccupied Lacerta ocellata Daudin, 1802). Bonn zoological Bulletin 57 (2): 307-328 In Mertens & Wermuth (1960) there are considered many new discoveries of the European herpetofauna (especial- ly new descriptions of many lizard subspecies of «Lacer- ta» sicula, L. erhardii, L. melisellensis, L. lilfordi, etc., from Mediterranean islands, by Cyrén, Miller, Wettstein, Radovanovic, Eisentraut, Buchholz, in various papers each, since the second list of Mertens & Miiller (1940). Nevertheless this «Dritte Liste» characterizes the relative- ly stable generic and specific systematics and nomencla- ture in lizards between 1940 and 1990. 2.5 Towards a final breaking up of the genus Lacerta by new methods and techniques The basic works of Arnold (1973, 1989) and Bohme (1971) broke new ground in the systematics of the Lac- ertidae. New techniques (genital-morphological, karyolog- ical, electrophoretical, albumin-immunological, genetic features) and modern univariate and multivariate statis- tics were executed. ©ZFMK Taxonomic history of the genus Lacerta 319 Fig. 12. Schema of the pileus scutellation in Merrem (1820: p. XII-XII and fig. p. 191 upper part). (Pen-) lithography. — The decisive step ahead in the schematic depiction of lizards was made by Merrem (1820). Based upon his similar system in sna- kes (Merrem 1790 and 1820), he gave names to the pileus shields of an adult Lacerta ocellata (now: Timon lepidus; see his pages XI and XII) and depicted their abbreviations in this figure. The description covered seven types of scutes with the letters A (Wir- belschilder — Scuta vertebralia), B (Hinterhauptschilder — Scu- ta occipitalia), C (Augenbrauenschilder — Scuta superciliaria), E (Stirnschilder — Scuta frontalia posterioria), F (Schnautzen- schilder — Scuta frontalia anterioria), G (Rtisselschild — Scutum rostrale, L (Nasenlécherschilder — Scuta nasalia). This system was later on differentiated and improved by Milne Edwards (1829: pls. 5—8) who depicted and described also the shields of the lower sides of head, body and limbs. The concept of Mer- rem (1820) and Milne Edwards (1829) remain valid today. The first results of Arnold’s (1973, 1989; see Fig.1) elab- orate researches, based mainly on morphology were the revalidation of the old Waglerian genus Podarcis and of Gallotia Boulenger, 1916 (then a subgenus) besides two very preliminary groups, named “Lacerta groups [ and II’. The taxonomic tentativeness at that time found its way in- to the comprehensive “Handbuch der Reptilien und Am- phibien Europas” (Bohme in BOhme 1981, Bohme, 1981). Mayer & Bischoff (1996) (re-) established further sepa- rations from the so far comprehensive genus Lacerta (Zootoca, Omanosaura, Timon, Teira). They visualized a phylogenetic tree of the Lacertidae from the relationships of their serum albumins. Bonn zoological Bulletin 57 (2): 307-328 The numerous and very detailed morphological works of Arribas (1997/1999) resulted in the splitting off of the mainly SW-European genus /berolacerta Arribas 1997, and above all of the mainly Caucasian genus Darevskia Atribas, 1997 from Lacerta. Thereby also a very old con- troversy (especially of Méhely and Boulenger; see chap- ter 2.4) on the muralis-like “Archaeolacertae” could be fin- ished. The name Darevskia was given in honour of the great Russian herpetologist IS. Darevsky (1925-2009) who had detected parthenogenesis in these lizards, and therewith in reptiles (see Darevsky 1967; Schmidtler 2010). Beginning with the comprehensive work of Harris et al. (1998) new genetic methods have also been adopted in the systematics of the lacertids and they have caused here, like everywhere in systematics, a revolutionary situation. DNA sequences from parts of the 12S, 16S and cytocrome b mi- tochondrial genes, together with morphological informa- tion, were used to estimate the relationships within the family. This work was continued by Arnold et al. (2007; Fig. 20 hoc loco). DNA sequences indicated that the Lac- Fig. 13. Schema of skull bones (“Lacerta horvathi’’= Ibero- lacerta horvathi) in Méhely (1909: Taf. X, upper part) on the basis of Siebenrock (1894). — In the middle of the 19" century important osteological investigations also were executed in la- certids. They allowed taxonomic research in the higher catego- ries but also within lacertid genera and species, after a reasona- ble schematization in osteology, above all in skull terminology, had been found. ©ZFMK 320 Josef Friedrich Schmidtler 7887. OTD oa LT N erie wy iy iv x iy RTE “ ae Rs WE ite n\ Sr a Laf, XU. rs Nema io Ce 52a, y tee an Ven a F te - Fig. 14. Schema of the dorsal pattern in some Podarcis by Eimer (1881: Taf. XII). Lithograph. It was the research since the middle of the 18 century which revealed the crucial importance of the dorsal pattern especially in the specific and infraspecific taxono- my of the current genus Podarcis. Eimer (1881: Taf. XIII) named the different longitudinal zones (“I bis VI erste bis sechste Zo- ne”) which usually exhibit a system of narrow light longitudinal streaks (nrs. I and IH, ““Grenzlinien”) and dark bands (nrs. II,” in- neres / 4uBeres Band’). Méhely (1909: Fig. 1) eased his terminology and gave it the presently valid content; the seven light stre- aks and dark bands were named after their initial points at the pileus shields (like “Supraciliarstreifen” and “Occipitalband”); see also Schreiber (1912: Fig. 68; p. 333-335) and Mertens (1915: Fig. 3). ertidae contain two subfamilies, Gallotiinae and Lacerti- nae, the latter comprising two monophyletic tribes, the Eremiadini of Africa and arid southwestern and central Asia, and the Lacertini of Europe, northwestern Africa and southwestern and eastern Asia. Relationships within the 108 species of Lacertini were explored using mtDNA for 59 nominal species. The morphology of the tribe was re- viewed and also used to assess relationships. The Lacer- tini were assigned to 19 monophyletic units of 1 to 27 species. There were described five new Lacertini-genera out of the old collective genus Lacerta: Dalmatolacerta, Dinarolacerta, Hellenolacerta, Iranolacerta, Phoenicolac- erta (see Fig. 20 for a complete listing of current genera). The new generic concept does not include subgenera ex- cept in [berolacerta (Pyrenesaura Arribas, 1997). The genus Lacerta is presently restricted to eight species, the majority of them being polytypic: Lacerta agilis Linnaeus, 1758 (type species), L. bilineata Daudin, 1802, L. media Lantz & Cyrén, 1920, L. pamphylica Schmidtler, 1975, L. schreiberi Bedriaga, 1878, L. strigata Eichwald, 1831, L. trilineata Bedriaga, 1886, L. viridis Laurenti, 1768. Bonn zoological Bulletin 57 (2): 307-328 Thus the genus Lacerta appears to have finished its re- duction through the centuries (Fig. 1) and to have stabi- lized at a level a little above the species level (according to the biological species concept). It seems however, that the species systematics of the eight species of Lacerta has not yet drawn to a close. 3. THE INTERACTION OF VERBAL DESCRIP- TIONS AND ILLUSTRATIONS IN LACERTID TAX- ONOMY 3.1 General notes Zoological publishing in a modern sense, and together with it, zoological book illustration, started at the end of the middle ages, at the beginning of the renaissance era in the 16' century (Nissen 1978: 113). They included above all Belon, Gessner, Rondelet and Aldrovandi — all of them physicians — who did no more than see their cru- cial challenge in lining up reports and opinions of ancient OZFMK Taxonomic history of the genus Lacerta 321 De Animalibus in geuere. 83 Aninalinne Tabula generalis. Animalia fune vel { Sanguinea, eaque vel Pulmone refpirantia, Corde ventriculis przdito, { Duobus, Vivipara, [ Aquatica ; Cetaceum genus, || Terreftria, Quadrupedia, vel, ut Manat: etiam < complectamur , pilofa. Animalia hujus {| generis amphibia terreftribus annnme- | T Bea | Ovipara, Aves. Unico, Quadrupedia Vivipara & Serpentes. Branchiss. repirantia, Pifces fanguinei preter i Cetaceos omnes. LBivang nia ements AN es { Mojora, quz vel || € Molliz, Manrdscy Poly pus, Sepia, Loiligo. ee Maarunsseoxa, Locutta, Aftacus Cancer. ‘ Teftacea, "OsegxoSeeue, que vel univalvia, vel |! bivalvia, vel eurbinata. , Minora, Infeca, Fig. 15. From Ray (1693): First attempt to exhibit affinities or relationships in animals by a tree-like diagram; comprising also amphibians and reptiles. 4, Familia. 29 authors without criticism. They made rather use of these earlier authors descriptions to identify the indigenous fau- na. They began to understand the pedagogic function of images and the importance of accuracy in representing the natural things in order to objectively describe them. The most important collection of natural history coloured paintings, among them 50 tables with amphibians and rep- tiles, originates from Ulisse Aldrovandi (1522-1605), hav- ing been detected for science and described in the last years. “Their number and quality allow this collection of images to be considered as the first attempt to organize an iconographic atlas of the Italian and Mediterranean fau- na and, without any doubt, the first collection of herpeto- logical images realized with relatively modern criteria” (Delfino, 2007; Delfino & Alessandro, 2008; Alessandri- ni & Ceregato, 2007). The lack of a species concept and the lack of the understanding of the animal organism, as well as the use of different engraving- and printing tech- niques (above all wood cuts) caused an excessive simpli- fication in most contemporary publications and rendered the figures useless to represent the distinctive character of a species (cf. Gessner’s figure; Fig. 2 hoc loco). The different insights of a painter and a mere engraver as well as the different qualities of a water colour and an engrav- ing are shown by a comparison of an Aldrovandi water colour (Fig. 3). It was about 250 years later, when the zoologist Hermann Schlegel (see Schmidtler, 2007) delved into the theory of natural science images (German translation in Nissen, Lacertini. Lingua tenuis, furcata, protractilis, scuta abdominalia et caudalia la- teralibus majora, haec omnia verticillata. squamae dorealibus sequales; cauda hicavinata —- Capitis < scuta, dorsalibus majora; collare nullum - Fig. 16. distinctum 5 poo Gula non dilatsbilis. a upmambis, (quadricarinata - Dracaena. rotundata - - Lacerta. - - - - = Tachidromus. Diagram of the family “Lacertini” in Oppel (1811): Oppel (1811: 20) established the family “Lacertini” (now: Lacerti- dae), among six families within his order ““Squamata’’. The “Lacertini” comprised four genera, two of them (Zupinambis and Dra- caena) belonging now to other families, before Duméril (1806: No. 49-51) had established two families only (“Planicaudati” and “Tereticaudati’’) in his order “Saurii’’.— Similar trees or identification keys like this and Duméril’s were the forerunners of an evo- lutionary view in herpetology (see Schmidtler 2009). Bonn zoological Bulletin 57 (2): 307-328 ©ZFMK Ww ho bo 1 Stannn, Ul. Gtannn. X. Ordnuna, E. Ovounung, @®reehkone, Kialoten. I. Ordnuag, Hlugeidechfer. ILE. Stamm. IV. Stamne. LE. Ordmung. Flugeidechfer. Sramn Stat WEE, Ovdnung, EF. Oronung. — ¥, Ovdnung. Chamileone, CECidech fen. divokodile. War vive. IXY, Ordming, Megalefaurier, 1a. Ordnung. WE. Ordnung. MeereiDechfen. Bepfe. V. Stain, Le. Ovtnung, HO. Ovdnung. YF. Ovdnung. Schildkroten. Schlangen. Srifche. UWL Ordnung. Sulawawder, dT, Ordnung, Cicilien. Fig. 17. Diagram from Kaup (1836): His “III. Ordnung E1- dechsen” (,,Lacertae“, text p. 26) corresponds to the rank of the present day family Lacertidae, comprising several genera like the “eigentliche Eidechsen” = Lacerta. His “Stamm” corresponds nearly to a current order. In this diagram there is exhibited the famous and strange dead end of “natural philosophy”; display- ing more a mysticism of numbers than a concept of natural sci- ence: Like in birds, mammals and amphibians as well, there exists in Kaup only a strict number of five “Stémme” and in each there are enclosed strictly three “Ordnungen”! See chapter 3.4. This mysticism does not proceed in the number of lacertid ge- nera. See also Fitzinger (1843) and the critique in Mertens (1973). acerla Arico ts | Notopholis / \ Gtxnyprt Fig. 18. Preertidn Josef Friedrich Schmidtler 1978: 231). According to him it is the function of such a figure to supersede the subjects difficult to be seen or ex- amined in nature, in order to recognize them after the de- piction and to be able to derive their shape, colours, pro- portions and other features as exact as possible. The per- fection of this claim calls for a full interaction of text and illustration in the scientific, artistic and technical aspects — in this article being demonstrated on the basis of the Lin- nean genus Lacerta. When the taxonomic importance of the different struc- tures, like scales, spotting and colouration had been rec- ognized about 1800, there originated also the need to dis- play them separately from each other. The joint — natura- listic — appearance interfered with their independant per- ceptibility, for instance because of their poor visibility (e.g. sutures of scutes), overlapping with spotting, light reflec- tion (Figs 8, 9), etc. At the same time abstracted figures (diagrams) were also used to exhibit relationships and / or identification keys, etc. (Figs 15—20). —— Puedes SS oN Be yi | Podareis / Nnmon j orgllalips From Camerano (1877: Tav. I, excerpt): Another rather popular kind of diagram displaying the relations between taxa. The small circles within the large circles symbolise related taxa of the four lacertid genera (Lacerta, Podarcis, Notopholis, Timon). The lines between such encircled taxa designate important morphological resemblances. Bonn zoological Bulletin 57 (2): 307-328 OZFMK Taxonomic history of the genus Lacerta 323 The first engraving — and printing — techniques were wood cuts (15'* century), and shortly later, copper engravings. Around 1800 copper engraving was refined (etchings) and the first very expensive and rare colour prints in herpetol- ogy were based upon them (Daudin 1802; see Schmidtler 2007) after centuries of hand colouring. Lithography (the first lizards in Schmid 1819) was invented and improved more and more up to the time of chromolithography (see Fig. 8). Bewick (e.g. 1809-1816; Fig. 7) renewed the wood cuts (“Wood engraving”). In the second half of the 19th century the first photographs appeared (see Niekisch, 2010; Fig. 9 hoc loco) and revolutionised the book-illus- tration also in natural science and herpetology together with new letterpress printing techniques. These technical advances were attended by the expansion of some zoo- logical / herpetological disciplines — or facilitated their proliferation — like ethology, husbandry, ecology. It is noteworthy to emphasize that each of the engraving- and printing techniques displays its technical, artistic and economical strengths and weaknesses as measured by the different requirements they have to satisfy (see Schmidtler, 2007). Book illustrations were always expensive and this was the most important reason why the informative val- ue of many important works had to suffer immensely. Greek Archipelago, S. Russia, Constantinople, Asia Minor, 3.2 Naturalistic figures As was explained above, the lack of species recognition was especially responsible for rendering many figures use- less in representing distinctive characters. This was espe- cially the case for many of the reptiles, being regarded as abhorrent or less important (except the venomous snakes), compared to mammals or birds — up to the Linnean times and later. A good example is Gessner’s fabulous creature (Fig. 1), called “‘Lacertus viridis”, and typically attended with a po- em advertising a medical and cosmetic prescription. It is only the indentations across the tail which suggests the possible belonging to the current Lacertidae. Shortly before Linnaeus, the “Thesaurus” of the wealthy pharmacist Seba (1735) described many exotic animals from Seba’s “Wunderkammer”. This voluminous work de- picted also many mythical creatures — besides some in- digenous reptiles, e.g. the male of a “Lacerta viridis” (= Lacerta agilis), being identifiable only by means of the distinctive dorsal pattern and the green and brown colours (see Schmidtler, 2004: Abb. 1). i 8 rudss Spanish Peninsula, Baleares, c mays OA ftaly with Corsica & Malia. i a portschinskit | dofilippii PaaeecosecocOsoass Sc aE ie ae ee Sei SEAS Dh eanleay RSE = | ‘ i Se | saxicola ' ' H o-2 5 ot “ ' <] q = 1 horvathi | bedriage sardoa ' hispanica — monticola t > er , : (South Hungary) ' ES 3 chalybdea . caitcasica ; t = <= 1 ee ee Ss erhard; | siliguerta wa | ' ea teroclvphi } ' \ hieroglyphica } : g ares ee aces toca oe lee Soh = || insulanica nigriveniris _filfolensss a lilfordi = | is He ky ; a 6 H | quadrilineata 1 pityusensis | 2 a 3S ’ : j i ; oe = | | i. brveggemanni AS ees hiolepis bocavii vaucheri ' = ee 4 S : 2 2 ES ae a CP TSR SROs Soca dee see Orne! SEE apa oe eeSoeer nets F Si 3 be serpa eee wt @ = sateoncese 4 Zi 5 8 } typicg N » : R ui Easi Coast & Islainds of the Adriatic, be Seeds event (31hl laatier | Li (iran?) 9) Sie? @igss-ssee+ La eee ante on RR rp NRE SUE EEA Bote a > er er er brev ceps S oe is campestris _ fimane pe —— tieliselle: Fig. 19. From Boulenger (1913: 205-206): Linear diagram of the forms of Lacerta muralis: Citation: “The preceding diagram expresses their affinities, as I conceive them, and also their distribution. It will also enable the reader to see at a glance how my views on the possible derivation of these forms differ from those advocated by Prof. Méhely.” The “varieties” of Boulenger’s on- ly species “Lacerta muralis” comprise four current genera (Podarcis, Darevskia, Iberolacerta, Archaeolacerta) with at least 17 current species and some more subspecies. Bonn zoological Bulletin 57 (2): 307-328 ©ZFMK 324 B oe perenne GOMotia galloti Psammodromus algirus = Omanosaura jayakari 100 ke eran _— === Omanosaura cyanura 68/51 | #797 i y AW cota Re : = Lacerta agilis 0.05 subs./site _ Fig. 20. 0 Podarcis muralis ————— aoe SCelorcis perspicillata Teira dugesii === Timon princeps == Timon lepidus 100 beens TION pater Josef Friedrich Schmidtler Gallotiinae Eremiadini ees Lakydromus sexlineatus > Archaeolacerta bedriagae Zootoca vivipara Podarcis taurica dUpHyAIVy] 1938] IBUIGAIIV'T aI Phoenicolacerta laevis gem. nov. = Iberolacerta bonnali Iberolacerta horvathi Dalmatolacerta oxycephala gen. nov. ——==- Apathya cappadocica Hellenolacerta graeca gen. nov. Anatololacerta danfordi gen. nov. Parvilacerta fraasii gen. nov. Algyroides marchi Franolacerta brandtii gen. nov. Darevskia chiorogaster — Darevskia saxicola From Arnold et al. (2007: Fig. 1B): The most actual phylogenetic tree of the Lacertidae comprising in detail the tribe Lacertini (among them Lacerta and the new genera Phoenicolacerta, Dalmatolacerta, Hellenolacerta, Anatololacerta, Iranolacer- ta), having been broken up from it). The relationships are indicated by DNA sequences (ML tree from a reanalysis of the mt DNA data set of Harris et al. (1998) based on cytochrome b, 12 S RNA and 16 S RNA). Different probability values resulting from Baye- sian analysis are indicated. The changes in scientific insight are visible by the differ- ences of the earlier drafts and the final hand coloured en- graving in Roésel v. Rosenhof’s (1758) famous frontispiece (see Fig. 4). Garsault (1764: pl. 688; fig. 5 hoc loco), a forerunner of Laurenti (1768), and a splitter like him (Chapter 2), moved an almost correct drawing of lacertids forward. In contrast to this figure, Laurenti’s (1768: Tab.I, Fig.IID) Seps caerulescens (= Lacerta agilis) has accomplished perfec- tion for the first time in the history of a lacertid drawing. Laurenti’s (1768) image remained unique for some decades. Even the excellently hand coloured copper en- Bonn zoological Bulletin 57 (2): 307-328 gravings of male and female specimens of Lacerta agilis in Wolf in Sturm (1799) and Bechstein (1800) display some deficiencies in pileus scutellation (cf. also Schmidtler 2004). The black / white and coloured engrav- ings in Daudin (1802) are comparably of a very different quality. The colours of his excellently drawn shape of L. ocellata (1802: pl. XX XIII; without the blue ocellae) re- veals that he did not see a live specimen. But the shape of the adult male is to the point. In Daudin (1801—1803), part of the so called Sonnini edition, for the first time the progressive but extremely expensive technique of colour printing was used in herpetology (see Schmidtler 2007). ©ZFMK Taxonomic history of the genus Lacerta 325 On the contrary, the hand coloured engravings in Sonni- ni & Latreille (1801) are very small — and bad; likewise the figures in the numerous popular editions of the French natural histories, named “Buffon — Cuvier — Lacepéde’”’, are out of the question. Their images were cribbed per- manently and often lost their quality step-by-step up to an unrecognisable condition. In the second half of the 19th century research on colours and patterns, the biological reasons and causes of their adaptation, became important for the evaluation of infra- specific variation and biology in general (Eimer, 1881: Taf. XHI-XV). Subsequently, many subspecies, especially within the current genus Podarcis (then mostly “Lacerta lilfordi, Lacerta melisellensis, Lacerta muralis, Lacerta sicula”), were based upon minute differences in scale counts, colouration and pattern (see Mertens & Wermuth 1960). As a result of the genetic revolution in the last years the importance of naturalistic figures in lacertid system- atics is on the decline. At the same time top-quality pho- tos (Fig. 10) have gained in importance especially in pop- ular vivaristic publications. 3.3 Schematic figures The abstraction of systematically important features, be- ing more or less hidden to the unprejudiced observer, is a condition for successful species recognition. The first noteworthy attempts towards a schematisation of zoolog- ical / herpetological features were displayed by Linnaeus in his earlier editions of the “Systema Naturae” (Fig. 11). It is above all the scale counts of ventrals and subcaudals in snakes which are explained in his table III (Linnaeus 1756). Only these scale counts are given in the diagnoses of snakes (see also Linnaeus 1758 and 1766). The decisive step forward in lizards was made by Mer- rem (1820). Based upon his similar system in snakes (Mer- rem 1790, 1820; see Schmidtler 2006) he gave names to the pileus shields of an adult Lacerta ocellata (now: Ti- mon lepidus; see his page XI and XII) and depicted their abbreviations in this figure. This description (see Fig. 12) covered seven types of scutes with the letters A (Wirbelschilder — Scuta vertebralia), B (Hinter- hauptschilder — Scuta occipitalia), C (Augenbrauen- schilder — Scuta superciliaria), E (Stirnschilder — Scuta frontalia posterioria), F (Schnautzenschilder — Scuta frontalia anterioria), G (Rtisselschild — Scutum rostrale, L (Nasenlécherschilder— Scuta nasalia). This system was later on differentiated and improved by Milne Edwards (1829: pls. 5-8) who depicted and described also the shields of the lower sides of head, body and limbs. The concept of Merrem (1820) and Milne Edwards is valid up to now. Bonn zoological Bulletin 57 (2): 307-328 In the middle of the 19 century important osteological investigations also were executed in lacertids. They al- lowed taxonomic research in the higher categories but al- so within lacertid genera and species, after a reasonable schematization in osteology, above all in skull terminol- ogy, had been found (Fig. 13 from Méhely). They brought about the famous and interminable controversy of L. v. Mehely (“splitter”) and G.A. Boulenger (“lumper’’) on the then intractable “Muralis-Frage” (see Méhely 1909; Boulenger 1920; among others). It was the research since the middle of the 18" century which revealed the crucial importance of the dorsal pat- tern especially in the specific and infraspecific taxonomy of the current genus Podarcis. Eimer (1881: Taf. XIH; Fig. 14 hoc loco) named the different longitudinal zones (“I bis VI erste bis sechste Zone”) which usually exhibit a sys- tem of n narrow longitudinal light streaks (nrs. I and III, “Grenzlinien”’) and dark bands (nrs. II,” inneres / 4uBeres Band”). Méhely (1909: Fig. 1) eased his terminology and gave it the presently valid content. The seven light streaks and dark bands were named after their initial points at the pileus shields (like “Supraciliarstreifen” and “‘Occipita- band”’); see also Schreiber (1912: Fig. 68; p. 333-335) and Mertens (1915: Fig. 3). Admittedly, morphological schemata (scutellation, dorsal pattern) like these have lost their crucial taxonomic im- portance in the 19th and 20‘ centuries during the last two decades because of the reasons given above (see Section 252) 3.4 Diagrams Semi-verbal depictions known in many different shapes (concerning biology as a whole) are book illustrations in broader terms. Contrary to naturalistic figures or the schemata discussed above, phylogenetical trees, based up- on genetic research, have become indispensable parts of comprehensive taxonomic work in the last years (Fig. 20). In many analyses the genetic distances currently have to- tally replaced the traditional taxonomic decisions based upon morphology and reproduction biology — be it ap- propiate or not. Tree-like diagrams, comprising also reptiles, trace back to Ray (1693) (see Fig. 15). They have an enormous im- portance in the history of general biology, not only in lizards. In the field of herpetology, lizards included, they became common practice since the basic works of Duméril (1806) and Oppel (1811; Fig. 16 hoc loco; see also Schmidtler 2009) and Cuvier in Cloquet (1819). Strange to say, it was not clear in those pre-evolutionary times, if the trees should represent identification keys only, or if ©OZFMK 326 Josef Friedrich Schmidtler they should depict relationships (too) when illustrating the Linnean hierarchical system by diagrams. Gould (2003: 105) explained the secret of the then triumph of the Lin- nean categories (from species to class) being nested into one another, by the circumstance that this system later on was capable of being converted into a phylogenetic inter- pretation (see also Schmidtler 2009: 500, Figs 7a—d). The subsequent diagrams are not yet phylogenetic trees in a strict sense; especially the one by Kaup (see Fig. 17), who was a follower of the fanciful “natural philosophy”. 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VII.2010 Accepted: 20.[X.2010 ©ZFMK Bonn zoological Bulletin | Volume 57 | Issue 2 | pp. 329-345 Bonn, November 2010 A brief history of Greek herpetology Panayiotis Pafilis !.2 'Section of Zoology and Marine Biology, Department of Biology, University of Athens, Panepistimioupolis, Ilissia 157—84, Athens, Greece 2School of Natural Resources & Environment, Dana Building, 430 E. University, University of Michigan, Ann Arbor, MI — 48109, USA; E-mail: ppafil@biol.uoa.gr; pafman@umich.edu Abstract. The development of Herpetology in Greece is examined in this paper. After a brief look at the first reports on amphibians and reptiles from antiquity, a short presentation of their deep impact on classical Greek civilization but also on present day traditions is attempted. The main part of the study is dedicated to the presentation of the major herpetol- ogists that studied Greek herpetofauna during the last two centuries through a division into Schools according to researchers’ origin. Trends in herpetological research and changes in the anthropogeography of herpetologists are also discussed. Last- ly the future tasks of Greek herpetology are presented. Climate, geological history, geographic position and the long human presence in the area are responsible for shaping the particular features of Greek herpetofauna. Around 15% of the Greek herpetofauna comprises endemic species while 16% represent the only European populations in their range. THE STUDY OF REPTILES AND AMPHIBIANS IN ANTIQUITY Greeks from quite early started to describe the natural en- vironment. At the time biological sciences were consid- ered part of philosophical studies hence it was perfectly natural for a philosopher such as Democritus to contem- plate “on the Nature of Man” or to write books like the “Causes concerned with Animals” (for a presentation of Democritus’ work on nature see Guthrie 1996). The very name of the discipline of herpetology derives from the Greek words epzeto (reptile) and Adyog (science) while the term amphibian reflects the typical dual (aquatic and ter- restrial) life style of frogs (from the Greek augi — both— and Biog — life). The first formal Greek herpetologist was Aristotle himself. In his books on animals (History of Animals, Generation of Animals and On the Parts of Animals) the father of zoology discussed the morphology, physiology and classification of reptiles and amphibians. Nicander was fascinated by the lethal power of snakes, focusing on the venom of serpents in two of his surviving poems (Theriaca, Alexipharmaca, see Knoefel & Covi 1991). In late antiquity Pausanias, though he wasn’t a naturalist, gave interesting information on the fauna and flora in various locations in Greece through the ten books of his notorious Description of Greece (EAAddoc¢ TMeptuiyots). Bonn zoological Builetin 57 (2): 329-345 Therein one could find citations to the Greek herpetofauna such as the Seriphian frogs or the tortoises of Arcadia. REPTILES AND AMPHIBIANS IN GREEK CULTURE Snake venom and the ability for ecdysis had deeply im- pressed ancient Greeks who incorporated reptiles in many of their myths. Snakes were considered magical creatures, capable of both good and evil, and were associated with chthonic religious beliefs. In Minoan Crete snakes repre- sented the underworld deities and were worshiped. Tens of statuettes depicting the Goddess of Snakes have been found in excavations all around the island. Ophion (from the Greek ophis — oqis meaning serpent), one of the mighty Titans, was the first ruler of Mount Olympus be- fore he was cast down by Cronus and Rhea. According to legend the first king of Athens, Kekrops, was half-snake half-man (Supvng meaning double nature) and thanks to his wisdom he decided to offer his city (known as Kekropeia at the time which afterwards changed to Athens to honor the patron goddess) to Athina instead of Poseidon when the two immortals were fighting over its possession. On the other hand Medusa (or Gorgon) the mythical monster that had snakes instead of hair, could turn anyone who ©ZFMK 330 Panayiotis Pafilis looked at her into stone. Perseus, using his shield as mir- ror, made Medusa look at herself and then decapitated her. He then offered her head to Athina who put it on her own shield (known as gorgoneion) so as to petrify her enemies. A serpent-like dragon, Python, was sent by Hera after Leto, mother of Apollo and Artemis, to punish her for hav- ing an affair with Zeus. Young Apollo took revenge for his mother by killing Python at Delphi, where the serpent dwelled. Since then the priestess of the oracle was named Pythia. The etymology of the name derives from the verb pythein (mbevv, “to rot”), referred to Python’s flesh in the state of decomposition. The priest of Poseidon Laocoon warned the Trojans about the Trojan Horse and tried to convince them to burn it. Athina, who was supporting the Greek army during the War of Troy, sent two snakes to strangle and kill Laocoon together with his sons. Greeks were aware not only of the lethal power of ven- om but also of its healing properties. In the statues of Hygeia, the goddess of health (the meaning of the word in Greek), a snake is lying on her shoulders. Aesculapius, the god of medicine and son of Apollo, was carrying al- ways his famous rod, a snake-entwined staff (the species was Zamenis longissimus). In his most magnificent tem- ple in Epidaurus, that used to function as a hospital, a strange construction known as tholos (dome) was erect- ed. Patients spent the night inside tholos together with tens of snakes that were believed to heal them. Two small snakes were coiled around Hermes wand, symbolizing the wisdom with which he spoke, since he was considered, together with Athina, god of eloquence. A fascinating story about the symbolic role of reptiles in antiquity comes from the island of Aegina. During the pe- riod of Aegina’s naval acme (6 century B.C.) the is- landers coined silver staters depicting the sea turtle Caret- ta caretta. However a terrapin (Zestudo sp.) replaced the sea turtle when the neighboring Athens inaugurated its long period of thalassocracy in Greek seas (after 480 B.C.): Due to the arid climate Greeks were more familiar with reptiles than amphibians. Thus only few references are known from antiquity, like the Aristophanes’ comedy “Frogs” (Batpayot) or the silver stater that Seriphians coined (ca 530 B.C.) to honor their local hero Perseus since frogs were associated with his cult (Pausanias, 2nd century A.D.). Frogs from Serifos Island were famous in antiquity for not croaking (another story linked to Perseus legend) and the expression “Seriphian frogs” was used as a popular proverb during ancient times for people refus- ing to talk. With the prevalence of Christianity reptiles become the personification of evil, starting from the Original Sin. Saint Bonn zoological Bulletin 57 (2): 329-345 George and Saint Demetrius, the so called militant saints, are depicted as dragon slayers, symbolizing the triumph of Good, as expressed by the Greek-Orthodox Church, over Evil, the former idolatry faith. The Serpentine col- umn, dedicated by Greeks in 479 B.C. to Apollo’s altar at Delphi to commemorate the victory over the Persians at the battle of Plataea, was formed by three intertwined snakes (Tpixdprnvoc Oguc), meaning three-headed snake). Constantine the First moved the column to the Hippo- drome of his new capital. However the people of Constan- tinople destroyed the higher part of the column (the heads of the snakes) since they thought it was the representa- tion of the devil. On the other hand the Apostolic Fathers recognizing the wisdom of snakes were advising the first Christians to be “prudent as the serpent” (Ignatius of An- tioch to Polycarp of Smyrna). Traces of the ancient beliefs still echo in folklore and traditions. The presence of geckoes in a house is considered good fortune. In many households in Cyclades people used to fill with milk a small cup for the “snake of the house” (in Greek onitdgid0, Zamenis situlus). The most amazing case though comes from the island of Cephalonia where pagan creeds survive together with christian rituals at the temple of Madonna of the Snakes (Ilavayta yn gidovoa). According to the legend a monastery stood at the very same place. When pirates disembarked close to the spot and tried to conquer and harry the treasures of the monastery, nuns prayed for help and Virgin Mary sent snakes that surrounded the building and scared away the pirates. Every year at August the 15th (when Greek Orthodox Church celebrates the Dormition of Holy Mary) locals collect cat snakes (Telescopus fallax) days prior to the feast and put them by the icon of the Virgin. Pilgrims touch these snakes and even let them coil around their shoulders or hands since they believe that they will protect them from sickness. GEOGRAPHY, BIOGEOGRAPHY AND SPECIES RICHNESS Greece is one of the small European countries with a to- tal area of around 132,000 km?. However its unique lo- cation at the biogeographical crossroads of three conti- nents, each making its distinct biological contribution, makes the country an invaluable site for biodiversity (Lymberakis & Poulakakis 2010). The rough geological mosaic encompassing mountain chains that separate the country into clearly distinct climatic zones and the large number of islands (approximately 8000, most of them in Aegean Sea) have a huge impact on the flora and the fau- na (Hausdorf & Hennig 2004). Hundreds of endemic species are hosted in both the mainland and the islands highlighting the region as a hot spot of endemism. ©OZFMK A brief history of Greek herpetology 331 The climate is typical Mediterranean with long, dry and hot summers and mild winters (though in the north and the mountainous parts of the country winter period could be harsh). These conditions are ideal for reptiles, which thrives in the hospitable Greek habitats but also support various amphibian species and populations. Despite the small size of the country, Greece is home to one of the richest herpetofaunas of Europe hosting 64 species of rep- tiles and 22 of amphibians. Ten of the reptilian species are endemic and 11 maintain their only European populations in Greece, whereas the respective figures for amphibians are 3 and 2, respectively. The majority of Greek reptiles and amphibians has Palearctic origin and are common in most of Europe or the Balkans (e.g. Natrix natrix or Bombina bombina). However, Greece hosts also species of Asian (e.g. Mon- tivipera xanthina, Lyciasalamandra luschani) or even African (Chameleo africanus) origin. Definitely the most interesting group is the one comprised of the endemic species, the majority of which are islanders, such as Pelo- phylax cerigensis, Podarcis levendis or Macrovipera schweizeri. The range and particularities of Greek reptiles and amphib- lans are, in a considerable degree, the result of the recent geology of the eastern Mediterranean Basin. During the Messinian salinity crisis, the Mediterranean Sea underwent long periods of desiccation that, in Aegean Sea, led to the emergence of landmasses that become islands. The old- est Greek islands (Crete, Skyros and Karpathos) remain to this status even after the Zanclean flood. The Ice Age periods during the Pleistocene with their consecutive freezing and warming conditions had a strong impact on the area, shaping glacial refugia that harbored many cold- intolerant species, which afterwards reinvaded the rest of the Balkans (e.g. Rana graeca). Many islands were con- nected either to mainland Greece or Asia Minor as a con- sequence of the low sea level during the last Ice Age pe- riod. Nowadays the herpetofauna of these islands still re- flects this geological incident with islands closer to Greece having a clearly “European” composition (e.g. Evvoia, Thassos), whereas those next to Asia Minor show a more “Asian” character (e.g. Lesvos, Chios, Samos). This sep- aration between European and Asian herpetofaunas 1s fur- ther supported by the existence of a deep-water trench run- ning over the Aegean Sea from southeast to northwest, separating the eastern “Asian” cluster from the western “European” one with only few exemptions. Another im- portant geological factor is the intense volcanism of the region. The Aegean volcanic arch, spanning the southern part of the area, was formed during the Pliocene as a con- sequence of the northward subduction of the African plate beneath the Aegean one (Fytikas et al. 1984). Milos Arch- ipelago, a small but extremely important in terms of en- Bonn zoological Builetin 57 (2): 329-345 demism island group, was separated from the rest of the Cyclades by middle Pleistocene as a result of volcanic ac- tivity (Sondaar et al. 1986; Dermitzakis 1990). In summary, most endemic species are concentrated to the oldest islands where the long history of isolation provid- ed the necessary conditions for speciation. The astonish- ing variety of subspecies in the islands, for instance 19 for Podarcis erhardii and 13 for Cyrtopodion kotshyi reflects the importance of insularity in the evolution of different morphs. In mainland Greece endemic species are located in the southernmost part of the country, Peloponnese, thanks to historical biogeographical reasons (glacial refugia) matched by a fair period of isolation. Last, but certainly not least, humans had a significant con- tribution in shaping the Greek herpetofauna. In the Aegean Sea navigation started quite early (around gth millenium B.C., Kotsakis 1990; Simmons 1991). Voyagers carried materials (e.g. marble or pottery) that offered an excellent opportunity for transportation of small-bodied species or their hidden eggs (typical examples are Hemidactylus tur- cicus and Tarentola mauritanica and most probably Lau- dakia stellio). In some other cases humans may deliber- ately transport reptiles or amphibians related to religious beliefs. Apart from dispersal, human activities favored rep- tiles with the deforestation of the largest part of the coun- try, providing opportunities for thermoregulation and for- aging. Thousands of kilometers of dry-stone walls all around the country, and especially in the Aegean islands, offer ideal hiding places and support thriving populations. On the other hand touristic development with its acces- sory consequences (water over-pumping, wetland drainage, habitat degradation), over grazing and intensive agriculture has largely altered the landscape, influencing negatively upon reptiles and, mostly, amphibians. THE FRENCH MOREA EXPEDITION The Morea (the Greek vernacular name for Peloponnese) Expedition (French: Expédition de Morée) accomplished by the French Army at the end of the Greek War of Inde- pendence. After the naval battle of Navarino where the united Franco-Russo-British fleet destroyed the Ottoman fleet, French expeditionary corps disembarked at south- ern Peloponnese to secure the evacuation of the area from the Turks. Following the example of the successful Napoleon’s Egyptian Campaign where a scientific com- mittee accompanied the French troops, a scientific mis- sion escorted the expedition in Peloponnese. The Head of the 17 experts of different disciplines that comprised the mission was the naturalist Jean Baptiste Bory de Saint Vin- cent. Bory collected hundreds of plants and animals that were sent to France for further identification and classi- ©ZFMK fication. It was from these specimens that the herpetology of Greece began formally in 1833, when the first endem- ic species to Peloponnese were described by Bory and his colleague Gabriel Bibron, who also participated in the Morea expedition. Bibron worked extensively on Her- petology and helped his mentor Duméril in the publica- tion of the first herpetological monograph Erpétologie generale (1834-—-1854) where many species distributed in Greece were described. The Morea Expedition covered not only Peloponnese but also numerous Greek islands. The importance of this mis- sion was crucial and later studies on Greek herpetofauna were largely based on the Expedition’s observation. Bibron and Bory described in total three species (A/gy- roides moreoticus, Podarcis peloponnesiacus and Ophiomorus punctatissimus), while later Duméril and Bibron, using specimens from Corfu, described one species (Algyroides nigropunctatus). THE GERMAN SCHOOL It is widely accepted that Greek herpetology, at least dur- ing its early period, literally “spoke German”. Eminent herpetologists from Germany, Austria and Switzerland worked (and are still working) extensively on Greek rep- tiles and amphibians, setting the basis for herpetology in the country. Maybe the underlying reason should be seeked in the first king after the War of Independence, Ot- to the First, son of Ludwig of Bavaria (Wittelsbach House), who brought with him hundreds of Germans to staff the administration of the new country. In this session the most important contributions in Greek herpetology were presented. The first German naturalists who arrived in the country and presented information on Greek amphibians and reptiles were not herpetologists but ornithologists (Erhard, Reiser) or botanists (Heldreich, Herzog). Hence many of their first observations proved to be incorrect since they were not familiar with herpetological systematics. Jacques von Bedriaga wrote the first major monograph on Greek reptiles and amphibians in 1881. After receiving his PhD Thesis from the University of Jena he started to trav- el very frequently to both Italy and Greece. The fruits of these trips was his “Die Amphibien und Reptilien Griechenlands” which was published in Moscow in three volumes. His special interest on lacertids is best reflect- ed in the description of four new species, two of which are endemic to Peloponnese and Milos Island (namely Hellenolacerta graeca, Podarcis milensis, Lacerta trilin- eata and P. erhardii). Though Bedriaga was born in Rus- sia, where he also took his bachelor’s degree at the Uni- Bonn zoological Bulletin 57 (2): 329-345 332 Panayiotis Pafilis versity of Moscow, he became scientifically active in Ger- many and published most of his works in German. That’s why his name is herein included in the so-called German School. Oskar Boettger, though never visited Greece, made the second important contribution to Greek herpetology. Dur- ing the years he was infirm and remained at home, he re- ceived numerous specimens sent by his many friends and colleagues. Among them von Oertzen shipped him rep- tiles and amphibians he collected while in Greece. Boettger worked on this collection and later published his findings (1888, 1891). On of the most prominent European herpetologists, Robert Mertens, worked also on the Greek herpetofauna. He re- alized at least three herpetological excursions in the coun- try, which later resulted in a series of paper (1959, 1961, 1968a, 1972). Using types and specimens from the large collections of the Senckenberg Museum in Frankfurt he also wrote systematics articles (1955, 1968b). His most significant contribution though, was the publication, to- gether with Miiller (1928, 1940) and Wermuth (1960), of the European checklists of amphibians and reptiles. This book has been a useful reference for researchers of the Greek herpetofauna. At this point it is worthy of mention- ing that Miller himself contributed one of the first her- petological papers on Greek herpetofauna in 1908. Karl Buchholz and Ulrich Gruber, both curators of her- petology in the Zoologisches Forschungsmuseum Alexan- der Koenig (hereinafter ZFMK), dedicated a large part of their research on Greek reptiles and especially to the is- land populations. Buchholz undertook numerous herpeto- logical excursions to Greece and collected many speci- mens (being an excellent markeman he shot his targets from long distance). His collections were published in a series of paper on the Aegean reptiles (1960, 1961, 1962a,b). Gruber focused also on insular populations (Gru- ber & Fuchs 1977, Gruber 1979) and following Werner’s example, specialized in the North Sporades island group (Gruber & Schultze-Westrum 1971, Gruber 1986). Hans Schneider, one of the leading researchers of amphib- ians, worked closely with Sofianidou and Kyriakopoulou- Sklavounou analyzing the acoustic properties of various species of frogs (1984, 1985, 1988, 1993). Acknowledg- ing the impact of his research on Greek herpetology he was invited as the pleninary speaker at the 10'* Meeting of Societas Europea Herpetologica in Crete in 1999 (Bioa- coustic studies in European Anurans). Many more German herpetologists, professional or ama- teurs, did research on Greek species and it would be 1m- possible to mention all of them in this brief paper. In any ©ZFMK A brief history of Greek herpetology 333 case it would be an omission not to mentioned B. Schnei- der who reported on the herpetofaunas of many Greek is- lands (e.g. 1986, 1995), A. Beutler who was interested al- so in insular populations (1979, Beutler & Froer 1980) and B. Trapp who investigated the Greek population of Chamaeleo africanus (e.g. 2003, 2004) and also wrote a book on Greek amphibians and reptiles in German (2006). Wolfgang Bohme is maybe the last of the Mohicans of the one-time all-potent German School. His engagement with Greek herpetofauna dates back to the time he was a stu- dent at the Christian-Albrechts University of Kiel. At 1969 he traveled with a friend to the Syrian borders of Turkey. On their way back they visited Thessaloniki and planned a field trip to study the endemic species of Peloponnese. Unfortunately their old Volkswagen “beetle” let them down in Athens so they had to cancel their excursion un- til the engine could be fixed. However they didn’t waste their time and attempted a herpetological survey of the area surrounding Acropolis. Their persistence was reward- ed with observations on Chalcides occelatus and Zame- nis situlus while they also discovered a dense population of Lacerta trilineata (specimens of this population can be found in the collections of ZFMK). In 1971 Wolfgang Bohme took office as Curator of the Herpetological Collection in ZFMK. His predecessors, Karl Buchholz and Ulrich Gruber, were keenly interest- ed in Greek herpetofauna, as mentioned above, and thus enhanced considerably the collections of the Museum. Thanks to them the newly appointed BOhme was able to immerse himself in the Greek collections during his cu- ratorship. It was in a series of Pseudepidalea viridis spec- imens collected by Buchholz in Peloponnese that BOhme discovered two misplaced adult individuals of Pelobates syriacus, the first record of this species in Greece (1975). The aborted field trip to Peloponnese finally took place in 1996, after his participation in the Congress of the Hel- lenic Zoological Society in Athens where he presented a paper on the Cypriot herpetofauna. During this trip Bohme went to Sparta and Mystras and observed many endemic species in situ. But another chance to visit Peloponnese would come from the far past. The former director of ZFMK, the archeozoologist Gtin- ther Nobis, had a house near Pylos. During his vacations he shot a black-and-white photo of a chameleon and up- on his return to Bonn gave it to BOhme. Since morpho- logical details were not discernible, B6hme assumed it to be C. chamaeleon and consequently published this record in a brief note (1989). In 1997 Bohme visited Nobis so as to have a first-hand examination of the species. Dur- ing this visit he met Andrea Bonetti and George Chiras who led him to the chameleon habitat where they soon de- Bonn zoological Bulletin 57 (2): 329-345 tected the first male individual. To their surprise instead of the typical small occipital flaps of C. chamaeleon, they found a tarsal spur, characteristic of the African species C. africanus. At the time the range of this species was be- lieved to be restricted only to Africa. Bohme and his col- leagues assumed that C. africanus was introduced to the area as result of the trade between Alexandria and Pylos, since the Gialova lagoon (the only place where the African chameleon is distributed in the country) is located to the exact site of the former ancient harbor of Nestor’s Palace (Bohme et al. 1998). The results were later verified with mtDNA analysis (Kosuch et al. 1999). This fascinating dis- covery came to corroborate the human influence on species dispersal in the Mediterranean Sea. Together with Evgeny Roitberg and his former PhD stu- dent Andreas Schmitz, now curator of herpetology in Geneva, Bohme traveled to Greece once more in 1999 to attend the 10'* Meeting of SEH in Iraklion, Crete. They made herpetological observations in Macedonia and at Mount Olympos. The last SEH Meeting in Kussadasi (2009) gave another opportunity to visit Greece. On his way back from Turkey, Bohme stopped, with his phd stu- dent Philipp Wagner, at various localities in northeastern Greece (Thrace and Macedonia). Last, but certainly not least, the impact of the Handbuch der Amphibien und Reptilien Europas (1981, 1984, 1986, 1993a) in which Bohme edited the volumes for snakes and lizards (and also contributed personally some species ac- counts — 1984, 1993b,c), has been catalytic for the devel- opment of herpetology in Greece. Data on ecology, sys- tematics, physiology and behavior were for the first time gathered and accessible to researchers. Besides the above, Wolfgang Bohme has another, more “indirect”, nonetheless important, relationship with Greek herpetology. During all the years he served as Head of the Herpetology Section in ZFMK (1971—2010) and Vice Di- rector of the Museum, he facilitated in every possible way researchers who were working on specimens from Greece. Many Greek herpetologists visited numerous times the rich herpetological collections of the Museum and retrieved valuable information on diet, reproduction, morphology, ontogeny, intra- and inter-population varia- tion, anatomy and phylogeny of Greek amphibians and reptiles. These data led to the publication of various sci- entific papers that considerably enlarged our knowledge of the Greek herpetofauna. THE AUSTRIAN SCHOOL Franz Werner was one of the most prolific and influential herpetologists who worked on Greek amphibians and rep- OZFMK 334 Panayiotis Pafilis tiles. Starting from 1894 he was an avid and consistent re- searcher of Greek herpetofauna and remained active till his death (1939), publishing a series of papers (1912, 1927, 1933, 1937, 1938). He was the first to describe the her- petofauna of Ioanian islands (1894) and he also expand- ed his studies to the Aegean Sea. Maybe his most impor- tant manuscript was the one published in English, some- thing quite unusual for a German-speaking scientist at the time, by the University of Michigan (1930). Therein he describes the findings of the visit he made at some Aegean Islands in 1927. In this mission he had the chance to col- laborate with K.H. Rechinger while informative photos shot by Prof. Schoenwetter illustrated the final paper. In his long herpetological pursuits in Greece Werner de- scribed four new species: Lacerta anatolica, L. oertzeni (a tribute to von Oertzen), Podarcis gaigeae (endemic to Skyros Archipelago and dedicated to Helen Gaige) and Macrovipera schweizeri. After his death his sons donat- ed his huge personal collection to the Natural History Mu- seum of Vienna, which since then is one of the wealthier in specimens coming from Greece. Werner’s pupil Otto von Wettstein followed up with en- thusiasm the work of his teacher on Greek reptiles. He took over as Curator of vertebrates at the Natural Histo- ry Museum of Vienna in 1920 and published his first pa- per on the herpetofauna of Crete in 1931. In his studies he emphasized the reptilian and amphibian populations of the Aegean islands of which he was a regular visitor. In 1942 he participated in a scientific mission to Crete that was conducted by a German Wehrmacht biological re- search squad. Without doubt his most important paper was the emblematic Herpetologia Aegea (1953). In the 182 pages of this landmark effort, Wettstein presented in the most detailed way, full of knowledge, all the information on the zoogeography of the Aegean Sea herpetofauna. The legacy Wettstein left to the Natural History Museum of Vienna is enormous and, luckily, his interest in Greek herpetofauna survived among his successors till today. Heinz Grillitsch, the actual Head of the Herpetological Collection since 1984, investigated aspects of the Greek herpetofauna (Grillitsch & Tiedemann 1984, Grillitsch & Cabela 1990, Grillitsch & Grillitsch 1991). Within his re- sponsibilities lies the heavy burden to maintain and pre- serve the huge collection, one of the greatest in Greek specimens. Werner Mayer from the molecular systemat- ic lab of the Museum has been working on the ecology and distribution of reptiles and continues to study the phy- logenetic relationships of numerous Greek lizards (May- er 1986, 1993, Mayer & Beyerlein 2002, Mayer & Arribas 2003). Franz Tiedemann, who 1s collaborating closely with the Museum, has conducted numerous studies on various aspects of herpetology on Greek species (e.g. Tiedemann & Haupl 1980, 1982, Tiedemann & Grillitsch 1986). Bonn zoological Bulletin 57 (2): 329-345 There are many more Austrian herpetologists that need to be mentioned here like Peter Keymar, who frequently vis- ited Greece and published papers on Greek amphibians and reptiles (1984, 1986a, b, 1988) or Thomas Bader and Christoph Riegler (2004, 2009) who described the herpeto- fauna composition of Rhodes Island. A special citation should be made to the very active Austrian herpetologi- cal group www. herpetofauna.at. In their excellent website one may find a wide variety of photos of Greek reptiles and amphibians since the members of the group have re- peatedly visited Greece. THE “INTERNATIONAL BRIGADES” Besides the predominant German and Austrian Schools that shaped the history of Greek herpetology, researchers from many other European countries made important con- tributions to the study of amphibians and reptiles of the country. Dodecanese islands during the first decades of the 20th century were under Italian rule and Italian naturalists de- scribed the herpetofauna of the region. Enrico Festa of the Museum of Zoology in Turin made herpetological obser- vations in the island of Rhodes that later were published by Calabresi (1923b) who also write his impression of a survey on Samos Islands (1923b). At the same period Et- tore Zavattari published a study on the fauna of the “Ital- ian islands of the Aegean Sea” (1929). Augusto Cattaneo is one of the most prolific authors on the distribution of Greek reptiles and amphibians, especially in the insular country (e.g. 1984, 1997, 1999, 2007). Another Italian her- petologist who investigated the range of Greek herpeto- fauna is Pierangelo Crucitti (e.g. 1990). The United Kingdom is represented by a handful of very productive herpetologists. Adrian Hailey (now at the Uni- versity of West Indies, Trinidad and Tobago), who worked for a long period at the University of Thessaloniki, em- phasized his research on the tortoise populations in Greece (e.g. Hailey 2000, Hailey & Willemsen 2003) while he al- so examined the metabolism of Laudakia stellio in col- laboration with Nikos Loumbourdis. Richard Clark wins easily the title of the champion of publications on the dis- tribution of Greek amphibians and reptiles. Starting from 1967 he wrote over 20 papers (e.g. 1968, 1971, 1989, 1996, 2000) covering most places of the country. Finally David Buttle traveled around Greece and published many new localities regarding the distribution of Greek herpeto- fauna (e.g. 1989, 1994, 1997). Nicholas Arnold with the different editions of his excellent guide on European am- phibians and reptiles (1985, 2004) offered an important reference book to herpetologists working on Greek species. ©ZFMK A brief history of Greek herpetolog 335 In the 1970’s Hans Lotze did many field trips in Greece and gave considerable information about snakes (e.g. 1974, 1977). Peter Beerli of Florida State University stud- ied Aegean water frogs (Beerli et al. 1996) and even de- scribed two new endemic species using molecular biolo- gy tools (Beerli et al. 1994): Pelophylax cerigensis and Pelophylax cretensis. But the real star of Swiss herpetol- ogists that involved the study of Greek species was un- doubtedly Hans Schweizer, the famous “Schlangenhansi”. Schweizer, an amateur herpetologist with a particular pref- erence to vipers, had already a reputation among the Eu- ropean herpetological community when in 1931 visited Milos Island. After spending considerable time walking throughout the island and observing lizards and snakes, he noted the striking differences between the local and mainland herpetofaunas. He begun to publish his findings (1932, 1935, 1938, 1957) and also contacted profession- al herpetologists around Europe, with whom he had a reg- ular correspondance, and started sending specimens. It was from one of those samples that Miller described the en- demic Milos grass snake in 1932 and dedicated it to him (Natrix natrix schweizeri). Thanks to Schweizer, Milos Is- land gained its reknown as herpetological hot spot in Mediterranean. Besides the grass snake, two more species bear Schlangenhansi’s name: Macrovipera schweizeri and Lacerta trilineata hansschweizeri. Otto Cyrén, one of the pioneers of Greek herpetology, was born in Sweden but spent many years of his life in Ger- many and consequently wrote in German his papers on Greek and Balkan herpetofauna (1928, 1933, 1935). Goran Nilson of Géteborg University, a viper expert, has exam- ined various aspects of the biology of Macrovipera schweizeri with his Greek collaborators Dimaki, Ioanni- dis and Dimitropoulos (Andren et al. 1994, Nilson et al. 1999). A younger representative of Swedish herpetology is Anna Runemark of Lund University who is doing her PhD thesis on the sexual isolation between mainland and inland populations of Podarcis gaigeae (Runemark et al. 2008). Two herpetologists from the Netherlands have studied the Greek herpetofauna. Ronald Willemsen focused on the study of Mediterranean tortoises (e.g. Willemsen 1991, 1999, Willemsen & Hailey 2002) while Henk Strijbosch examined the distribution and ecology of lacertids (Stri- jbosch et al. 1989, Strijbosch 2001). The Czechoslovakian Stepanek traveled to Greece and published an important contribution to the knowledge of Greek herpetofauna in 1944, along with two other papers (1934, 1938). Mario Broggi from Liechtenstein is a reg- ular visitor to Greece and has published over 15 papers on local herpetofaunas around the country (e.g. 1978, 1988, 1997, 2009). The Danish Henrik Bringsge is anoth- Bonn zoological Bulletin 57 (2): 329-345 er researcher that wrote on different species of reptiles and amphibians (e.g. 1986, 1997, 2004). Jeroen Speybroeck from Belgium has visited Greece many times and runs a well organized website with great photographs of Greek amphibians and reptiles (http:/www.hylawerkgroep.be/jeroen). GREEKS ON GREEK HERPETOLOGY Until the late 1960’s only foreign scientists, mostly from Central Europe, were researching on the Greek herpeto- fauna. In 1968 John Ondrias of the University of Patras (which hosts the oldest School of Biology in Greece) pub- lished the first list of amphibians and reptiles. That was the starting point that instigated many Greek zoologists to get involved in herpetological studies. Theodora Sofi- anidou of the University of Thessaloniki carried out the first dissertation on herpetology in 1977. Since then 18 re- searchers defended their PhD theses on herpetological sub- jects. Namely (in order of seniority): Loumbourdis (1981), Kyriakopoulou-Sklavounou (1983), Xyda (1983), Chondropoulos (1984), Tzannetatou-Polymeni (1988), Valakos (1990), Asimakopoulos (1992), Maragou (1997), Adamopoulou (1999), Vassara (1999), Kassapidis (2001), Poulakakis (2003), Pafilis (2003), Tsiora (2003), Sotiropoulos (2004), Mantziou (2006), Dimaki (2007) and Simou (2009). Bassilis Chondropoulos published the checklists of Greek lizards and snakes in 1986 and 1989 respectively. These papers remained for a long period the most dependable source for the distribution of the Greek herpetofauna. Sofi- anidou wrote the first complete herpetological monograph in Greek in 1999 on Testudo marginata. In 2000 Achil- leas Dimitropoulos and Yannis Ioannides published their work on the reptiles of Greece and Cyprus (in Greek), the first herpetological book to appear in Greece. Nowadays Greek herpetologists are working under the auspices of Universities, non-governmental organizations and Museums. The major groups of herpetological re- search are located at three Universities. At the Universi- ty of Athens (the oldest in the country) Professors of Ecol- ogy Ioannis Matsakis and Moisis Mylonas though not her- petologists encourage young people to work in the field and do indepth research during preparations of their dis- sertation. Rosa Maria Tzannetatou-Polymeni and Sratis Valakos, pupils of the aforementioned, became faculty in 1990 and 1992 respectively and with their turn supervised new herpetological PhD theses. Tzannetatou-Polymeni (assistant professor) is an expert on both Lyciasalaman- dra species and is actually supervising a PhD thesis on the endemic Helversen’s salamander (Karpathos and Kas- sos islands). Valakos (associate professor) laid a founda- tion for an active group that has already produced five dis- OZFMK 336 Panayiotis Pafilis sertations while two more are in process. Together with colleagues from other institutions he published an accom- plished guide for the amphibians and reptiles of Greece (in English) in 2008. Earlier, with his collaborators, wrote the first volume on a local herpetofauna (2004, in both Greek and English). His research focuses on the environ- mental physiology and phylogeny of lacertid lizards (e.g. Valakos 1989, Valakos & Mylonas 1992, Valakos et al. 2007). Angeliki Xyda, former faculty (now retired), con- ducted studies on the ecology of Laudakia stellio (e.g. 1986). Professor Mylonas moved to the University of Crete at 1992 and set the basis for a new herpetological nucleus. Three dissertations have been completed so far whereas more PhD candidates are still working on their theses. Pet- ros Lymberakis, curator of vertebrates at the Natural His- tory Museum of Crete (belonging to the University of Crete) deals with numerous aspects of herpetology (e.g. Lymberakis et al. 2007, 2008). Nikos Poulakakis (assis- tant professor) has worked extensively on the reconstruc- tion of the phylogenetic histories of various amphibians and reptiles (e.g. Poulakakis et al. 2003, 2005a, b, 2008). At the Aristotelian University of Thessaloniki the research group of Sofianidou and Kyriakopoulou-Sklavounou be- gan a series of papers on frogs in collaboration with Hans Schneider. Fruit of their work, based on bioacoustics, was the description of a new species (Pelophylax epeiroticus — 1984). Sofianidou (now retired) supervised two disser- tations and worked mainly with amphibians (e.g. Sofian- idou & Kyriakopoulou-Sklavounou 1983, Sofianidou 1996). She was also one of the editors and main contrib- utors to the Atlas of Amphibians and Reptiles in Europe (Gasc et al. 1997). Kyriakopoulou-Sklavounou (associate professor) supervised one PhD thesis while studying life- history traits and genetic differentiation of Greek frogs (e.g. Kyriakopoulou-Sklavounou 1992, Kyriakopoulou- Sklavounou et al. 2000, 2003). Nikos Loumbourdis (pro- fessor) studies the metabolism and overall physiology of amphibians and reptiles (e.g. Loumbourdis & Hailey 1985, Loumbourdis 1997, 2005, 2007). Besides the aforementioned foundations the Goulandris Natural History Museum hosts a group of active herpetol- ogists: Dimitropoulos who contributed many new locali- ties for reptiles (e.g. 1986, 1990), Ioannides wrote on the herpetofaunas of numerous areas and also on the ecolo- gy of reptiles (e.g. Ioannides et al. 1994, Ioannides & Bousbouras 1997) and Maria Dimaki, who has focused on chameleons (e.g. Dimaki et al. 2000a,b). Panayiota Maragou of the WWF Hellas studies the ecology of lac- ertids endemic to Peloponnese (e.g. Maragou et al. 1996, 1999), while Chloe Adamopoulou (Zoological Museum Bonn zoological Bulletin 57 (2): 329-345 of the University of Athens) is emphasizing on Podarcis milensis (e.g. Adamopoulou et al. 1999, Adamopoulou & Valakos 2005). Dimitris Margaritoulis of Archelon did an important work on the conservation of sea turtles (e.g. Margaritoulis et al. 1986, Margaritoulis 2005). US based Greek Johannes Foufopoulos (assistant profes- sor, University of Michigan) is investigating the evolution and physiological adaptations of lizards on islands of Aegean Sea (e.g. Foufopoulos 1997, Foufopoulos & Ives 1999) in close collaboration with herpetologists in Greece. Recently two more members of the Greek her- petological community became faculty: Konstantinos Sotiropoulos (University of Ioannina, lecturer) who stud- ies genetic differentiation and phylogenetic relations in newts (e.g. Sotiropoulos et al. 2001, 2008a,b, 2009) and Panayiotis Pafilis (University of Athens, assistant profes- sor), focusing on functional ecology and conservation physiology of lacertids (e.g. Pafilis et al. 2005, 2007, 2008, 2009). The increasing number of people involved in herpetolog- ical studies in Greece is also reflected in the organization of three Congresses: the First (1992) and the Sixth (2008) Symposia on the Lacertids of the Mediterranean Basin (both held in Lesvos Island) and the 10‘ Ordinary Gen- eral Meeting of SEH in Crete (1999). Some of the con- tributions presented during the last were published in a volume under the general title Hereptologia Candiana (Lymberakis et al. 2001). The threatened species of the Greek herpetofauna have been recorded in the two editions of the Red Data Book of threatened species of Greece. In the first edition (Karandinos & Paraschi 1992) eight species (seven rep- tiles and one salamander) are listed as threatened while in the second edition (Legakis & Maragou 2009) twelve reptiles and six amphibians are characterized as critical- ly endangered, endangered or vulnerable. An important step in the history of Greek Herpetology was the foundation of the Hellenic Herpetological Society (So- cietas Herpetologica Hellenica, EAdnvucy Epretodoyuxy, Etatpe/a— http://www.elerpe.org) in 2000. The members of the Society are scientists who are involved in the study of amphibians and reptiles but also amateur herpetologists (as non-full members) who are interested in conservation and natural history. At this point Arche/on, the Sea Tur- tle Protection Society of Greece (http://www.archelon.gr), should be mentioned as well. Thanks to the efforts of this pioneer group (founded on early 1980’s), Greek public opinion was sensitized towards the conservation of Caretta caretta. ©ZFMK A brief history of Greek herpetology 337/ PAST, PRESENT AND FUTURE OF GREEK HERPETOLOGY Unlike herpetologies of other European countries, herpeto- logical publications in Greece up to the 1970’s dealt with systematics, focusing on the discovery and description of new species. The majority of studies concerned the dis- tribution of various taxa and the description of local her- petofaunas, with emphasis on the islands. Nonetheless, during the last decade, herpetologists are covering success- fully a wide spectrum of biological aspects including mo- lecular biology, genetic differentiation, environmental physiology, functional ecology, immunology and the over- all picture has been reversed (Fig. 1). 120 - | eee 1880- 1900- 1960- 1970- 1900 1960 1970 1980 1980- 1990 1990- 2000 2000- 2010 Fig. 1. Histogram of publications (total 606 papers) made by foreign (light bar) and Greek herpetologists (spotted bar). The number of species inhabiting Greece has been raised throughout the years (see Appendix I). During the first two decades of 19th century only the species described by the classical taxonomists (Linnaeus, Laurenti and Pallas) were known from Greece. The French Morea Expendition led to the description of the first endemic species and since then the study of the Greek herpetofauna became method- ical and continuous. New species are rather rare and their description is based on cutting edge technology tools, like molecular inference, paired though with typical anatom- ical-morphological studies (e.g. Beerli et al. 1994). This is also the case for the recently described lacertids Podar- cis cretensis and P. levendis, the first Greek species that were published by an exclusively Greek group (Lymber- akis et al. 2008). Until today foreign researchers were publishing most of the papers on the Greek herpetofauna. However this trend has changed during the last 20 years and today the scien- tific work of Greek herpetologists has yielded a continu- ously growing number of papers (Fig. 2). It is important to mention that most Greek researchers are working in col- laboration with colleagues from Europe and North Amer- Bonn zoological Bulletin 57 (2): 329-345 ica, keeping alive the international interest for the Greek herpetofauna and exchanging ideas and methods. Habitat degradation, environmental pollution, introduced species and the non-stop, greedy development of tourism (principal source of money for Greek economy) stress the imperious need for conservation studies in the immediate future. Though knowledge of species distribution is in sat- isfactory level, the evaluation of populations’ status 1s still very poor. Amphibian and fresh water turtle populations are known to decrease as a consequence of water pollu- tion and the desiccation of water bodies. The problem is much more intense in the islands because of tourism-re- lated activities (excessive withdrawal of groundwater and construction projects on wetlands areas). Reptile popula- tions are threatened by wildfires that the last 10 years de- stroyed a significant part of Greek forests and also, in the case of small islets, by overgrazing. Greece hosts some very important nesting beaches for Caretta caretta, endan- gered as well by tourism and fishing. In order to protect and maintain one of the richest European herpetofaunas special conservation projects should be undertaken short- ly with the contribution of herpetologists from all fields. 120 100 80 60 40 everwelN ian a we ilies 1 1880- 1900- 1960- 1970- 1980- 1990- 2000- 1900 1960 1970 1980 1990 2000 2010 Fig. 2. Chart of publications concerning systematics and dis- tribution (light bar) and non-systematic and distributional records (spotted bar) of a total of 606 papers. REFERENCES Adamopoulou C (1999) Structure and composition of animal ground communities with an emphasis on the lizard Podar- cis milensis (Sauria: Lacertidae), in island ecosystems of the Aegean Sea. Dissertation in Greek. 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Comparative Biochemistry and Physiology (A) 147: 1-10 Valakos ED, Pafilis P, Sotiropoulos K, Lymberakis P, Marangou P, Foufopoulos J (2008) Reptiles and Amphibians of Greece, Chimaira Publications, Frankfurt am Mainz Vassara E (1999) Intrapopulation study of the frog Bombina va- riegata (Linnaeus 1758) in Greece. Dissertation in Greek. Uni- versity of Thessaloniki Werner F (1894) Die Reptilien- und Batrachierfauna der Ioni- schen Inseln. Verhandlungen der Zoologisch-Botanischen Ge- sellschaft in Wien 44: 225-237 Werner F (1912) Beitraege zur Kenntnis der Reptilien und Am- phibien Griechenlands. Archive fuer Naturgeschichte 78: 167— 180 Werner F (1927) Beitrage zur Kenntnis den Fauna Griechen- lands. Zoologische Anzeiger LXX, pp. 135-143 Werner F (1930) Contribution to the knowledge of the reptiles and amphibians of Greece, especially the Aegean islands. Oc- casional papers of the University of Michigan Museum of Zo- ology 211: 1-47 Werner F (1933) Ergebnisse einer zoologischen Studien- und Sammelreise nach den Inseln des Aegaeischen Meeres. I. Rep- tilien und Amphibien. Sitzungsberichte der Mathematisch-Na- turwissenschafftlichen Klasse-Abteilung B. 143: 105-131 Werner F (1937) Beitraege zur Kenntnis der Tierwelt des Pelo- ponnes, der Inseln Kythira und Euboea sowie der kleinen In- seln im Saronischen Golf. Sitzungsberichte der Mathematisch- Naturwissenschaftlichen Klasse Abteilung B. 146: 135—153 Werner F (1938) Ergebnisse der achten zoologischen For- schungsreise nach Griechenland (Euboea, Tinos, Skiathos, Thasos usw.). Sitzungsberichte der Mathematisch-Naturwis- senschafftlichen Klasse Abteilung B. 147: 151-173 Wettstein O v (1931) Herpetologie der Insel Kreta. Annalen des Naturhistorischen Museums Wien 45: 159-172 Wettstein O v (1953) Herpetologia Aegaea. Sitzungsberichte der Mathematischen-Naturwissenschafftlichen Klasse Abteilung B. 162: 651-833 Willemsen RE (1991) Differences in thermoregulation between Testudo hermanni and Testudo marginata and their ecologi- cal significance. Herpetological Journal 1: 559-567 ©ZFMK A brief history of Greek herpetology 343 Willemsen RE (1999) Variation of adult body size of the tortoise Testudo hermanni in Greece: proximate and ultimate causes. Journal of Zoology 248: 379-396 Willemsen RE, Hailey A (2002) Body mass condition in Greek tortoises: regional and interspecific variation. Herpetological Journal 12: 105-114 Xyda A (1983) Differentiation between the local Greek and Cypriot populations of Agama stellio (Reptilia: Agamidae). Dissertation in Greek. University of Athens APPENDIX Xyda A (1986) Supplementary evidence on the biometry and ecology of the lizard Stellio stellio of Greece and Cyprus. Bi- ologia Gallo-Hellenica 12: 451-458 Zavattari E (1929) Ricerche faunistiche nelle Isole Italiane dell’ Egeo. Anfibi e Rettili. Archivo Zoologico Italiano 12/13: 161-166 Received: 30.VII.2010 Accepted: 31.VII.2010 Table 1. List of species known from Greece with distribution and date of original description. Date Species Author Range Group 1758 Anguis fragilis Linnaeus Mainland Greece, Thassos and Corfu islands —_ Rept: Anguidae 1758 Bombina variegata Linnaeus Mainland Greece, not in Peloponnese Amph: Discoglossidae 1758 Bufo bufo Linnaeus Mainland Greece and large Aegean islands Amph: Bufonidae 1758 Caretta caretta Linnaeus All Greek seas Rept: Chelontidae 1758 Chamaeleo chamaeleon Linnaeus Chios and Samos Islands Rept: Chamaeleonidae 1758 Chelonia mydas Linnaeus All Greek seas Rept: Cheloniidae 1758 Dolichophis jugularis Linnaeus Islands of southeastern Aegean Sea Rept: Colubridae 1758 Emys orbicularis Linnaeus Mainland Greece, Samos, Samothraki, Rept: Emydidae Kos, Lesvos and Evvoia islands 1758 Eryx jaculus Linnaeus Throughout the country excluding Crete Rept: Boidae 1758 Hemidactylus turcicus Linnaeus Throughout the country Rept: Gekkonidae 1758 Hyla arborea Linnaeus Mainland Greece and large islands Amph: Hylidae 1758 Lacerta agilis Linnaeus Northern borders in high elevations Rept: Lacertidae 1758 Laudakia stellio Linnaeus Only European population Rept: Agamidae Eastern Aegean Sea Islands, Corfu, Thessaloniki and central Cyclades 1758 Lissotriton vulgaris Linnaeus Mainland Greece and large Ionian islands Amph: Salamandridae 1758 Natrix natrix Linnaeus Throughout the country excluding Crete Rept: Colubridae 1758 Rana temporaria Linnaeus Northern borders with Bulgaria Amph: Ranidae 1758 Salamandra salamandra Linnaeus Mainland Greece Amph: Salamandridae 1758 Tarentola mauritanica Linnaeus Western Peloponnese, Crete and Ionian Islands Rept: Gekkonidae 1758 Testudo graeca Linnaeus Mainland Greece and many islands Rept: Testudinidae 1758 Trionyx triunguis Forsskal Introduced, Kos island Rept: Trionychidae 1758 Trachylepis auratus Linnaeus Rhodes, Kos, Symi and Samos islands Rept: Scincidae 1758 Vipera ammodytes Linnaeus Throughout the country excluding Crete, Rept: Viperidae Milos Archipelago and eastern Aegean Sea islands 1758 Vipera berus Linnaeus Macedonia and Thrace in high elevations Rept: Viperidae 1758 Zamenis situlus Linnaeus Throughout the country Rept: Colubridae 1761 Bombina bombina Linnaeus Borders with Bulgaria, River Evros Amph: Discoglossidae 1761 Dermochelys coriacea Vandelli All Greek seas Rept: Dermochelyidae 1768 Chamaeleo africanus Laurenti Only European population, Rept: Chamaeleonidae a restricted zone in southeastern Peloponnese 1768 Coronella austriaca Laurenti Epirus, Macedonia, Thrace, Thassos Rept: Colubridae and Samothraki islands 1768 Hierophis gemonensis Laurenti Throughout mainland Greece Rept: Colubridae excluding Macedonia and Epirus, Ionian islands and Crete Bonn zoological Bulletin 57 (2): 329-345 ©ZFMK 344 Panayiotis Pafilis Date Species Author Range Group 1768 Lacerta viridis Laurenti Mainland Greece excluding Peloponnese Rept: Lacertidae 1768 Mesotriton alpestris Laurenti Mainland Greece Amph: Salamandridae 1768 Natrix tessellata Laurenti Throughout the mainland country, Rept: Colubridae Crete and some Aegean and Ionian islands 1768 Podarcis muralis Laurenti Throughout mainland Greece Rept: Lacertidae and Thassos island 1768 Pseudepidalea viridis Laurenti Mainland and insular Greece Amph: Bufonidae 1768 Triturus carnifex Laurenti Epirus, Macedonia and Corfu island Amph: Salamandridae 1768 Zamenis longissimus Laurenti Throughout the mainland country, Rept: Colubridae Corfu and Paxoi islands 1774 Pelophylax ridibundus Pallas Eastern Macedonia and Thrace Amph: Ranidae 1775 Chalcides ocellatus Forsskal Attica and close islands, Rept: Scincidae Crete, eastern Peloponnese 1775 Pseudopus apodus Pallas Mainland Greece and in many large islands Rept: Anguidae 1789 Dolichophis caspius Gmelin Throughout the country excluding Crete, Rept: Colubridae Rhodes and the majority of Peloponnese 1789 Elaphe quatorlineata Lacepéde Throughout the country excluding Crete Rept: Colubridae and Rhodes 1789 Eurotestudo hermanni Gmelin Mainland Greece, Zakynthos, Cephalonia, Rept: Testudinidae Corfu and Evvoia islands 1789 Hierophis viridiflavus Lacepéde Introduced, Gyaros Island Rept: Colubridae 1789 Platyceps najadum Gmelin Throughout the mainland country Rept: Colubridae and in some Aegean islands 1795 Testudo marginata Schoepff Endemic, Mainland Greece Rept: Testudinidae excluding Thrace and many Aegean islands 1802 Rana catesbeiana Shaw Introduced, Crete Amph: Ranidae 1804 Malpolon monspessulanus Hermann Throughout the country Rept: Colubridae excluding Crete and Cyclades 1814 Elaphe sauromates Pallas Thrace and Thassos island Rept: Colubridae 1814 Podarcis tauricus Pallas Throughout mainland Greece Rept: Lacertidae and Ionian islands 1820 Typhlops vermicularis Merrem Throughout the country excluding Crete Rept: Typhlopidae 1831 Telescopus fallax Fleischmann Throughout the country Rept: Colubridae 1832 Ophisops elegans Ménétriés Only European population, Rept: Lacertidae islands of northeastern Aegean Sea 1833 Ablepharus kitaibelii Bibron & Bory Throughout the country Rept: Scincidae / terra typica in Greece 1833 Algyroides moreoticus Bibron & Bory Endemic, Peloponnese and Rept: Lacertidae few Ionian Islands 1833 Mauremys rivulata Valenciennes Throughout the country Rept: Geoemydidae 1833 Ophiomorus punctatissimus Bibron & Bory Only European population, Rept: Scincidae Peloponnese, Kythira and Kastelorizo islands / terra typica in Greece 1834 Darevskia praticola Evermann Eastern Thrace near river Evros Rept: Lacertidae 1834 Hemorrhois nummifer Reuss Only European population, Rept: Colubridae Islands of southeastern Aegean Sea 1833 Podarcis peloponnesiacus Bibron & Bory Endemic, Peloponnese / Rept: Lacertidae terra typica in Greece 1835 Vipera ursinii Bonaparte Central and northern Greece Rept: Viperidae in high elevations 1838 Eirenis modestus Martin Only European population, Rept: Colubridae Thrace and eastern Aegean Sea islands 1839 Algyroides nigropunctatus Duméril & Bibron Western mainland Greece Rept: Lacertidae (excluding Peloponnese) and Ioanian Islands / terra typica in Greece 1840 Rana dalmatina Bonaparte Discontinuous range in mainland Greece Amph: Ranidae Bonn zoological Bulletin 57 (2): 329-345 ©ZFMK A brief history of Greek herpetology 345 Date Species Author Range Group 1849 Montivipera xanthina Gray Only European population, Rept: Viperidae Thrace and eastern Aegean Sea islands 1870 Cyrtopodion kotschyi Steindachner Throughout the country Rept: Gekkonidae 1870 Trituris karelinii Strauch Macedonia and Thrace Amph: Salamandridae 1876 Podarcis erhardii Bedriaga Throughout mainland Greece, Rept: Lacertidae Sporades and Cyclades / terra typica in Greece 1881 Hellenolacerta graeca Bedriaga Endemic, Peloponnese Rept: Lacertidae 1882 Pelophylax bedriagae Camerano River Evros, east Aegean Islands Amph: Ranidae 1882 Podarcis milensis Bedriaga Endemic, Milos Arhipelago Rept: Lacertidae / terra typica in Greece 1884 Blanus strauchi Bedriaga Only European population, Rept: Amphisbaenidae Islands of southeastern Aegean Sea 1886 Lacerta trilineata Bedriaga Throughout the country Rept: Lacertidae / terra typica in Greece 1889 Pelobates syriacus Boettger Localities in northern Greece Amph: Pelobatidae and Peloponnese, Lesvos island 1891 Lyciasalamandra luschani Steindachner Only European population, Amph: Salamandridae Kastellorizo island 1891 Rana graeca Boulenger Mainland Greece Amph: Ranidae 1894 Anguis cephallonicus Werner Endemic, Peloponnese, Rept: Anguidae Ithaca, Cephalonia and Zakynthos islands 1900 Anatololacerta anatolica | Werner Only European population, Samos island Rept: Lacertidae / terra typica in Greece 1904 Anatololacerta oertzeni Werner Only European population, Ikaria, Symi Rept: Lacertidae and Rhodes islands / terra typica in Greece 1930 Podarcis gaigeae Werner Endemic, Skyros Arhipelago Rept: Lacertidae / terra typica in Greece 1935 Macrovipera schweizeri Werner Endemic, Milos Archipelago Rept: Viperidae and Siphnos island / terra typica in Greece 1940 Pelophylax kurtmuelleri Gayda Mainland Greece, Thassos and Amph: Ranidae Zakynthos islands, most Cycladic islands 1963 Lyciasalamandra helverseni Pieper Endemic, Karpathos, Kassos Amph: Salamandridae and Saria islands / terra typica in Greece 1984 Pelophylax epeiroticus Schneider, Western mainland Greece Amph: Ranidae Sofianidou & Kyriakopoulou- Sklavounou 1994 Pelophylax cerigensis typica in Greece 1994 2008 Pelophylax cretensis Podarcis cretensis Beerli, Hotz, & Uzzell Beerli, Hotz, Tunner, Heppich & Uzzell Lymberakis, Poulakakis, Endemic, Karpathos and Tunner, Heppich Endemic, Crete / terra typica in Greece Endemic, Crete / terra typica in Greece Amph: Ranidae Rhodes islands / terra Amph: Ranidae Rept: Lacertidae Kaliontzopoulou, Mylonas & Valakos Lymberakis, Poulakakis, Kaliontzopoulou, Mylonas & Valakos 2008 Endemic, islets Pori and Lagouvardos Rept: Lacertidae close to Antikythira / terra typica in Greece Podarcis levendis Bonn zoological Bulletin 57 (2): 329-345 ©ZFMK | | tal | ik ‘a i i 3) | {| i \} i Hi Hitues Hi Pee FSD | Weith y i | | | | | i | i | il ae fe \ i ti HHI i} i 1} mi Bonn zoological Bulletin Volume 57 Issue 2 pp. 347-357 | Bonn, November 2010 The history of reptiles and amphibians at Frankfurt Zoo Manfred Niekisch Zoo Frankfurt, Bernhard-Grzimek-Allee 1, D-60316 Frankfurt am Main; E-mail: manfred.niekisch@stadt-frankfurt.de Abstract. Reptiles and amphibians were kept in Frankfurt Zoo from the very beginning in 1858. Information on the col- lection is somewhat fragmentary but still sufficient to draw a picture on its development from then until today. Starting with just a few reptile cages in the monkey house, reptiles soon became a major attraction for the visitors, and a special section in the aquarium building was opened for them in 1904. Knowledge about how to keep reptiles and amphibians was still very poor, but evolved steadily, and shortly before World War II Frankfurt Zoo was famous for its impressive collection of herps, especially crocodilians. Completely destroyed in 1944, the zoo re-opened only a few months after the end of war, but it was not until 1957 that the reptile hall on top of the newly erected aquarium building, which now was called “Exotarium”, could be opened. Having undergone a number of improvements and renovations in the last four decades, the Frankfurt Exotarium today has a number of remarkable breeding results and is putting — as the whole zoo —a focus on nature conservation. Key words: Exotarium, terrarium, herpetological collection, breeding success, nature conservation THE START WITH A FEW SPECIES Frankfurt Zoo was opened on 8 August 1858, the second zoo in Germany after Berlin’s. Reptiles and amphibians were exhibited here from the beginning. They were shown in a wing of the monkey house and consisted mainly of European species. The first Frankfurt Zoo guide published in 1860 mentions a few tailed amphibians and lizards, snakes and turtles (see Table 1). That chapter on herps al- so mentions that there were plans to replace the small “rep- tiles cage” and aquarium by a bigger facility. The newts and the salamanders were at that time kept in the aquar- tum together with the Great Loach (Cobitis fossilis). With regard to the salamanders, it was stated that the animals could be found “on the leaves” in the aquarium, but it is not clear what this means exactly. Were they presented on leaves floating on the water surface? Unfortunately, aside from this first small “inventory” of the reptiles and amphibians kept in Frankfurt Zoo, a sys- tematic list was started only in the 1950s. Daily reports contained information on new acquisitions, deaths and births, and an electronic register for herps has been start- ed only recently. So there is no detailed, continuous doc- umentation of the herpetological species kept here from the beginning until today. Substantial information on rep- tiles and amphibians in the Frankfurt collection is Bonn zoological Bulletin 57 (2): 347-357 scattered over a wide range of articles, zoo guides, and annual reports, allowing one to gain an overall picture with limited, but nevertheless interesting data. Despite the growing importance and attractiveness of Frankfurt Zoo’s herpetological section, not much attention was paid to it in the zoo publications, as will also be shown later. For example, the book published on the occasion of the 100th anniversary of Frankfurt Zoo (Zoologischer Garten der Stadt Frankfurt am Main 1958) has no picture of a reptile or amphibian and only one — rather unimpor- tant — view of the interior of the reptile building from 1957-58. THE FOUNDING OF THE “DEFINITIVE” ZOO This first zoo was a huge success as it awoke much inter- est among the citizens of Frankfurt, but the terrain at the “Leers’scher Garten” was small and could be rented for only ten years. As a consequence, the founders of the zoo decided to find a new location for a bigger and “defini- tive” zoo. In 1865, the Frankfurt Zoological Society and the Senate of the City of Frankfurt signed a contract to establish a new Zoo at the “Pfingstweide”, (then) outside ©ZFMK 348 Manfred Niekisch Table 1. Herpetological species as mentioned in the first guide to Frankfurt Zoo (Weinland 1860). Scientific name* Common name Remarks (translated from German) (translated from German) Lacerta viridis Pseudopus Pallasii (scheltopusik, horned serpent) Tropidonotus natrix var. bilineata Coronella laevis Testudo graeca Testudo polyphemus Glass snake Ringed snake Smooth snake Greek tortoise Gopher tortoise Triton cristatus, igneus, taeniatus Salamandra maculata European green lizard our specimens come from Vienna eats grass, outside during the summer on the flamingo meadow during the summer Our German water salamanders Common European salamander on the leaves * Note: Scientific names in the whole article, when in quotes, and in this table are given as they are mentioned in the respective publication and have not been transferred into modern nomenclature. the City of Frankfurt. Reptiles and fishes had turned out to be a real attraction for the visitors, and so, from the be- ginning, the plan for the new zoo included designs for a herpetological exhibition and aquaria. Due to a number of complications and especially as a consequence of the Fig. 1. The ,,romantic Aquarium tower of Frankfurt Zoo abo- ve the lake in 1880. Bonn zoological Bulletin 57 (2): 347-357 wars between Prussia and Austria (1866) and France and Germany (1870-71), this contract never materialized and the zoo remained — with an extension of the old contract —at the “Leers’scher Garten” for a few more years. Fi- nally, under a new contract, the new zoo was opened at the Pfingstweide on 29 March 1874, at the same place that had already been envisaged before the wars. Frankfurt Zoo has remained at this site until today. Thanks to an initia- tive of Bernhard Grzimek immediately after the Second World War it was enlarged and now covers eleven hectares. The city of Frankfurt has grown around it, so to- day Frankfurt Zoo is in a central location. SLOW START FOR THE TERRARIUM SECTION Despite all the plans and good intentions, the construction of a number of enclosures for mammals and birds and, in particular, a new aquarium and terrarium building had to be postponed due to financial and other constraints once the zoo had moved to its new destination in 1874. But at least there was substantial planning, and the knowledge about how to keep fish and herps as well as the develop- ment of technical means was rapidly increasing right throughout that period. This is also indicated by the fast growing number of associations of aquarium and terrari- um hobbyists in Germany in the last two decades of the 19th century. To provide the financial means for the aquarium building, the members of the administrative council and supervi- sory board provided a loan of 50,000 Reichsmarks. In 1877, the building, comprising two freshwater and 12 sea- water aquaria and (as far as is known) a few terraria was ©ZFMK Herpetology at Frankfurt Zoo 349 finally able to be opened. The building was placed inside an artificial hill, so the walls were insulated and the tem- perature could be kept relatively constant. The issue of ma- jor concern and of utmost importance for the aquarium section, namely water, was solved by erecting a tower with water tanks inside filled with ground water. This simple technique, based on gravity, is still functioning today, guar- anteeing the constant and uniform flow of water into the filters and aquaria. The tower was made to look like an old castle or ruin, and, together with the hill it stands on and the neighbouring lake, it catered nicely to the romantic taste of that time (Fig: 1): An extra entrance fee was charged for the aquarium in or- der to pay the loan back. So the aquarium had its own en- trance fee, and only in 1992 was this practice abandoned and no extra fee was charged any more for visiting the aquarium. DIFFICULT TIMES, BUT A STEADY INCREASE IN EXPERIENCE The herpetological section had been planned as the sec- ond storey of the aquarium building and could only be built later. It finally opened on 15 May 1904. In the ten years before that, the reptiles seem to have had a rather difficult life at Frankfurt Zoo. Especially in winter, many of these animals died because of the poor conditions they were kept in. It was Wilhelm Haacke, director of the zoo from 1888 to 1893, in particular, who expanded the rep- tile collection. During the summer months, when the mon- keys were kept in outside enclosures, he used the mon- key house to put boxes with reptiles and amphibians on exhibit. As can be read in the 1895 zoo guide, the “col- lection that was outstanding because of its richness ... [was] usually set up in May, temperatures allowing, and remained there until October, a few boxes (containing the giant snakes and bigger lizards) even remaining on show during the winter”. For that latter purpose, a heated plat- form (“Warmetisch”’) had been built in 1891. Fig. 2. Bonn zoological Bulletin 57 (2): 347-357 In the newly opened “Reptile Hall” sunlight was seen an important factor for the well being of reptiles (1904). OZFMK 350 Manfred Niekisch 5 Panag) 42 ae . Fig. 3. It is plain that mortality was high. The 1895 zoo guide (the first to be published after ten years!) lists, however, an im- pressive number of species — or to be more precise: two crocodiles, eleven turtles, 17 snakes, 22 lizards and ten Fig. 4. (1912). A big specimen of Python reticulatus in its terrarium Bonn zoological Bulletin 57 (2): 347-357 San Rit “< we, LS W 7h Mixed life in a tropical jungle environment: Tiliqua rugosa (?), Cordylus giganteus and Macroscincus coctei (1912). anuran species and three species of urodela. A few of the comments and specimens are truly remarkable. So “thanks to the goodness of Mr Schmacker from Shang- hai”, the collection contained “the first specimen of the Chinese alligator brought live to Europe” and “two giant Aldabra turtles (7estudo elephantina)’. With regard to the latter, the 1895 zoo guide states that “only a few decades will pass until this turtle of such incredible dimensions will have become extinct”. “One of the rarest species at the Zo- ological Garden, the snake-necked turtle, Hydraspis hi- laire’’”’, was kept together with Chelydra serpentina. “The Sinai lizard (Uromastix ornatus)” is described as a “very strange animal. It feeds on rose petals in summer and on acacia and lettuce in winter; as soon as the sun shines on its back, it opens certain depressions in the skin and the body assumes a very beautiful colour”. The lizard species ranged from Anguis fragilis to “Silubosaurus stokes” and from “Lacerta muralis” to “Tiliqua gigas”. Aside from alpine salamander, Japanese giant salamander, bull frog and Leptodactylus, all amphibians kept at that time were species that occurred wild in the Frankfurt area. ©ZFMK Herpetology at Frankfurt Zoo 351 Fig. 5. (1912). Tropical jungle landscape made of aquaria and plants Haake (who by the way had been vividly recommended by Ernst Haeckel for the post of zoo director) had a well- developed collector’s mentality but his attempt to estab- lish systematic collections of birds and herps did not re- ceive much of a positive response from visitors to the zoo, and he quit the job in 1893. His immediate successor, Adalbert Seitz, then started to develop a completely new concept for a reptile exhibit, recognizing that these ani- mals needed sunlight. A glasshouse called the “reptile hall” was erected on top of the aquarium building and inaugu- rated on 15 May 1904 (Fig. 2). In the venomous snakes section, increased security measures were introduced in the year 1906 to offer the keepers better protection. The number of species had by then risen to twelve turtles, 28 snakes, 25 lizards and 13 amphibians. Evidently there was some “fluctuation” in the crocodile species, as the 1905 Z00 guide states: “Mostly different species such as the al- ligator, Nile crocodile, dwarf crocodile etc. are on exhib- it”. The Chlamydosaurus kingi kept in the collection is said to be the first specimen “to have reached the European continent alive”. In 1907, even before becoming zoo di- rector in 1908, the then zoo assistant, Kurt Priemel, start- ed changing the concept again. He wanted to show the vis- itors the diversity of life, abandoning the approach of sys- tematic collections. He built a second glasshouse next to the first one, added to the reptile hall 40 aquaria for trop- ical fish (Fig. 5) and a tropical wetland area for crocodiles as well as big terraria for giant snakes (Fig. 4) and turned the reptile house, together with the aquarium, into the “biggest and most diverse of all such institutions on the continent” (Scherpner 1983). As visitors to the zoo had to pay an additional entrance fee for the aquarium and ter- rarium building, visitor numbers could be easily moni- tored. The new and enlarged building attracted more than 80,000 people every year. Bonn zoological Bulletin 57 (2): 347-357 The zoo did not suffer any major physical damage dur- ing the First World War, but the economically difficult post-war era obliged director Priemel to be creative. He made an interesting contract with the animal catcher and dealer John Hagenbeck. Frankfurt hosted reptiles import- ed by Hagenbeck and, in exchange, got the pre-emption rights and a reduced price on the specimens Priemel want- ed to buy. It is reported that visitors were quite astonished by a sign saying “for sale” on a big container full of gi- ant snakes which “none of the visitors managed to count” (Scherpner 1983). Soon the reptile collection had reached an impressive di- mension, and its increasing importance is also document- ed by a number of articles on it which were published in the Zoo’s own “Mitteilungen aus dem Frankfurter Zoo” and elsewhere. One of the authors is Robert Mertens who did evidently have a close relationship with the zoo, since he authored seven papers between 1921 and 1925 specif- ically about the species and specimens kept at Frankfurt Zoo, and more precisely on the freshwater turtles (1921), giant snakes (1921, 1924 — the latter one not mentioned by Schirner 1977), venomous snakes (1925), news from the reptile house (1922), new animals (1922) and on Cer- atophrys ornata (1922). An exhaustive paper on the whole collection of reptiles by Richard Wieschke (1925) gives, like the articles just mentioned, short notes and comments on the different species shown and mentions, as a special attraction, a giant salamander from Japan which was ex- hibited in an aquarium in the lower basement. From that description of the collection one can deduce that, in 1925, there were more than 40 snake species, 28 lizard species, more than 20 turtles and tortoises (including a loggerhead sea turtle), and seven crocodile species on exhibit. An in- teresting detail is the mentioning of a female reticulated python 8 m in length which, after having undergone “dif- ficult surgery”, had not taken any food for 16 months be- fore she finally accepted a piglet. Six anuran and 2 urode- lan species are specifically mentioned as part of the col- lection “plus the numerous European frogs, toads, sala- manders and newts”. Among the amphibians mentioned are “two giant bull frogs..., and, even more impressive, two South American horned frogs”, as well as African clawed frogs “which because of their hopping movements under water soon got the name ‘water monkeys’”, besides Pipas, a Japanese giant salamander and a Proteus. Whereas Robert Mertens 1s well known among herpetol- ogists even today and does not need to be introduced to the reader, a few words have to be said about Wieschke. In one of the articles, his name is given as “Fritz”, in the other article as “Rich[ard]’”. It was not possible to find out if Fritz and Rich[ard] Wieschke were the same person — and “Fritz” a printing error? — or whether and how they ©ZFMK 352 Manfred Niekisch were related. “Fritz” could not be identified at all, where- as it is known that Richard was a volunteer assistant, help- ing out quite actively with many activities in the zoo such as the administration of the library, keeping the register of animals and observing them. He published several small papers in the Mitteilungen, the last one appearing in No- vember 1928, and, as far as is known, died at the age of 23 ani l929: A remark in a review authored by zoo director Priemel on issues | and 2 of Wilhelm Klingelhoffer’s Terrarienkunde in the Mitteilungen from May 1925 gives an interesting insight into the concept of reptile keeping. Priemel wel- comes Klingelhoffer’s approach to arranging the contents of the terraria in such a way that they resemble the habi- tat of the species in the wild. Furthermore, he writes that terraria for schools should always be arranged so that they imitate nature, but then he goes on to write: “Unfortunate- ly, the containers in public exhibitions cannot follow this principle, as so many inhabitants must be kept in them in order that visitors can observe the major part of them at any time of the day”. on “ea Fig. 6. the nght foreground. Bonn zoological Bulletin 57 (2): 347-357 In the following years, the collection continued to grow and attracted more and more visitors. Various publications talk about anacondas, chameleons, Gila monsters and sea turtles all becoming part of the collection and, of course, about “Komo”, the tame Komodo monitor lizard which came to Frankfurt in 1927, only 15 years after this species had been discovered. It was brought from Komodo to Frankfurt by Robert Mertens. Frankfurt already had some experience in taming monitor lizards. Two Varanus sal- vator had come to Frankfurt Zoo in 1922—23, when they were just 25 cm long. They were quite aggressive and, while one died, the other one had grown to a length of 1.35 m by 1926 and thanks to “persistent, gentle, careful treatment and care” had become tame. Whenever the door to his enclosure was opened or his name (“Bubchen’”’, lit- tle boy) was called, he climbed onto the keeper’s shoul- der and allowed himself to be carried around (Fig. 7). In that time before World War II, discussions arose about the rights and wrongs and justifications of keeping ani- mals “in captivity”, and, as one of the arguments in favour of zoos, curator Gustav Lederer (1937) published infor- mation about the longevity of reptiles at Frankfurt Zoo. Different species, different sizes: A look into the “world renown” crocodile collection in 1925. Note the Macroclemys in ©ZFMK Herpetology at Frankfurt Zoo 353 —— Fig. 7. Varauns salvator “Bubchen” and his keeper — in the truest sense of the word! — in 1929. In that year, the zoo was home to an Alligator mississip- piensis, a Trionyx triunguis and two Heloderma suspec- tum that had been living at the zoo since 1905, 1912 and 1927 respectively, as well as a Chinese alligator. The lat- ter had moved to Frankfurt in 1910 when the Berlin Aquar- tum had to be closed because of financial problems and Frankfurt took over its entire reptile collection. This alli- gator had come to the Berlin Aquarium in 1886 and so had lived for 30 years in Frankfurt when he died in 1940. The already highly diverse collection of crocodiles was enriched by a Jomistoma schlegeli in 1937, raising the number of species kept to eight. In addition, young and old, small and large individuals were all kept together (Fig. 6). This crocodile collection is repeatedly referred to as a major attraction and as “world renowned” (Lederer 1937), but, looking at it today, it certainly must be regarded as highly problematic from a zoological point of view as well as from the aspects of animal welfare. Wieschke (1927) mentions, for example, that the “newly created tropical swamp area for crocodiles” is host to “numerous species, among others a large number of American alligators (A/- ligator mississippiensis), one of the few surviving speci- mens of the Chinese alligator, Nile crocodiles, saltwater crocodiles and African dwarf crocodiles” — each in the plu- Bonn zoological Bulletin 57 (2): 347-357 ral! A completely new approach to keeping crocodiles was only introduced around 1975, when this collection was fi- nally dissolved to create a larger crocodile enclosure. With the retirement of Kurt Priemel in March 1938, the dynamic development of the herp collection came to an end, as his successor, Georg Steinbacher, was more of a “bird man” and evidently not much interested in reptiles. The first bombs hit Frankfurt Zoo in October 1943, caus- ing some limited damage including to the aquarium build- ing, but most, if not all, of the reptiles survived. The ven- omous snakes, however, now had to be put down for se- curity reasons. A few months later, the disastrous bomb- ing of Frankfurt on 18 March 1944 completely destroyed the zoo and the aquarium with all its animals (Fig. 9). ON THE WAY TO A MODERN ZOO: THE PERIOD AFTER WORLD WAR II The reconstruction of the zoo, under its new director Bern- hard Grzimek, started immediately after the end of the war. The zoo re-opened on | July 1945, offering its visitors a few animals and a lot of entertainment in the form of all kinds of cultural events, circus shows, carousels and so on. The re-building of the aquarium started in 1951 and the shell of a “24 m long tropical swamp area destined for the keeping of crocodiles, turtles and so on” had been com- pleted in 1952. Precisely | chameleon, four snakes, 26 tur- tles and seven crocodiles were housed in the preliminary terraria in 1953. From then on, more and more reptiles and amphibians were acquired by or donated to the zoo. The building with the new big reptile hall was opened official- ly on 27 August 1957 and by the end of that year, it had had 282,084 visitors. There were “giant crocodiles able to kill a human being” and “gigantic land tortoises 200 years of age” living in the (altogether eleven) “climatic landscapes”, with plant arrangements giving the visitor the illusion of being in a tropical jungle. As the “aquarium” was now housing fish and other aquat- ic animals, as well as penguins in an Antarctic environ- ment and a few other birds in the tropical section, along with many reptile and amphibian species, it was decided to give it a new name to better reflect the situation and intention of the building. Since 1954, this building has therefore been known as the “Exotarium’”’. The innovative ideas and plans for the Exotarium were basically devel- oped by Gustav Lederer, who had already been the key person for the “pre-war” aquarium under director Priemel. After the war, he became the zoo’s chief curator. How far- sighted and innovative his thinking was and how careful- ly he observed his animals is reflected, for example, in his paper on the “importance of light in animal keeping” (Le- derer 1927). ©ZFMK 354 Manfred Niekisch soe et wh wt Whee PU} % be Fig. 8. An unidentified keeper working in the terrarium section (1936). At the suggestion of Bernhard Grzimek and in recogni- tion of his merits and the quality of his scientific publi- cations, he received an honorary doctorate from Frankfurt University in 1953. He retired on 30 September 1958, af- ter having served Frankfurt Zoo for 45 years, accompa- nying it through two world wars and all its ups and downs. Gustav Lederer died at the age of 69 on 13 February 1962. THE EXOTARIUM TODAY Despite all changes, improvements and renovation activ- ities in the 1980s and 1990s, the concept of the herpeto- logical section of the Frankfurt Exotarium until today es- sentially goes back to Gustav Lederer. He was followed by curator Dieter Backhaus who, in 1973, handed over to Hartmut Wilke. It was still a time of much “trial and er- ror’, since knowledge regarding the keeping of reptiles was still limited. In 1960, a few adult and juvenile spec- imens of Amblyrhynchus cristatus were even exhibited, but did not survive the first two years. Reptiles and amphibians were selected for their “didac- tic, zoogeographical and ecological aspects” and the zoo “dispensed with animals which were always hiding away during opening hours”. Backhaus, as well as his succes- Fig. 9. | View of the reptile hall after the bombing in 1944. Bonn zoological Bulletin 57 (2): 347-357 ©ZFMK Herpetology at Frankfurt Zoo 355 Fig. 10. The Exotarium Tower overlooking the “seal cliffs” (2008). Photograph: Sabine Binger. sor, constantly tried to improve the living conditions of the animals, trying out all sorts of lamps, heating equip- ment and other means to improve the climate control of the terraria. They also did lots of work on nutrition and disease prevention and carried out the associated physi- cal changes and improvements to the building, such as spe- cial rooms to prepare food, raise foraging animals and raise newly born reptiles and amphibians. Terraria were equipped with appropriate soil substrate for digging species as well as stones and trees for climbing species, in addition to hiding places and other structures, paving the way to modern reptile keeping. Another remarkable change was the renovation of the crocodile enclosure which had made it necessary to give up the crocodile col- lection in around 1975 and to send the gharial (which had been living at Frankfurt Zoo since 1958) to the Gharial Breeding Centre in Orissa, India in 1979. After the reno- vation of the building had been completed, Nile crocodiles returned to the zoo in 1977, but the enclosure turned out to be unsuitable for that aggressive species. Finally, in 1990, Frankfurt Zoo started keeping Australian freshwater crocodiles, which started breeding regularly in 1994 and still do so today. This is one of the many breed- ing successes at the Frankfurt Exotarium since Rudolf Wicker became its curator in 1984. He took over at a time when again some necessary renovation work had started, and so the opportunity to build a big landscape terrartum Bonn zoological Bulletin 57 (2): 347-357 for freshwater tortoises (1987) was seized. The group of Cyclura cornuta then consisted of shy and aggressive an- imals. They had come to the zoo in 1974, but Wicker re- placed them by ten new animals imported from the zoo in Santo Domingo. These animals laid eggs for the first time in 1987, but the keeping facility was not the most favourable in many aspects, and there was little breeding success. Just a few weeks after they had been moved to anewly built enclosure in 1991, they started breeding suc- cessfully and have done so ever since. Other remarkable breeding successes of the last two decades have been the Phelsuma klemmeri from Mada- gascar, Varanus salvator cumingi and Erymnochelys madagascariensis, as well as the Laemanctus serratus and Petrosaurus thalassianus that were all bred in Frankfurt for the first time ever under human care. Our specimens of Crocodylus johnsoni form the only breeding group out- side Australia, and Frankfurt Zoo keeps and breeds Ctenosaura bakeri, the highly endangered iguana from Utila island. Especially as Frankfurt Zoo has made nature conservation in situ and ex situ one of its top priorities, its close coop- eration with the Customs Service at Frankfurt Airport must also be noted. Every year, this results in hundreds of rep- tiles being seized from travellers or commercial shipments at Frankfurt Airport and being brought to the Exotarium OZFMK oS) n on Fig. 11. (2008). Photograph: Sabine Binger. — even rare animals such as a few Psammobates from two different species. Some of the shipments seized contain quite a number of specimens, for example 300 Geoche- lone elegans or more than 70 Cordylus mossambicus and C. rhodesianus and, repeatedly, also large numbers of poi- son arrow frogs. Particularly with regard to the more com- mon species and relatively high numbers of specimens, it is extremely difficult to find appropriate people and in- stitutions willing and able to take them on. All these an- imals are lost to the natural world as they cannot usually be taken back and released into the wild. One exception was the case of five hawksbill turtles (Eretmochelys imbricata) in 2009. Dogs trained to detect CITES species at Frankfurt Airport discovered the eggs in the luggage of a tourist. The eggs were brought to the Exotarium, and, as they looked good, were put into an in- cubator. During the following days, the turtles hatched and were kept in an aquarium until they had reached a length of around 20 cm. As it was known from which beach in the Seychelles they had been collected, they could be sent Bonn zoological Bulletin 57 (2): 347-357 Manfred Niekisch sda aoe. “ : ~ The entrance to the Frankfurt Exotarium still preserves the charm and character of the 1950s when it was rebuilt back and were released into the sea by the local authori- ties. The media attention was huge, and so this success- ful, but quite untypical, story could be accompanied by the message that one should not take home souvenirs of endangered and protected species. Generally, species conservation aspects today play an im- portant role in Frankfurt Zoo, and this, of course, also ap- plies to the Exotarium. This building, with its long and interesting history and its rich collection of reptiles and amphibians, is certainly one of the best places in Frank- furt Zoo to demonstrate to the visitors the multitude of forms, colours, adaptations and other expressions of the diversity of life. Today, there is neither a separate entrance fee nor any counting of the number of visitors to the Ex- otarium, but, in all probability, it may be assumed that al- most all the visitors to Frankfurt Zoo (more than 900,000 per year!) also visit the Exotarium. Showing around 170 adult specimens of 29 amphibian species and more than 400 reptiles from 68 species, it is and remains one of the main attractions of Frankfurt Zoo. ©ZFMK Herpetology at Frankfurt Zoo 357 Sabine Binger. REFERENCES Anonymous (1895) Fiihrer durch den Zoologischen Garten zu Frankfurt am Main. Lederer G (1927) Die Bedeutung des Lichtes in der Tierpflege. Blatter ftir Aquarien- und Terrarienkunde 37: 36-42, 63-64 Lederer G (1937) Merkwiirdigkeiten und Seltenheiten im Frankfurter Tiergarten-Aquarium. Frankfurter Zoo-Zeitung 13: 37-38 Mertens R (1921) Die StiRwasserschildkréten des Frankfurter Zoologischen Gartens. Lacerta 1: 55—56 Mertens R (1921) Die Riesenschlangen des Frankfurter Zoolo- gischen Gartens. Blatter ftir Aquarien- und Terrarienkunde 32: 277-278 Mertens R (1922) Neues aus dem Reptilienhause des Zoologi- schen Gartens in Frankfurt am Main. Naturwissenschaftlicher Beobachter 63: 43-44 Mertens R (1922) Neue Tiere. Naturwissenschaftlicher Beobach- ter 63: 173 Mertens R (1922) Schmuckhornfrésche (Ceratophrys ornata Bell) im Frankfurter Zoologischen Garten. Naturwissenschaft- licher Beobachter 63: 208 Mertens R (1924) Riesenschlangen. Mitteilungen aus dem Frank- furter Zoo 2: 6—7, 10-11 Mertens R (1925) Giftschlangen. Mitteilungen aus dem Frank- furter Zoo 4: 6-11 Bonn zoological Bulletin 57 (2): 347-357 Priemel K (1925) Buchbesprechung “Terrarienkunde”. Mittei- lungen aus dem Frankfurter Zoo 5: 20-21 Scherpner C (1983) Von Burgern ftir Biirger — 125 Jahre Zoo- logischer Garten Frankfurt am Main Schirner E. (1977) Die Veroffentlichungen von Robert Mertens. Courier Forschungsinstitut Senckenberg 20: 12-104. Frank- furt am Main Weinland D F (1860) Fiihrer durch den Zoologischen Garten in Frankfurt am Main. Verlag der Zoologischen Gesellschaft, Frankfurt Wieschke F (1924) Echsen des Frankfurter Aquariums — Mit- teilungen aus dem Frankfurter Zoo 5: 14-16 Wieschke R (1925) Die Reptilienabteilung unseres Zoo. Mittei- lungen aus dem Frankfurter Zoo 2: 12-13, 20-28 Wieschke R (1927) Die Tropische Sumpfanlage ftir Krokodile. Mitteilungen aus dem Frankfurter Zoo 4: 2 Zoologischer Garten der Stadt Frankfurt am Main (Ed.) (1958) Hundertjahriger Zoo in Frankfurt am Main Zoologischer Garten der Stadt Frankfurt am Main (Ed.) (no year) Jahresbericht des Zoologischen Gartens der Stadt Frankfurt a. M. 116-130 fiir 1974-1991. Report of the Zoological Gar- dens [sic!] of Frankfurt 116-130 for 1974-1991 Received: 25.VIII.2010 Accepted: 25.1X.2010 OZFMK 0 Nii Te el Bonn zoological Bulletin Volume 57 Issue 2 pp. 359-366 Bonn, November 2010 Tetramorium boehmei sp. n. — a new ant (Hymenoptera: Formicidae) species from the Kakamega Forest, Western Kenya Francisco Hita Garcia, Georg Fischer, Patrick Kick, Birthe Thormann & Marcell K. Peters Zoological Research Museum Koenig, Adenaueralle 160, D-53113 Bonn, Germany. E-mails: f.hita.zfmk@uni-bonn.de; Georg.Fischer@gmx.de; patrick_kueck@web.de; birthe.thormann.zfmk@uni-bonn.de; m.peters.zfmk@uni-bonn.de Abstract. Tetramorium boehmei Hita Garcia & Fischer sp. n. — a new ant species from the Kakamega Forest in Western Kenya is described. The new species can be placed in the 7etramorium camerunense species group and differs signifi- cantly from the other members of the group by its highly reduced sculpturation on head and mesosoma. With only two available specimens sampled in undisturbed primary forest, Zetramorium boehmei sp. n. seems to be a relatively rare en- demic species. Additionally, a first key to the Tetramorium species groups found in the Kakamega Forest is provided. Keywords. Ants, Kakamega Forest, species group key, taxonomy, 7etramorium, Tetramorium camerunense species group. INTRODUCTION The ant genus 7etramorium Mayr, 1855 is almost global- ly distributed, and with over 430 described species one of the most species-rich genera worldwide (Bolton 1995; B Bolton, Isle of Wight, pers. comm. 2010). The Afrotrop- ical zoogeographic region holds the largest diversity with over 210 listed Tetramorium species (Bolton 1976, 1980, 1985, 1995; Hita Garcia et al. 2010). The Kakamega Forest, one of the last indigenous forests in Kenya, and its animal diversity have received consid- erable scientific attention in the last decades (e.g. Claus- nitzer 1999, 2005; Copeland et al. 2005; Hita Garcia et al. 2009; Kitihne 2008; Schick et al. 2005; Tattersfield et al. 2001; Wagner & Bohme 2007; Zimmermann 1972). Generally, the forest is considered to be the eastern-most relict of the equatorial Guineo-Congolian lowland rain for- est belt (Kokwaro 1988; Wagner et al. 2008; Zimmermann 1972). The strong biogeographic affinities to West and Central African forests can be clearly seen in some fau- nal elements like reptiles, dragonflies, and ants (Claus- nitzer 2005; Hita Garcia et al. 2009; Wagner et al. 2008). The ant fauna proved to be remarkably diverse with 288 species from 52 genera and 11 subfamilies constituting the second highest species richness reported for the Afrotrop- ical zoogeographic region (Hita Garcia et al. 2009). By far the most species-rich genus in Kakamega Forest was Tetramorium with more than 40 species belonging to 14 Bonn zoological Bulletin 57 (2): 359-366 species groups (Hita Garcia et al. 2009; FHG, unpub- lished). The Zetramorium camerunense species group was well represented with four species: Tetramorium lu- cayanum Wheeler, W.M., 1905, Tetramorium cf. gegaimi Forel, 1916, and two undescribed species. Recent taxonomic work was primarily focused on the Tetramorium weitzeckeri species group with the descrip- tion of Tetramorium snellingi Hita Garcia, Fischer & Pe- ters, 2010 and a species group revision for the whole Afrotropical region (FHG, unpublished). However, around 10 species or 25% of the Tetramorium fauna of the Kakamega Forest still remain undescribed. With this work we present a first preliminary key to the 7etramorium species groups present in the Kakamega Forest and de- scribe a new species belonging to the 7’ camerunense species group. MATERIAL AND METHODS The type material has been deposited in the following in- stitutions: NMK: National Museums of Kenya, Nairobi, Kenya ZFMK: Zoological Research Museum Koenig, Bonn, Ger- many ©ZFMK 360 Fancisco Hita Garcia et al. Both, holotype and paratype, were measured with an Olympus SZX 12 stereomicroscope equipped with a dual- axis optical micrometer at a magnification of 90x. The fol- lowing measurements and indices, in parts adapted from Bolton (1980) and Gusten et al. (2006), were used: Head length (HL): maximum distance from the mid-point of the anterior clypeal margin to the mid-point of the oc- cipital margin, measured in full-face view. Head width (HW): width of head directly behind the eyes measured in full-face view. Scape length (SL): maximum scape length excluding basal condyle and neck. Eye length (EL): maximum diameter of compound eye measured in oblique lateral view. Pronotal width (PW): maximum width of pronotum meas- ured in dorsal view. Weber’s length (WL): diagonal length of mesosoma in lat- eral view from the postero-ventral margin of propodeal lobe to the anterior-most point of pronotal slope, exclud- ing the neck. Propodeal spine length (PSL): in dorsocaudad view, the tip of the measured spine, its base, and the centre of the propodeal concavity between the spines must all be in fo- cus. Using a dual-axis micrometer the spine length is measured from the tip of the spine to a virtual point at its base where the spine axis meets orthogonally with a line leading to the median point of the concavity. Petiole length (PTL): maximum length of petiolar node measured in dorsal view. Petiole height (PTH): maximum height of petiolar node measured in lateral view from the highest (median) point of the node to the ventral outline. The measuring line is placed in an orthogonal angle to the ventral outline of the node. Petiole width (PTW): maximum width of petiolar node measured in dorsal view. Postpetiole length (PPL): maximum length of postpetiole measured in dorsal view. Postpetiole height (PPH): maximum height of the post- petiole measured in lateral view from the highest (medi- an) point of the node to the ventral outline. The measur- ing line is placed in an orthogonal angle to the ventral out- line of the node. Bonn zoological Bulletin 57 (2): 359-366 Postpetiole width (PPW): maximum width of postpetiole measured in dorsal view. Ocular index (OI): EL / HW * 100 Cephalic index (CI): HW / HL * 100 Scape index (SI): SL/ HW * 100 Propodeal spine index (PSLI): PSL / HL * 100 Petiolar node index (PeNI): PTW / PW * 100 Lateral petiole index (LPel): PTL / PTH * 100 Dorsal petiole index (DPel): PTW / PTL * 100 Postpetiolar node index (PpNI): PTW / PW * 100 Lateral postpetiole index (LPpI): PPL / PPH * 100 Dorsal postpetiole index (DPpI): PPW / PPL * 100 Postpetiole index (PPI): PPW / PTW * 100 Measurements and indices are presented as minimum and maximum values. Additionally, all measurements are ex- pressed in mm and presented with three decimal places. The digital colour images were produced with a QImag- ing Micropublisher 5.0 RTV camera attached on a LEICA Z6 APO stereo-microscope and mounted with Syn- croscopy Auto-Montage software (version 5.03). The mounted images were processed for publication with Adobe Photoshop CS2 and ImageJ. All images present- ed in this work are also online available at Antweb (Fish- er, 2002). Furthermore, holotype and paratype are unique- ly identified with specimen-level codes (e.g. CASENT0217239) affixed to each pin. Total genomic DNA was extracted from two dissected sin- gle legs of the holotype, using the Qiagen DNeasy®Blood&Tissue Kit, following the manufacturers’ protocol. DNA was eluted with 50 ul buffer AE; this step was repeated once to maximize yield. A ca. 650 bp long fragment of the 5’-region of the cy- tochrome c oxidase subunit I (COI), the standard DNA barcode-marker for animals, was amplified using the primers LCO 1490 and Nancy (5’-GGT CAA CAA ATC ATA AAG ATA TTG G-3’ and 5’-CCC GGT AAA ATT AAA ATA TAA ACT TC -3’; Folmer et al. 1994) and the Qiagen® Multiplex PCR Kit. Amplification reactions were carried out in a 20 pl volume containing 10 ul QIAGEN Multiplex PCR Mastermix, 2 111 Q-Solution, 1.6 pl of each primer (both 10 pmol/ul), and 2.5 tp] DNA template, and filled up to 20 ul with sterile HO. The PCR temperature profile consisted of an initial denaturation at 95° (15 min), followed by 40 cycles at 94° (35 s, denaturation), 48.5° (90 s, annealing), 72° (90 s, extension), and a final exten- sion at 72° (10 min). PCR success was checked by elec- trophoresis on an 1.5% agarose gel containing ethidium bromide. The PCR product was purified using 3 pl of the ExoSAP-IT® PCR purification reagent following the man- ufacturers’ protocol. ©ZFMK A new species of Zetramorium from the Kakamega Forest 36] The sample was bidirectionally sequenced by a commer- cial company (Macrogen Inc., Seoul, Republic of Korea; http://www.macrogen.com) using PCR primers. BLAST search confirmed belonging of the sequence to the genus Tetramorium. The sequence is deposited in GenBank (ac- cession number HM753586). KEY TO THE TETRAMORIUM SPECIES GROUPS FOUND IN KAKAMEGA The following key to species groups is adapted from Bolton (1976, 1980) and specific for the Kakamega For- est, though it also works for Western Kenya in general: 1. Whole body covered with regularly branched hairs, either bifid or trifid, giving the ant a woolly or furry APPS ALAN CE teeter ene aesere oe elaeines. iene Seen ctes 2 — Hairs generally simple, rarely bizarrely modified, but never regularly branched bifid nor trifid as above . 3 2 Antennae 11-segmented; elongate simple hairs pres- ent along the antennal scapes and upper borders of the DROWN CTGTIVEYS, ccsnooseccosasdascocospsecedz0so000 T. ericae group — Antennae 12-segmented; elongate simple hairs absent along the antennal scapes and upper borders of the PO MCaleCARMNAC Heese ceseseee cee eee T. gabonense group Smeemtcmmac lil =seommemte dees esac eccesecesstesccascrsenessstses 4 SP AMLemmMac I 2-SeCmeMte ds a-..-cresccersceecessce-eeeeeseeeeeree 5 4 Petiolar node squamiform to high nodiform, never blocky nodiform with sharply defined angles. paqgacadoodadoOdaed Here reRe nee eee eRe OE Se T. weitzeckeri group — Petiolar node strongly blocky nodiform, generally with sharply defined angles. ... 77 angulinode group 5 Lateral portion of clypeus prominent, raised to a tooth or crest in full-face view; in dorsal view the lateral clypeal portions rise to a high peak in front of the an- tennal insertions and then slope down towards the me- dian part of the clypeus. ............ T. sericeiventre group — Lateral portion of clypeus not modified as above. . 6 6 Antennal scapes very long (always SI > 120); frontal carinae weakly developed and short, at most reach- ing the posterior eye margins. ....... T: aculeatum group — Antennal scapes distinctly shorter than above (always SIO) strontalicarinae wantablet eee ssere-eeeeeeee-c- a 7 Propodeum armed with a pair of small triangular teeth or denticles which at most are as large as the pro podeallObesh eres c eee eee seer eee senses 8 — Propodeum armed with a pair of medium-sized to long spines which are noticeably larger than the propodeallObes: 2s es. vicssscecascn a5. sscasscanckseecess eae hones cose 12 Bonn zoological Bulletin 57 (2): 359-366 8 Anterior clypeal margin with median impression. Bear cence: Metre ssute ae eniny tnaieess T. dumezi group (in parts) = AntentorclypealitmareimCmtines es 2 ; > Santorini ~ x (Thera) = 0 3 ( ioe Crete j whe iS) Fig. 1. | Map with the locations of the Ionian and Aegean is- lands mentioned in the text. is limited in its south-eastern European distributional range to the continental Balkan peninsula. No moles have ever been reported from the eastern Mediterranean islands, with the only exception of the Balkan mole Talpa stankovici V. Martino & E. Martino, 1931 on the islands of Corfu (Niethammer 1962, 1990; KryStufek 1999a) and Cephalonia (Catsadorakis 1985; Giagia-Athanassopoulou 1998; Stamatopoulos, in verbis). Wettstein (1942) observed another species, the crested porcupine Hystrix cristata Linnaeus, 1758, but mentioned that local people referred to its presence on the Eastern Aegean islands of Ikaria and Lesbos. He (Wettstein 1942) added that this might have been the result of confusion with a hedgehog, the today dispersed Northern white-breasted hedgehog Erinaceus roumanicus Barrett-Hamilton, 1900 (Krystufek et al. 2009). Effectively, the Greek term used to indicate the hedgehog is skanzohiros, which means “spiny pig”, which is probably the reason of a confusion with the English “porcupine” (and/or the Italian “porcospino” and the French “porc-épic’’). Moreover, the common porcupine has never been reported from the Balkan peninsula (Masseti et al. in press), while the Indian crested porcupine, Hystrix indica Kerr, 1792 is known from Anatolia with an occurrence further east to the Near East, including Arabia, Kashmir, Nepal and through peninsular India to Sri Lanka (Harrison & Bates 1991). These publications are probably the baseline of several unproven reports. Cheylan (1988) still quoted the occurrence of “Hystrix cristata” (sic) on the Eastern Aegean islands of Rhodes, Ikaria and Lesbos. The occurrence of Microtus subterraneus (de Sélys-Longchamos, 1836) was reported from Euboea by Cheylan (1988), while Niethammer (1982) and Krystufek (1999b) mentioned it as absent from Bonn zoological Bulletin 57 (2): 367-373 the entire Mediterranean coast and islands (see also Masseti 2009). A label without specimen, written by Ioannis C. Ondrias himself, in the mammal collection of the University of Patras (coll. no. 3158) reports the occurrence of the common vole Microtus arvalis (Pallas, 1779), from the area of Mytilene in south-eastern Lesbos. But, according to Stella Fraguedakis Tsolis (in litteris 13th July 2006), this species does not appear to exist or to have ever existed on this island. Furthermore, the specimen to which the label referred has unfortunately been lost. Contrary, the presence of Gunther’s vole M. guentheri (Danford & Alston, 1880) is known from Lesbos (Stamatopoulos & Ondrias 1995), but according to Krystufek & & Vohralik (2005) this is the only record from all Mediterranean islands so far. THE INSULAR EDIBLE DORMICE Erroneous evaluations, or rather inattentive reading of pub- lications of early authors have supported cultural models which are still difficult to eradicate, e.g. the consideration of the diffusion of several species of glirids in the Greek islands. One example is the erroneously supposed occur- rence of the forest dormouse Dryomys nitedula (Pallas, 1778). Erhard (1858) reported the occurrence of Myoxus nitela Schreber, 1782, a species of glirid, similar in name to the forest dormouse from Andros, Naxos and Siphnos, where it occurred in orchards and orange groves. This re- port supported the assumption that this rodent occurs on these islands, but in reality the taxonomic classification does not correspond to that of the forest dormouse. Ac- cording to Ellerman & Morrison-Scott (1951) Myoxus nitela is indicated as one of the synonyms of Eliomys quercinus (Linnaeus, 1766; garden dormouse), a species currently unknown in the Aegean area and being wide- spread in the central-western Mediterranean basin. Here it is not found further east than Dalmatia and the north- western Balkan Peninsula. Although according to Krystufek (1999b), this forest dormouse does not occur on Mediterranean islands, Cheylan (1988) reported it from Euboea. Recently, the presence of the forest dormouse was reported on the island of Andros, which is still an uncon- firmed record (Chondropoulos & Fraguedakis-Tsolis, in verbis). We have, on the other hand, known for some time of the presence of the edible dormouse on islands such as Crete (Zimmermann 1953; Kahmann 1959; Niethammer & Krapp 1978; Catsadorakis 1994), Euboea (Ondrias 1966), Corfu (Niethammer 1962; Niethammer & Krapp 1978) and Cephalonia (Niethammer & Krapp 1978; Cat- sadorakis 1985; Giagia-Athanassopoulou 1998). On the latter island, its occurrence has been recently confirmed by H. Pieper (in litteris), whereas Dimaki (1999) provid- ed arguments for the existence of the species on Andros. According to H. Alivitzatos & A. Lane (in verbis), the ed- ©ZFMK Homeless mammals from Ionian & Aegean islands ible dormouse is also present on the island of Thassos where they mentioned its occurrence in the surroundings of the village of Panaghia, on 30 August 2000. Wettstein (1942) reports the occurrence of a dormouse, possibly the forest dormouse, from Rhodes, but according to other au- thors the species is still unknown here (cf. Festa 1914; De Beaux 1929; Zimmermann 1953). A remarkable human impact on the geographical distribution of some dormouse species in the Mediterranean region was observed by Carpaneto & Cristaldi (1994), Colonnelli et al. (2000) and Masseti (2005). The population density can be document- ed since antiquity through historical and biogeographical analyses, supported by paleontological and archaeozoo- logical data. Furthermore, ethnozoological enquiries doc- ument the utilisation of dormice for food or medicine, through traditional captive-breeding techniques, up to very recent historical times. HOMELESS GREEK ISLAND CARNIVORES IN THE EUROPEAN MUSEUMS Several European natural history museums conserve ma- terial collected on the Greek islands which create prob- lems in the attempt to arrive at their origins. This is the case, in the lynx, Lynx lynx (Linnaeus, 1758), collected on the island of Corfu and part of the collection of the Mu- seum Alexander Koenig in Bonn, registered under the col- lection number ZFMK 93423. The specimen was pur- chased by Jochen Niethammer during the mammalogical exploration of the island. But the occurrence of the lynx on Corfu was very questionable and immediately resolved by the collector himself. Niethammer reported that he had bought it at the market, where he had been told that it orig- inated from northern Greece, more specifically from Macedonia. In other cases specimens represent species which are in fact completely unknown to the islands which they are reported to originate from. In some cases, species have recently become extinct, like jackals from Corfu rep- resented in the collections of the Museum Koenig (ZFMK 61193, 93420). Dispersed in the Balkan and Ana- tolian peninsulas, the Golden or Asiatic jackal Canis au- reus Linnaeus, 1758 has been reported from Corfu (Ni- ethammer 1962; Douma-Petridou 1977; Adamakopoulos et al. 1991; Demeter & Spassov 1993), Cephalonia (Deme- ter & Spassov 1993), Lefkada (Douma-Petridou 1977; Demeter & Spassov 1993) and Kythera (Jameson 1836, 1937), while other authors mentioned its occurrence on Ikaria (Atanassov 1955) and Skyros (Werner 1928; Wettstein 1942; Atanassov 1955). Ioannidis & Giannatos (1991) surveyed with positive results the island of Samos where jackals exist in the same habitats as in the rest of the southern Balkan Peninsula. Following the account of the expedition to the Greek archipelago published by the botanist Joseph P. de Tournefort (1717), Clarke (1801) ob- Bonn zoological Bulletin 57 (2): 367-373 369 Fig. 2. Stuffed specimen of the badger Meles meles collec- ted on the island of Santorini (Thera) in 1859, and part of the collection of the Zoological Museum of the University of Athens (ZMUA 128) (photo Anastasios Legakis; courtesy Zoological Museum of the University of Athens). served that “Samos is infested with wolves”. Anyway, this record should refer to jackals rather than wolves. There is in fact no evidence for the occurrence of the latter canides on the Greek islands of the late Holocene. Accord- ing to Ioannidis & Giannatos (1991), the jackal no longer exists on Corfu, Kythera, Skyros and Ikaria, where it pos- sibly became extinct in very recent historical times, but jackals vanished from Corfu not before 1991-1992 (Grémuillet, in verbis). The only Aegean islands where the species still survives are Euboea (Demeter & Spassov 1993) and Samos (Laar & Daan 1967; Douma-Petridou 1977; Adamakopoulos et al. 1991; Ioannidis & Giannatos 1991; Demeter & Spassov 1993; Ioannidis et al. 1996; Dimitropoulos et al. 1998). Among the collections of the Greek museums, there are several specimens that provoke questions which are still far from having been satisfactorily answered. For exam- ple, there is a stuffed badger, Meles meles (Linnaeus, 1758) today on display at the Zoological Museum of the University of Athens (ZMUA 128, Fig. 2) and collected on the island of Santorini (Thera) by K. Bassiliou in 1859. This specimen is intriguing because of the old age and it is the only record of the badger from this island. Accord- ing to Schmalfuss (1991) the species is today unknown from Santorini. If the origin of the ZMUA specimen is cor- rect, the species must have become extinct around the end of the nineteenth century because Douglas (1892) did not mention the badger in his list of the insular mammals. San- torini should therefore be added to the distribution areas of the badger within the Aegean islands. Known in Greek as asvos, the badger was recorded from Cephalonia (Cat- sadorakis 1985), Rhodes (Festa 1914; Tortonese 1973) and Crete (Raulin 1859; Barrett-Hamilton 1899; Bate 1906, ©ZFMK 370 Marco Masseti Fig. 3. Detail of the early 16th century wall decoration sho- wing the “Life of St. Benedict” in the Great Cloister of the mo- nastery of Monte Oliveto Maggiore (Siena, Italy) painted by the Italian artist Giovanni Antonio Bazzi. 1913; Miller 1907, 1912; Zimmermann 1953; Ondrias 1965; Ragni et al. 1999) where it is locally indicated by the vernacular term arkalos. In the course of the present study, it was possible to confirm its occurrence on the is- lands of Tinos, where it is locally known as chakalos (Gaetlich, pers. com.), Euboea, Crete, Rhodes, and pos- sibly Andros (Gaetlich, pers. com.). There are unconfirmed records of badgers from Siphnos (Erhard 1858; Heldre- ich 1978; Cheylan 1988), but this does not exclude a pri- ori the possibility of a previously more widespread dis- tribution in the Aegean basin, and more specifically on the Cyclades. Moreover, the human practice of the 1mporta- tion of badgers onto the Greek islands is documented since prehistorical times. On Crete the oldest bones of M. meles were discovered in the Aceramic Neolithic levels at Knos- sos, while Ceramic Neolithic and later levels produced nu- merous remains of the species (Jarman 1996). Other os- teological material was found on the site of Aghia Tria- da, and Kavousi-Vroda and has been respectively referred to the Ancient Minoan period (about 3,000—2,200 B.C.) (Wilkens 1996), and to the Late Minoan HI C (Klipper & Snyder 1991; Snyder & Klippel 1996). It is not immedi- ately apparent why human should have wanted to intro- duce badgers onto the islands, which is suggested because otherwise they would not have been able to pass unob- served on the small boats employed to reach the new ter- ritories (Vigne 1988, 1995; Masseti 1995). Since very an- Bonn zoological Bulletin 57 (2): 367-373 cient times, they may have played an important role in hu- man societies, both symbolically and as food. Badgers might also have been utilised for their fur (Masseti 1995). Moreover, in medieval Europe another use of this mustelid has been documented. Wall paintings from the early 16th century (Fig. 3) at the monastery of Monte Oliveto Maggiore (Siena, Italy), painted by the Italian Gio- vanni Antonio Bazzi, better known as Sodoma, clearly show badgers as pets, very likely representing an authen- tic status symbol that underscored the affluence and so- cial position of their owner, the painter himself (Carli 1980). LEOPARDS REMARKS ON SAMOS - CONCLUDING A stuffed adult leopard (Fig. 4) is on display at the Natural History Museum of the Aegean in Mytelenti, on the Greek island of Samos (Masseti 2000). This specimen previously belonged to the Town Council (Greek: Nomarkia) and has been exhibited there for several decades (Ioannidis et al. 1996; Dimitropoulos et al. 1998). On its label it is classified as kap/ani, with the explanation that this is the Samian terminology indicating a species of panther. However, the word derives from the Turkish term kaplan, commonly used in Anatolia to indicate the tiger, and erroneously also the leopard (Danford & Alston 1880). On the basis of available information, it is today not possible to ascertain the age and the origin the specimen. It is said that the leopard was killed on the island between 1870 and 1880, but there is no evidence that this is correct. The title of one of the most famous novels of the contemporary Samian writer Alki Zei, To kaplani tis vitrinas (~The kaplani of the showcase), better known however as Wildcat under glass, was inspired by this leopard. Speaking of her childhood, the author described this kaplani, and since she was born in 1936, it can be presumed that the leopard is older. Unfortunately, the Samian specimen is of an unnatural shape because it has been rather inexpertly stuffed, and hardly recalls the form of a living individual. It has a total length of about 235 cm and tail length of 90 cm, apparently proving that this specimen is a large one. But since the skin of felids is extremely elastic, the original dimensions could have been altered during the taxidermic procedure. The coat colour has deteriorated due to bad preservation conditions, and its prolonged display under daylight. The hair of the skin is worn in patches, but it seems that originally the colouration was tawny or buff on the back and paler on the flanks, where it could have merged into the white of the belly. Today, the entire coat is uniform pale, with dark- brown rosettes along the flanks and the back, which are fairly large (about 34 cm in diameter), widely spaced and thinly rimmed, with the centres slightly darker than the ©ZFMK Homeless mammals from Ionian & Aegean islands 37] Fig. 4. The stuffed specimen of Asia Minor leopard, Panthera pardus tulliana Valenciennes, 1856, shown at the Natural Histo- ry Museum of the Aegean, Samos (Greece) (photo Marco Masseti; courtesy Natural History Museum of the Aegean, Mytelenii, Samos). ground tint. The coat is fairly short and full, the hair on the nape is long, and the tail is decidedly bushy. According to the colouration and coat pattern, this specimen could belong to the Anatolian leopard Panthera pardus tulliana, as mentioned by Valenciennes (1856), Pocock (1930) and Leyhausen (1991), and clearly distinct from other Near Eastern subspecies (Masseti 2000). It has also been said that the animal arrived at Samos from the opposite coast of Turkey, swimming across the channel separating the island from western Anatolia. In fact there is a deeply-rooted traditional belief on Samos which refer to leopards swimming from Anatolia in various periods. This was reported by Tournefort (1717) who confirmed this legend, observing that: ““‘// y passe quelques tigres qui viennent de terre ferme par le Petit Boghas’’. Petit Boghas was the name used at this time to indicate the above mentioned channel. Clarke (1801) followed this observation and mentioned that: “tigers sometimes arrive from the mainland, after crossing the little Boccaze; thereby confirming all observation made by the author in the former section, with regard to the existence of triggers in Asia Minor’. However, Tournefort (1717) report was probably not based on an own observation, but rather Bonn zoological Bulletin 57 (2): 367-373 inspired by local people. In any case, since the distance between the island and the mainland is not more than 1.7 km, it cannot be excluded that leopards could have reached the island by swimming, at various times. These felids are good swimmers and could have come e.g. from the Samsundag area (Masseti 2000) which was until the early 1970s the last western Anatolian stronghold of the species (Kumerloeve 1971; Avci 1978; Ulrich & Riffel 1993; Masseti 2000). Acknowledgements. I would like to express my appreciation and gratitude to the following friends and colleagues for their suggestions and assistance while the preparation of the present paper: Wolfgang Bohme and Rainer Hutterer, Zoologisches Forschungsmuseum Alexander Koenig, Bonn; Suleyman Karakaya and Suleyman Kagar, Forest Department of Antalya (Turkey); Ioannis C. Ondrias, Basil Chondropoulos, Stella Fraguedakis-Tsolis and A. 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Zeitschrift fiir Saugetierkunde 67: 1-72 Received: 01.1X.2010 Accepted: 03.X1.2010 OZFMK ‘ad vey eel ore aay a it Pl. f ve i j 4 , nia ial irae ‘ th) ab ribald ah Bonn zoological Bulletin (BzB) Instructions to authors Scope The Bonn zoological Bulletin (BzB), formerly “Bonner zoologi- sche Beitrage”, is an international, peer-reviewed, open access jour- nal publishing original research articles, reviews, and scientific notes dealing with organismal zoology. Focus of the BzB are (1) taxon- omy, (2) systematics and evolution, and (3) biodiversity and bio- geography, all with respect to terrestrial animals. Terrestrial animals as understood here include those inhabiting fresh or brackish wa- ters. Contributions from related fields like ecology, morphology, anatomy, physiology or behaviour are welcome when of clear rel- evance to the focus topics. 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WN WwW ‘Oo OND Contents Descriptions of New Taxa Joger, Ulrich & Bshaenia, Ismail: A new Tarentola subspecies (Reptilia: Gekkonidae) endemic to Tunisia Wilms, Thomas M., Shobrak, Mohammed & Wagner, Philipp: A new species of the genus Tropiocolotes from Central Saudi Arabia (Reptilia: Sauria: Gekkonidae) Lutzmann, Nicola, Stipala, Jan, Lademann, Ralph, Krause, Patrick, Wilms, Thomas M. & Schmitz, Andreas: Description of a new subspecies of Kinyongia uthmoelleri (Muller, 1938) (Squamata: Chamaeleonidae) with notes on its captive propagation Vogel, Gernot & David, Patrick: A new species of the genus Lycodon (Boie, 1826) from Yunnan Province, China (Serpentes: Colubridae) Wagner, Philipp & Wilms, Thomas M.: A crowned devil: new species of Cerastes Laurenti, 1768 (Ophidia, Viperidae) from Tunesia, with two nomenclatural comments History of Herpetology Schmidtler, Josef Friedrich: The taxonomic history of the Linnean genus Lacerta (Squamata: Sauria: Lacertidae) in the mirror of book-illustration Pafilis, Panayiotes: A brief history of Greek herpetology Niekisch, Manfred: The history of reptiles and amphibians at Frankfurt Zoo Non-herpetological articles Hita Garcia, Francisco, Fischer Georg, Ktick, Patrick, Thormann, Birthe & Peters, Marcell K. Tetramorium boehmei sp. n. — a new ant (Hymenoptera: Formicidae) species from the Kakamega Forest, Western Kenya Masseti, Marco: Homeless mammals from the lonian and Aegean islands 267 275 28) 289 297 307 329 347 359 367 re a es yo SG Ian eam ae vol tone vo Contents UN 567 9293 Ecology & Evolution Carretero, Miguel A., Cascio, Pietro Lo, Corti, Claudia & Pasta, Salvatore: Sharing recources in a tiny Mediterranean Island? ; Comparative diets of Chalcides ocellatus and Podarcis filfolensis in Lampione Kuhnel, Susanne, Reinhard, Sandy & Kupfer, Andreas: Evolutionary reproductive morphology of amphibians: an overview Checklists, Nomenclature & Distribution Koch, André, Auliya, Mark & Ziegler, Thomas: Updated checklist of the monitor lizards of the world (Squamata: Varanidae) 111 119 127 Ziegler, Thomas & Nguyen, Truong Quang: New discoveries of amphibiens and reptiles from Vietnam Dubois, Alain & Bour, Roger: The distinction between family-series and class-series nomina in zoological nomenclature, with emphasis on the nomina created by Batsch (1788, 1789) and on the higher nomencature of turtels 137 149 Jirkt, Miloslav, Mihalca, Andrei Daniel, Necas, Petr & Modry, David: An addition to the East African herpetofauna: the first record of Tarentola annularis relicta (Squamata: Gekkonidae) in Uganda Taxonomy Rédel, Mark-Oliver, Sandberger, Laura, Penner, Johannes, Mané, Youssouph & Hillers, Annika: The taxonomic status of Hyperolius spatzi Ahl, 1931 and Hyperolius nitidulus Peters, 1875 (Amphibia: Anura: Hyperoliidae) 173 177 Capula, Massimo & Corti, Claudia: Genetic variability in mainland and insular populations of Podarcis muralis (Reptilia: Lacertidae) 189 Arribas, Oscar J.: Intraspecific variability of the Carpetane Lizard (/berolacerta cyreni [Muller & Hellmich, 1937]) (Squamata: Lacertidae), with special reference to the unstudied peripheral populations from the Sierras de Avila (Paramera, Serrota and Villafranca) 197 Bare}, Michael F., Ineich, lvan, Gvozdik, Vaclav, Lhermitte-Vallarino, Nathaly, Legrand Gonwouo, Nono, LeBreton, Matthew, Bott, Ursula & Schmitz, Andreas: Insights into chameleons of the genus 7rioceros (Squamata: Chamaeleonidae) in Cameroon, with the resurrection of Chamaeleon serratus Mertens, 192 221 Descriptions of New Taxa GUnther, Rainer: Another new Cophixalus species (Amphibia: Anura: Microhylidae) from western New Guinea 231 Vences, Miguel, Kohler, Jorn, Crottini, Angelica & Glaw, Frank: High mitochondrial sequence divergence meets morphological bioacoustic conservatism: Boophis quasiboehmei/ sp. n., a new treefrog species from south-eastern Madagascar Bauer, Aaron M.: A new species of Pachydactylus (Squamata: Gekkonidae) from the Otavi Highlands of northern Namibia 241 257 yy Ministerium fiir Innovation, ' ' b Wissenschaft, Forschung i i is und Technologie des Landes eh RW. Lei i iz <@S” Nordrhein-Westfalen @ G emein sch aft | |