Bonn Zoological Bulletin formerly: Bonner zoologische Beitrage Volume 60 Issue 1 ZC 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 Editorial Board Dirk Ahrens, Insects: Coleoptera, ZFMK, tel. +49 2289122286, fax: +49 228-9122-332; d.ahrens.zfmk(@uni-bonn.de Wolfgang Boéhme, 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: +49 228-9122-212; r.elzen.zfmk@uni-bonn.de Bernhard Huber, Invertebrates except Insects, ZFMK, tel. +49 228-9122—294, fax: +49 228-9122-212; b.huber.zfmk@uni-bonn.de Rainer Hutterer, Mammals, ZFMK, tel. +49 2289122261, fax: +49 2289122212; r.hutterer.zfmk@uni-bonn.de Gustav Peters, Mammals, ZFMK, tel. +49 2289122262, fax: +49 228-9122-212; g.peters.zfmk@uni-bonn.de Bradley Sinclair, Canadian National Collection of Insects, Ottawa Plant Laboratory — Entomology, CFIA, K.W. Neat- by Bldg., C.E.F., 960 Carling Ave., Ottawa, ON, Canada K1A 0C6, tel. + 1 613-759-1787, fax: + 1 613—759— 1927, bradley.sinclair@inspection.ge.ca Dieter Stiining, Insects except Coleoptera and Dive ZFMK, tel. +49 228-9122-220, fax: +49 228-9122~ 212; d.stuening.zfmk(@uni-bonn.de a Philipp Wagner, Villanova University, Department of Biolo- gy, 800 Lancaster Avenue, Villanova, PA 19085-1699, USA; philipp.wagner.zfmk@uni-bonn.de Advisory Board Theo C. M. Bakker, Rheinische Friedrich-Wilhelms-Univer- sitat, Institut fiir Evolutionsbiologie & Okologie, 53113 Bonn, Germany, tel. +49 228—73—5130, fax: +49 228—73- 2321; t.bakker@uni-bonn.de Aaron M. Bauer, Villanova University, Department of Biolo- gy, 800 Lancaster Avenue, Villanova, PA 19085-1699, USA, tel. +1 610-519-4857, fax: +1 610-519-7863; aaron.bauer@villanova.edu Wieslaw Bogdanowicz, Museum and Institute of Zoology, Polish Academy of Sciences, Wilcza 64, 00-679 Warszawa, Poland, tel. +48 22—628—7304, fax: +48 22—629-6302; wieslawb@miiz.waw.pl Matthias Glaubrecht, Museum ftir Naturkunde Berlin, Leib- niz-Institut fir Evolutions- und Biodiversitatsforschung an der Humboldt-Universitat zu Berlin, Invalidenstrasse 43, 10115 Berlin, Germany, tel. +49 30—2093—8504/ 8400, fax: +49 030—2093-8565; matthias.glaubrecht@mfn- berlin.de Jeremy D. Holloway, The Natural History Museum, Depart- ment of Entomology, Cromwell Road, London, SW7 5BD, U.K.; j.holloway@nhm.ac.uk Tan Heok Hui, Raffles Museum of Biodiversity Research, National University of Singapore, Department of Biological Sciences, 6 Science Drive 2, #03-01, Singapore 117546, tel. +65-6516 1662, heokhui@nus.edu.sg Boris Krystufek, Slovenian Museum of Natural History, P. O. Box 290, Ljubljana, Slovenia; boris.krystufek@zrs.upr.si Wolfgang Schawaller, Staatliches Museum fir Naturkunde, Rosenstein 1, 70191 Stuttgart, Germany, tel. +49 711-8936—-221, fax: +49 711-8936-100; schawaller.smns@naturkundemuseum-bw.de Ulrich K. Schliewen, Department of Ichthyology, Bavarian State Collection of Zoology, Minchhausenstr. 21, 81247 Minchen, Germany, tel. + 49 89-8107—110; schliewen@zsm.mwn.de Michael Schmitt, Ernst-Moritz-Arndt-Universitat, Allge- meine & Systematische Zoologie, Anklamer Str. 20, 17489 Greifswald, Germany, tel. +49 3834—-86-4242, fax: +49 3834-86—-4098; michael.schmitt@uni-greifswald.de _W. David Sissom, Dept. of Life, Earth and Environmental Sciences, W. Texas A. & M. University, WTAMU Box 60808, Canyon, Texas 79016, USA, tel. +1 806-651-2578, fax: +1 806-651-2928; dsissom@mail.wtamu.edu Miguel Vences, Technical University of Braunschweig, Zoo- logical Institute, Spielmannstr. 8, 38106 Braunschweig, Germany, tel. + 49 531-391-3237, fax: + 49 31-391-8198; m.vences(@tu-bs.de Erich Weber, Eberhard-Karls-Universitat, Zoologische Schausammlung, Sigwartstr. 3, 72076 Tubingen, tel. +49 70712972616, fax +49 7071—295170; erich.weber@uni-tuebingen.de Editorial Welcome to the 60" volume of the Bonn zoological Bul- letin. All contents published since 1950 are now available online at the ZFMK homepage, providing open access to numerous contributions to organismal zoology. The jour- nal has now been added to the Directory of Open Access Journals, further improving the visibility of papers pub- lished in the BzB. Besides open access status, publication remains free of charge for authors, including colour illus- trations. In sum, the journal is increasingly attractive for the publication of sound scientific work from its three fo- cus fields, namely (1) taxonomy, (2) systematics and evo- lution, and (3) biodiversity and biogeography. Taken together, 32 scientific papers and 10 catalogues of type specimens housed at ZFMK were published in more than 730 pages in the first year after changing the BzB ti- tle from “Bonner zoologische Beitrage” to “Bonn zoolog- ical Bulletin”. The papers contain, among other topics, de- scriptions of more than a dozen species new to science, ranging from butterflies to fishes, frogs and snakes. The current issue adds new species records, a comprehen- sive checklist, descriptions of five new species, and fur- ther contributions to organismal zoology. A molecular study suggests that an Amazonian Pecari described a few years ago likely does not represent a valid species, and a case study conducted on a small tropical island reports on the accumulating species richness data in a herpetologi- cal field survey. Last but not least, it’s an honour to welcome two new board members. Philipp Wagner, currently at Villanova University (USA), is a herpetologist and well known to BzB readers. He has a special interest in agamid lizards from Africa, has acted as guest editor previously, includ- ing last year’s special issue “Herpetologia Koenigiana” in honour of Wolfgang Bohme, and will now join the Edi- torial Board. Tan Heok Hui from Raffles Museum of Bio- diversity Research in Singapore will support the BzB team as a member of the Advisory Board. He is famous among ichthyologists, especially for his taxonomic work on Asian freshwater fishes including sucker loaches, fighting fish, and the smallest fish species recorded so far, a cyprinid. His interests include also herpetology and vertebrates as well as invertebrates occurring in marine, brackish and freshwater habitats of Asia. A warm “welcome on board” to both colleagues! With best wishes to all readers, authors and “BzB-team” members, Tibi float Fabian Herder (Editor-in-Chief, Fish Curator at ZFMK) Bonn, April 2011 - 5 Ui Hig es iia va cae ae ah a ae "Giese hu Dene ae ba | yee Bonn zoological Bulletin Volume 60 Issue | | pp. 3-8 Bonn, May 2011 | Estimating the herpetofaunal species richness of Pangkor Island, Peninsular Malaysia Johan van Rooyen!, Chan Kin Onn3, L. Lee Grismer+ & Norhayati Ahmad> 'Netherlands Centre for Biodiversity Naturalis, section Zoological Museum Amsterdam, Mauritskade 61, NL-1092 AD Amsterdam, The Netherlands 2Tulpentuin 313, NL-2272 EH, Voorburg, The Netherlands; E-mail: j1.van.rooijen@hetnet.nl 3Institute for Environment and Development (LESTARI), Universiti Kebangsaan Malaysia, 43600 Bangi, Selangor Darul Ehsan, Malaysia; E-mail: kin_onn@yahoo.com 4Department of Biology, La Sierra University, 4500 Riverwalk Parkway, Riverside, California, 92515-8247 USA; E-mail: lgrismer@lasierra.edu 5School of Environment and Natural Resource Sciences, Faculty of Science and Technology, Universiti Kebangsaan Malaysia, 43600 Bangi, Selangor Darul Ehsan, Malaysia; E-mail: yati_68@yahoo.co.uk. Abstract. Herpetological surveys of Southeast Asian tropical ecosystems rarely, if ever, result in complete inventories. This is due to the fact that surveying to completion requires huge investments in terms of search effort. As a result, the presented species lists usually represent subsets of the total herpetofaunal assemblage and consequently do not shed light on the total species richness of the investigated area. This is regrettable as species richness is an elementary measure of biodiversity that underlies many ecological models and conservation strategies. By recording not just species but species per unit of search effort, an extended dataset results which can be used to generate estimates of total species richness. In this study, the herpetofauna of Pangkor Island, Peninsular Malaysia is used as an example. In 2009 and 2010, the first herpetological surveys were carried out on this small, 18 km2, island. Those surveys recorded 43 species of reptiles and 13 species of amphibians. In this study, total reptile species richness was estimated by fitting several models to the sam- ple-based rarefaction curve as well as by application of the nonparametric Chao-I estimator. Of the applied models, the 4-parameter Weibull function was shown to be superior, a finding that is in line with several other studies. Consequent- ly, the use of this model is recommended. On the basis of the fitted 4-parameter Weibull-function, 69 reptile species are expected to occur on Pangkor Island. As for amphibians, total species richness was estimated to be 17. As such, a re- markably extensive herpetofaunal assemblage inhabits this small island. Key words. Pangkor Island, Malaysia; amphibians, reptiles, species-richness, sample-based rarefaction curve, negative exponential function, 3-parameter Weibull function, 4-parameter Weibull function, Chapman-Richards model, Chao-I es- timator INTRODUCTION Herpetological surveys of species-rich tropical ecosystems rarely result in complete inventories (e.g. Lloyd et al. 1968; Murphy et al. 1994; Hofer & Bersier 2001; Van Rooijen 2009). This is due to the asymptotical nature of the species accumulation process in combination with lim- ited survey-investments. As a consequence, the use of es- timation techniques is unavoidable when the intended ob- jective is to assess species richness, an elementary meas- ure of biodiversity that underlies many ecological mod- els and conservation strategies. One such estimation tech- nique consists of fitting an appropriate function to the species accumulation curve. The asymptote of the fitted functions can then be regarded as an estimate of total species richness. Species accumulation curves are regu- larly applied in herpetofaunal surveys but they are most- ly used to arrive at qualitative judgements about the exhaustiveness of the survey (e.g. Murphy et al. 1994; Zug et al. 1998; Ziegler 2002). However, extensive and sophis- ticated literature exists pertaining to methods and models used to quantitatively estimate species richness, on the ba- sis of either rarefied species accumulation curves or abun- dance patterns (e.g. Colwell & Coddington 1994; Flather 1996; Gotelli & Colwell 2001; Longino et al. 2002; Brose et al. 2003). The offshore archipelagos of Peninsular Malaysia have been the subject of increasing interest in recent years. Many of these islands have never been surveyed and re- cent explorations are only beginning to uncover the hid- den diversity and endemism that they shelter (Chan et al. 2009a; Grismer 2008; Grismer & Norhayati 2008; Gris- mer & Pan 2008; Grismer et al. 2008, 2009a, b). One such 4 Johan van Rooien et al. ni Fig. 1. al., 2010; D: Dryophiops rubescens (Gray, 1835). island is Pulau Pangkor, approximately 18 km2 and situ- ated 3.5 km from the west coast of Peninsular Malaysia. The island’s interior remains heavily forested and main- tains a river system, Sungai Pinang, which supplies a sig- nificant source of permanent fresh water in the form of multiple streams. Chan et al. (2010) provided the first re- port on the (non-marine) herpetofauna of this island and documented 43 reptiles and 13 amphibians. A few illus- trations are provided in figures | and 2. The study described in this paper had a dual objective. First, data collected by Chan et al. (2010) were used to evaluate the performance of several estimators. Second, the most appropriate estimator was used to estimate the total herpetofaunal species richness harboured by Pangkor Island, West Malaysia. MATERIAL AND METHODS The data underlying this study were based on the surveys carried out by Chan et al. (2010). These were conducted in the periods March 15—17, May 4-8, June 13 to July 8, 2009, and February 22 to March 8, 2010. During the lat- ter two periods, only reptiles were surveyed. Marine rep- Bonn zoological Bulletin 60 (1): 3-8 A: View into the primary forest of Pangkor Island; B: Boiga drapiezii (Bote, 1827); C: Cnemaspis shahruli Grismer et tiles were ignored altogether. The predominantly applied survey method corresponds to visual encounter survey (VES), a simple method which has been shown to be ef- fective for surveying rainforest herpetofauna (Doan, 2003). VES was carried out both during day and night (e.g. Coddington et al., 1996; Doan, 2003) in a way similar to that applied by Minh (2007). Existing trails as well as sev- eral trails made by the Department of Forestry of Perak and Peninsular Malaysia were used. These trails traversed dipterocarp forest, mangrove forest and cultivated areas and provided ample access to forest streams. The second collection method consisted of searching for road-kills. The third entailed turning logs, fallen tree bark and sim- ilar objects in order to uncover animals hiding underneath. Sampling effort was expressed in terms of search-days where one search-day was defined as roughly 4.5 search- hours. As two of the surveys underlying the data collect- ed by Chan et al. (2010) focused solely on reptiles, sub- stantially more information was available regarding rep- tiles allowing for more thorough analyses. Therefore, this study focuses predominantly on reptiles, amphibians be- ing treated separately and in less detail. Sample-based rarefaction curves (Gotelli & Colwell 2001) were generated with the program EstimateS (Colwell ©OZFMK Estimating herpetofaunal species richness nn Fig. 2. A: Heosemys spinosa (Gray, 1831); B: Kaloula pulchra Gray, 1831; C: Naja sumatrana (Miller, 1890); D: Tropidolae- mus wagleri (Boie, 1827). 2005). Four models were fitted to the rarefaction curve. The first corresponds with a negative exponential model (Colwell & Coddington 1994; Flather 1996; Van Rooyen 2009). It is based on the assumption that the number of new species found per search day is proportional to the number of as yet undiscovered species, in mathematical terms: dY/dt = c (A—Y) where A is the total number of species present in the area under investigation, Y is the to- tal number of species found and c is a constant. This equa- tion can be represented as a negative exponential function (e.g. Van Rooyen 2009): Y= A (1—-e~*'). The basic assump- tion underlying the negative exponential model (hence- forth NE) may be overly simplistic given that abundance patterns are usually strongly skewed (e.g. Lloyd et al. 1968; Coddington et al. 1996; Limpert et al. 2001; Longi- no et al. 2002; Thompson et al. 2003). In order to model more complex species accumulation processes, the NE can be refined in various ways by adding one (d) or two pa- rameters (d and p), resulting in the Chapman-Richards model (henceforth CR), 3- and 4-parameter Weibull cu- mulative distribution functions (henceforth 3pW and 4pW): Y =A (1-e«t)* (CR), Y=A (1-e()“) pW), Y= A (1-eep)) (4pW) Bonn zoological Bulletin 60 (1): 3-8 The four models were fitted to the sample-based rarefac- tion curve using nonlinear regression analysis (e.g. Noru- sis and SPSS 1994) with SPSS (release 14 February 1996; SPSS Inc.). Extrapolation using different models for the species ac- cumulation process can provide different asymptotes and thus predict different values of species richness (e.g. Col- well & Coddington 1994; Flather 1996). Therefore, care has to be taken to select the most appropriate model in or- der to minimize bias. In this study, the appropriateness of each model was evaluated on the basis of three criteria. The first criterion was goodness-of-fit. The second crite- rion entailed the behaviour of the richness-estimate with increasing cumulative search effort. The final criterion was the difference with the nonparametric Chao-I estimator (Chao 1984; Coddington et al. 1996; Hofer & Bersier 2001; Veith et al. 2004). This estimator is an often applied representative of a class of estimators that uses a differ- ent approach as they are based on abundance patterns in- stead of the accumulation curve. The Chao-I estimator is based on the observed number of rare species, A = Y + (a?/2b), where A is the total number of species, Y is the observed number of species, a is the number of observed species represented by a single specimen and 6 OZFMK 6 Johan van Rooijen et al. is the number of observed species represented by exact- ly two specimens. RESULTS Figure 3 depicts the deviations of the fitted functions from the sample-based rarefaction curve (residuals) for reptiles. As expected, the NE exhibits a very poor goodness-of-fit. At first, the fitted curve is situated beneath the rarefaction curve, then above and finally drops beneath the rarefac- tion curve again. The suboptimal fit of the NE as well as the pattern of residuals are in agreement with results ob- tained with regard to avian diversity (Flather 1996) as well as snake diversity (Van Rooien 2009). Although the RC and 3pW fit substantially better than the NE, they exhib- it a similar pattern of residuals. The 4pW function obvi- ously exhibits a superior, near-perfect, fit. residual QRROOS negative exponential : —— 3-parameter Weibull -2,5 Miyake - 7 > 4-parameter Weibull ——— Chapman-Richards search days Fig. 3. | Deviations of the fitted models from the sample-based rarefaction curve (residuals). Figure 4 depicts the sample-based rarefaction curve as well as the fitted 4pW. The only noticeable deviation of the fit- ted 4pW from the rarefaction curve lies in the fact that it passes through (p,0) instead of the origin (0,0). The close fit of the 4pW is in agreement with results obtained in studies of avian diversity (Flather 1996), snake diversity (Van Rooijen 2009) and diversity of small reptiles (Thompson et al. 2003). The progression of the rarefac- tion curve clearly indicates that an asymptote has not yet been reached, thus the surveys have not been exhaustive. Figure 5 shows how the richness estimates (as opposed to observed richness or fitted model-values) develop with increasing search days during the second half of the sur- vey. The NE is neglected due to its poor fit. Evidently, the 3pW- as well as CR-based estimates are still increasing at the end of the survey whereas the 4pW-based estimate Bonn zoological Bulletin 60 (1): 3-8 acne | | | | | | 2 3 cumulative number of species 0 1 ZS OO. Fi ACI 9) AOR A 14 201314516 fa Seto m2 mee cumulative number of search days Fig. 4. Reptile species sample-based rarefaction curve with fitted 4-parameter Weibull function. has reached a more or less stable level. A similar result (unpublished) was obtained on the basis of a dataset that underlies a study of Bornean snakes (Van Rooijen 2009): the 4pW-based estimate reached an approximately stable plateau rather early with the CR- and 3pW-based estimates approaching the 4pW-based estimate with increasing search days. On the basis of these results, the 4pW-based estimate, 69, is assumed to be the least biased. Finally, the estimate based on the nonparametric Chao I estimator, 62, agrees reasonably well with the 4pW-based estimate. Taking the average, 65 reptile species are expected to inhabit Pangkor Island. As for amphibians species richness, the fitted 4pW and Chao I estimator both resulted in an esti- mate of 17 species. 2): ete cio cho OO OO OP CLO OOOO OOOO oo aoc o mS 65 | 60 4 @ 504 = ze | B ao n g 35 4 2 304 2 254 20 4 1) 3-parameter Weibull 10 * = * 4-parameter VWeibull 5 | —— Chapman-Richards | 0 oot : —————— 10 1 12 13 14 15 16 7 18 419 20 21 cumulative number of search days Fig. 5. Richness estimates in relation to cumulative search ef- fort. ©OZFMK Estimating herpetofaunal species richness a/ DISCUSSION Estimation by extrapolation, as opposed to interpolation, into the unknown entails a high degree of uncertainty. However, the alternative, surveying until completion, is simply unfeasible when the survey is aimed at the herpeto- fauna of a Southeast Asian rainforest ecosystem (Van Rooijen 2009). As such, there simply is no alternative. Thus, further refinement of estimation methods is of im- portance. That said, approximately complete species lists are obviously still crucial for many zoogeographic stud- ies. Such species lists can be (and are) compiled by com- bining results of various surveys. However, this usually entails the combination of species lists pertaining to dif- ferent locations. As such, completeness comes at a price as one has to accept a huge decrease in spatial resolution. In most zoogeographic studies however, a high resolution is not essential as species compositions of major land masses are compared (e.g. In Den Bosch 1985; How and Kitchener 1997; Inger and Voris 2001). At the spatial scale which is relevant for conservation though, complete species lists are a utopia in the majority of cases and es- timation will therefore gain in importance (Van Rooijen 2009). This study underscores the notion that a combina- tion of criteria needs to be applied to select the most ap- propriate estimator. The 4-parameter Weibull function ex- hibited the highest goodness-of-fit. Moreover, it produced the most stable estimate near the end of the survey (fig- ure 5). Finally, it resulted in a richness-estimate that agreed well with the estimate based on the nonparametric Chao- I estimator. Even then, the estimate may be downwardly biased due to suboptimal sampling. First of all, although searches were carried out during both day and night, no searches were performed in the early morning (before dawn). Secondly, sampling of canopy-microhabitat was obviously unfeasible. Finally, as true species richness is underestimated by most estimators when sample size is small (Colwell & Coddington 1994; Canning-Clode et al. 2008), the estimate of amphibian species richness may be downwardly biased since it was based on only five search days. Nevertheless, the amphibian species richness appears to be rather low which is in agreement with the impression of herpetologists who carried out the surveys (Chan et al. 2010). The fact that the 4-parameter Weibull function exhibits such a good fit when applied to diverse ecological com- munities such as small reptiles in desert habitat (Thomp- son et al. 2003), birds (Flather 1996) and reptiles in rain- forest habitat (Van Rooijen 2009 and this report) is strik- ing. Two explanations can be put forward. First, relative species abundances follow very similar patterns over a wide range of ecological communities: relatively few species are abundant whereas most are rare (e.g. Hughes 1986). Thus, the shape of the species accumulation process Bonn zoological Bulletin 60 (1): 3—8 may also be expected to be rather uniform. Second, the higher the number of parameters of a model, the better the fit. Thus, the 4pW may simply be expected to fit better than similar functions with fewer parameters, irrespective of the field of application. 65 reptilian species are estimated to inhabit Pangkor Is- land, 43 of which have been recorded (Chan et al. 2010). A major part of the as yet unrecorded species will con- cern snakes as members of this taxon are notoriously hard to find due to their elusive habits and low densities (e.g. Lloyd et al. 1968; Inger & Colwell 1977; Hofer & Bersi- er 2001; Orlov et al. 2003; Van Rooijen 2009). This no- tion is strengthened by several indirect observations. Lo- cals provided accounts regarding observations of Boiga dendrophila (Bote, 1827), Cryptelytrops purpureomacu- latus (Gray, 1832), Ophiophagus hannah (Cantor, 1836) and Maticora bivirgata (Boie, 1827). In addition, there has been a visual record of a Coelognathus radiatus (Boie, 1827) (Schultz, pers. comm.). As amphibian species rich- ness is estimated to be at least 17, herpetofaunal species richness is expected to equal at least 82 species, which is quite impressive given the fact that Pangkor Island encom- passes merely 18 km?. On the other hand, whether Pangkor Island harbours a comparatively extensive her- petofauna is impossible to determine at the moment as species richness on other Malaysian islands has not yet been estimated. Estimating species richness of specific sites such as islands obviously has added value as species richness is an ele- mentary criterion a conservationist may use when select- ing sites and is crucial for many ecological studies. How- ever, the function fitted to the rarefaction curve not only provides an estimate of total species richness but also pro- vides insight in expected return on further investment: how many previously unrecorded species may be expected to be found with additional search effort? For instance, on the basis of the fitted 4-parameter Weibull function, ten previously unrecorded reptiles are expected to be found on Pangkor Island when 20 additional search days are in- vested. Such statistical expectations can be used as input for a cost-benefit evaluation when choices have to be made between different sites for the investment of survey-ca- pacity. Alternatively, one might determine how much search effort would have to be invested to bring the sur- vey to some specified level of completeness. REFERENCES Brose U, Martinez ND, Williams RD (2003) Estimating species richness: sensitivity to sampling coverage and insensitivity to spatial patterns. Ecology 84: 2364-2377 Canning-Clode J, Valdivia N, Molis M, Thomason JC, Wahl M (2008) Estimation of regional richness in marine benthic com- ©ZFMK 8 Johan van Rooyen et al. munities: quantifying the error. 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Hamadryad 23: 111-120 Received: 21.06.2010 Accepted: 14.03.2011 Corresponding editor: F. Herder OZFMK Bonn zoological Bulletin Volume 60 Issue | pp. 9-16 Bonn, May 2011 New records of snakes from Cat Tien National Park, Dong Nai and Lam Dong provinces, southern Vietnam Peter Geissler '!, Truong Quang Nguyen !, Nikolay A. Poyarkov 34 & Wolfgang Bohme ! | Zoologisches Forschungsmuseum Alexander Koenig, Adenauerallee 160, 53113 Bonn, Germany; E-mails: pgeissler84@yahoo.de; w.boehme.zfmk(@uni-bonn.de 2 Institute of Ecology and Biological Resources, 18 Hoang Quoc Viet, Hanoi, Vietnam; E-mail: nqt2@yahoo.com 3 Lomonosov Moscow State University, Biological faculty, Department of Vertebrate Zoology, Leninskiye Gory, Moscow, GSP-1, 119991, Russia 4 Joint Russian-Vietnamese Tropical Research and Technological Center of the A.N. Severtsov Institute of Ecology and Evolution, South Branch, 3, Street 3/2, 10 District, Ho Chi Minh City, Vietnam; E-mail: n.poyarkov@gmail.com. Abstract. We report ten new records of snakes from Cat Tien National Park, Dong Nai and Lam Dong provinces, south- ern Vietnam. The specimen of Zyphlops siamensis Gunther, 1864 from Cat Tien represents the second country record in Vietnam after more than one century. Our new record of Dendrelaphis ngansonensis (Bourret, 1935) leads to a consid- erable range extension of this species, so far known only from northern and central Indochina. Additional specimens of Oligodon deuvei David, Vogel & van Rooijen, 2008 are described, along with the first photograph of a living specimen, showing the coloration including the aposematically red-coloured underside of the tail. An updated checklist of snakes of the Cat Tien National Park is also provided. Key words: Squamata: Serpentes: Typhlopidae, Colubridae, Cat Tien National Park. INTRODUCTION Among the 545 species of reptiles and amphibians known from Vietnam, snakes have the highest species diversity (Nguyen et al. 2009). However, many snake species are often recorded only by a single or rather few specimens, and data on their distribution and natural history are still deficient. Though the Cat Tien National Park is the biggest and most important National Park for the lowland rain- forests of southern Vietnam (Polet & Ling 2004), its her- petofauna is still poorly studied (Nguyen 1988, Le et al. 1998, Nguyen & Ho 2002, Le 2007, Geissler et al. 2009). The most recent checklist of reptiles and amphibians of the Cat Tien National Park, which was compiled by Nguyen & Ho (2002), reported a total of 42 snake species. We herein provide an updated checklist of snakes from this national park with ten new records and discussion about the status of some rare and poorly known species in Viet- nam. MATERIAL AND METHODS Field work was conducted by Peter Geissler in July and August 2008 and from March until June 2009; by Niko- lay A. Poyarkov from November until December 2007, from February until April 2008, from February to April 2009, and in July 2010 in Cat Tien National Park, Dong Nai Province, southern Vietnam (Fig. 1). Species identi- fications were also based on the examination of specimens collected by colleagues from the Appalachian State Uni- versity (North Carolina, USA). A total of 54 specimens were examined and they were subsequently deposited in the collections of the Institute of Ecology and Biological Resources (IEBR), Hanoi, Vietnam; the Zoologisches Forschungsmuseum Alexander Koenig (ZFMK), Bonn, Germany; and the Zoological Museum, Moscow State University (ZMMU), Moscow, Russia. The following measurements were taken with a digital vernier calliper: SVL (snout-vent length); TL (total 10 Peter Geissler et al. East Sea (South China Sea) VI2°€ 2D, len Seat Paracel Island x 114€ Elevation (m) |. FI WF 1200+ Tt Gulf of 4 0: 2 (7S 150 R+—+ +S Kilometers Ship teens tail Spratly,islonds + | * Thailand 2 Sa hee ailond f : pe pms 0 150 300 Kilometors aire Fig. 1. | Map showing the study site (black diamond) in south- ern Vietnam: Cat Tien National Park. length); TaL (tail length). In addition, several scale counts were taken: VEN (number of ventrals); MD (middorsal scales); MBS (midbody scale rows, including the ven- trals); SC (subcaudals); IL (infralabials); SL (supralabi- als). TAXONOMIC ACCOUNT Typhlops siamensis Giinther, 1864 Specimen examined: One adult male (ZFMK 88922), collected by P. Geissler on the road in a rainy evening on 31 March 2009, at the Headquarters of Cat Tien Nation- al Park, Dong Nai Province (near 11°25719.3”N, 107°25°42.0”E, 104 m a.s.l.). Characteristic features: The morphological features of this specimen fit the descriptions of Giinther (1864) and Bonn zoological Bulletin 60 (1): 9-16 Wallach (2001): SVL 140.2 mm; TL 4.4 mm; preocular separated from nasal; inferior nasal suture contacting sec- ond supralabial; superior nasal suture extending horizon- tally to rostral, not visible from above; 22 scale rows around midbody (including ventral scale rows); 308 par- avertebral scales (306-368 according to Wallach 2001); dorsum uniformly dark brown, venter cream (see Fig. 2a). Remarks: Wallach (2001) limited the distribution of this species to Thailand and Cambodia, although it was record- ed from Vietnam by Nguyen & Ho (1996), see also Nguyen et al. (2009). The record is based on a specimen (ZISP 5426), which was collected by Tramond in 1879 and subsequently deposited in the collection of the Zoologi- cal Institute in St. Petersburg (Tirant 1885). Therefore, our record from Cat Tien National Park represents a rediscov- ery of this species in Vietnam after 130 years. In main- land Southeast Asia, both Typhlops diardi Schlegel, 1839 and Zyphlops muelleri Schlegel, 1839 are morphological- ly similar to Zyphlops siamensis. T: siamensis differs from T: diardi in having a lower number of scale rows around midbody (22 vs. 24-28). Bourret (1936) synonymized 7: siamensis with the subspecies 7. diardi nigroalbus Dumeril & Bibron, 1844. Typhlops diardi nigroalbus was recently synonymized with Zyphlops muelleri Schlegel, 1839 by Wallach (2001). 7: siamensis can be distinguished from 7. muelleri by having a lower count of MBS (22 vs. 24-30) and by the absence of a sharply bounded yellow ventral surface. However, 7. muelleri was not included in recent lists of the snake fauna of Vietnam (Ziegler et al. 2007, Nguyen et al. 2009). We thus follow Wallach (2001) to recognize 7’ siamensis as a distinct species. Future re- search on 7yphlops from southern Vietnam is required to show whether 7: muelleri actually occurs in Vietnam as affirmed by Wallach (2001) or these records are based on misidentified specimens of the closely related 7: siamen- sis. Boiga multomaculata (Boie, 1827) Specimen examined: One adult male (ZFMK 88923), collected by K. D. Le in August 2008 in the bamboo for- est, Cat Loc area, Lam Dong Province (near 11°37’22.5”N 107-17 5SI24Es 3 5emiarsi): Characteristic features: The morphological features of this specimen agree with the descriptions of Bourret (1936), Smith (1943), and Campden-Main (1970): SVL 588 mm; TaL 158 mm; | loreal; | preocular; 2 postocu- lars; 8 SL; 10 IL; 19 MD; 215 VEN; 93 SC; head with two dorsal dark brown stripes, from snout to neck. For colouration in life see Fig. 2b. ©ZFMK New records of snakes from southern Vietnam 1] Fig. 2. Calamaria pavimentata Dumeril & Bibron 1854 Specimen examined: One subadult specimen (ZFMK 88924), collected by K. D. Le in June 2008, in Cat Loc area, Lam Dong Province (near 11°37’22.5”N NO fila ie 2 E135 mais): Characteristic features: The morphological features of this specimen fit the descriptions of Bourret (1936), Smith (1943), and Ziegler et al. (2007): SVL 208 mm; TaL 15.9 mm; snout obtuse; no supranasals; no loreals; no tempo- rals; 13 MD (smooth); 158 VEN; 17 SC; dorsum dark brown or grey; 6 black dorsolateral stripes; yellow collar, narrowing dorsally; tail with 2 yellow cross bands. For colouration in life see Fig. 2 c. Remarks: This is the southernmost record of this species from Vietnam (compared with Nguyen et al. 2009). Bonn zoological Bulletin 60 (1): 9-16 a. Typhlops siamensis (ZFMK 88922); b. Boiga multomaculata (ZFMK 88923); ¢. Calamaria pavimentata (ZFMK 88924); d. Coelognathus flavolineatus (ZFMK 88898). Photographs: E. Galoyan & P. Geissler. Coelognathus flavolineatus (Schlegel, 1837) Specimen examined: One adult female (ZFMK 88898), collected by P. Geissler in April 2009, on the forest floor, in a mixed forest near Bau Sau Lake, Cat Tien National Park, Dong Nai Province (near 11°27°32.9"N 107°20’43.7”E, 167 ma.s.l.). Characteristic features: The morphological traits of this specimen agree with the descriptions of Bourret (1936) and Smith (1943): SVL 1,307 mm; TaL 374 mm; | elon- gated loreal; 1 preocular; 2 postoculars, 8 SL; 10 IL; 19 MD, keeled; 212 VEN; 96 SC, divided; dorsum brown, with a orange, black edged vertebral stripe; vertebral stripe desappearing in the posterior half of body, which is uni- formly dark brown. For colouration in life see Fig. 2d. ©ZFMK Fig. 3. (juvenile, ZFMK 91899); d. Oligodon deuvei(ZMMU NAP-02811); e. Oligodon ocellatus from Cat Tien National Park, Dong Nai Province (e); f. Xenochrophis flavipunctatuss (ZFMK 88914). Photographs: E. Galoyan, P. Geissler, W. Van Devender. Dendrelaphis ngansonensis (Bourret, 1935) Specimens examined: One adult male (ZFMK 88913), collected by P. Geissler on 22 July 2008, in a secondary forest near Bau Sau Lake, Cat Tien National Park, Dong Nai Province (near 11°27°32.9’N, 107°20°43.7”E, 160 m a.s.l.). It was found in the morning, basking on a sun- ny spot on the lava-rock-covered forest floor. Bonn zoological Bulletin 60 (1): 9-16 12 Peter Geissler et al. a. Dendrelaphis ngansonensis (ZFMK 88913); b. Lycodon subcinctus (adult, IEBR A.2010.42); ¢. Lycodon subcinctus Characteristic features: The characters of the specimen fit the descriptions given by Bourret (1935, 1936) and Ziegler & Vogel (1999): SVL 824 mm; TaL 419 mm; 25 dentary teeth; 15 MD; 188 VEN (165-199 in the descrip- tion of Ziegler & Vogel 1999); 153 SC; vertebral dorsals enlarged; dorsum bronze-brown; presence of a distinct black stripe from the posterior margin of the eye to the neck; scales on dorsilateral neck and anterior part of body with bluish margin. For colouration in life see Fig. 3a. ©ZFMK New records of snakes from southern Vietnam 13 Remarks: According to Ziegler & Vogel (1999) and Nguyen et al. (2009) this species has been known only from northern and central Vietnam and Quang Nam Province has been known to be the southernmost locali- ty of the species’ range. Our record from Cat Tien Nation- al Park extends the known distribution about 500 km southwards to the lowland forests of southern Vietnam (see Fig. 3). The distribution gap seen in the map leads to the suggestion that D. ngansonensis 1s also present on or along the Central Highlands as well as the Da Lat Plateau. Like in other localities in Vietnam (Ziegler & Vogel 1999), Den- drelaphis pictus (Gmelin, 1789) can be found in the same habitats with its congener, affirming the species status of D. ngansonensis, which was originally considered as a subspecies of D. pictus. Lycodon subcinctus Boie, 1827 Specimens examined: One adult male IEBR A.2010.42), collected by W. Van Devender in June 2004, near the head- quarters of Cat Tien National Park, Dong Nai Province (near 11°25°19.3”°N, 107°25’42.0”E, 104 m a.s.1.). One juvenile (ZFMK 91899), collected by W. Van Devender on 9 June 2004, near Suoi Rang Ranger Station, Cat Tien National Park, Dong Nai Province. One juvenile (IEBR A.2010.43), collected by W. Van Devender in 2006 in Cat Tien National Park, Dong Nai Province. Characteristic features: The morphological traits of the specimens (data given in the following order: IEBR A.2010.42/ZFMK 91899/IEBR A.2010.43) fit the descrip- tions of Smith (1943); Daltry & Wiister (2002), and Ziegler et al. (2007): SVL 651 mm / 276 mm / 228 mm; TaL 159 mm / 65 mm / 55 mm); head with broad blunt snout; preocular absent; prefrontal and loreal in contact with the eye; 8 SL, 4 suboculars; 17 MD, smooth; 201/202/201 VEN; precloacal scale divided; 71/72/87 SC, divided; juveniles black above with white cross bands (Fig. 3c); adult darker, cross bands absent on posterior part of body (Fig. 3b). Oligodon deuvei David, Vogel & van Rooijen, 2008 Specimens examined: One juvenile specimen (IEBR A.2010.16), collected by local people on 29 May 2004. An adult male (IEBR A.2010.17), collected by P. Moler in May 2004. Another adult male (ZFMK 91226), collect- ed by P. Moler in 2005. These specimens were found near the Headquarters of Cat Tien National Park, Dong Nai Province (near 11°27°32.9"N, 107°20’°43.7”E). One adult male (ZMMU NAP-02811), collected by N. Po- Bonn zoological Bulletin 60 (1): 9-16 yarkov on 8 November 2007, in Cat Tien Village on the eastern bank of the Dong Nai River, Dong Nai Province. Characteristic features: The morphological characters of four specimens from Cat Tien National Park agree with the description given by David et al. (2008b): SVL 283-361 mm; TaL 59.0—68.2 mm; 17 MD, 15 MD before vent; 14 maxillary teeth, the posterior two strongly en- larged; 8—9 IL; 8 SL; no presubocular; a conspicuous, pale yellow vertebral stripe, edged with two darker faint par- avertebral stripes or with lines of dark dots in the paraver- tebral region; 4-5 markings on dorsal head surface: one transverse band across the snout, one sagital blotch be- tween the orbits, two streaks behind the orbit, directed pos- teriorly downwards, and one broad arrow shaped blotch on the neck. In one specimen (ZMMU NAP-02811), three nuchal blotches are fused to one butterfly-shaped mark- ing, and a number of small irregular dots are present on the forehead. Measurements and selected scale counts, in comparison with data of the type specimens provided by David et al. (2008b), are given in Table 1. Colouration: In accordance with Deuve’s (1985) unpub- lished manuscript, our specimens have a grey dorsum. The yellow vertebral stripe is broad on the neck, narrowing backwards, and edged by two dark brown paravertebral stripes. Though not mentioned by Deuve (1985) or David et al. (2008b) our specimens have a dark grey dorsolater- al stripe (aligned dark brown dots in preserved specimens). Large dark blotches are present on upper tail surface in the holotype but they are lacking in our specimens. Ven- ter and lower surface of the tail are pinkish red with nu- merous rectangular and subrectangular blackish-brown spots. These spots are lacking in the posterior half of the tail (Fig. 3d). Remarks: Here we provide the first record for the Cat Tien National Park and the second collection of the species after the description of David et al. (2008b). Our data sup- port the sexual dimorphism observed by David et al. (2008b). Due to the lower number of subcaudals and the shorter tail length, we assume that the juvenile specimen (IEBR A.2010.16) is a female. The specimens in our col- lection slightly differ from the type series by having a higher TL/TaL ratio in males and by having eight supral- abials (this character only occurs in one of 17 specimens examined by David et al. 2008b). For the first time a colour photograph of a living specimen is provided show- ing the coloration of the dorsum as well as the red under- side of the tail, which is used by the snake for defensive display (Fig. 3d). Our observations in the field indicate that this snake species was mostly active during twilight, but was twice observed actively foraging at day time. O. deuvei was found along the riverside and neighbouring ru- ral areas. The stomach of ZMMU NAP-02811 contained ©ZFMK 14 Peter Geissler et al. Table 1. Measurements and selected scale counts of Oligodon deuvei specimens from Cat Tien National Park in comparison with the type series (for abbreviations see Material and Methods). Type specimens JEBR A.2010.17 ZFMK 91226 ZMMU NAP-02811 IEBRA. 2010.16 (juv.) (David et al. 2008) (3) (3) (3) (3) TL (mm) up to 333 535 (2) 342 384.2 328.5 108.8 SVL (mm) up to 275(¢) 302(2) 283 316 271 95.1 TaL (mm) up to 58(3) 51(Q) 59 68.2 57.5 BF TaL/TL (3): 0.158-0.172 (2): 0.132-0.149 0.173 0.178 0.175 0.126 VEN (3): 140-147 (2): 147-155 148 151 144 158 SC (3): 36-73 (Q): 31-38 42 40 40 33 SL 7-8 8/8 8/8 8/8 8/8 IL 8-9 9/9 9/9 9/9 9/9 one juvenile frog (Fejervarya limnocharis) and two unidentified anuran tadpoles. Oligodon ocellatus (Morice, 1875) Specimens examined: Two female specimens (ZFMK 88919-88920), collected by P. Geissler in Cat Tien Na- tional Park, Dong Nai Province. One female specimen (IEBR A.2010.54.), collected by W. Van Devender on 28 May 2004, near Bau Sau Lake, Dong Nai Province (near 12773219 CN 109520743. 7 (E67 murassis): Characteristic features: The morphological traits of the specimens (data given in the following order: ZFMK 88919 / ZFMK 88920 / IEBR A.2010.54) from Cat Tien National Park fit the descriptions of David et al. (2008a): SVL 405 mm / 199 mm / 209 mm; TaL 108 mm / 28.3 mm / 29.7 mm; | presubocular; 8 SL; 9 IL; 19 MD; 173 / 159/165 VEN; precloacal scale undivided; 59 / 59/41 SC; dorsum light brown, with 11—14 dark brown blotch- es, edged with black; dark brown cross bands along the dorsum; head dark brown. For colouration in life see Fig. Se: Remarks: The occurrence in the Cat Tien National Park, Dong Nai Province is the southernmost record of this species in Vietnam (compare with Nguyen et al. 2009). Xenochrophis flavipunctatus (Hallowell, 1861) Specimens examined: One juvenile specimen (IEBR A.2010.46), collected by W. Van Devender in 2006, in Cat Tien National Park, Dong Nai Province. One subadult specimen (ZFMK 88914), collected by P. Geissler on 19 Bonn zoological Bulletin 60 (1): 9-16 May 2009 at the headquarters of the Cat Tien National Park, Dong Nai Province (near 11°25°19.3”N, 107°25°42.0”E, 104 m a.s.1.). Characteristic features: The morphological characters of the specimens (data given in the following order IEBR A.2010.46./ ZFMK 88914) fit the descriptions of Bour- ret (1936), Smith (1943), and Ziegler et al. (2007): SVL 206 mm / 244 mm; TaL 76 mm/ 119 mm; | loreal; | pre- ocular; 3 postoculars; 8 SL; 10 IL; 19 MD, keeled; 136 / 125 VEN; precloacal scale divided; 77 / 85 divided SC; head and dorsum grey; light band from the eye to the an- gle of the mouth, edged with black; parietals with a light blotch, edged with dark brown (in juveniles); lateral sides of neck yellow; flanks with a series of black blotches. For colouration in life see Fig. 3f. DISCUSSION In their review of the herpetofauna of the Cat Tien Na- tional Park, Nguyen & Ho (2002) listed 42 species of snakes, which is almost 21% of the 203 snake species known from Vietnam (Nguyen et al. 2009, Orlov et al. 2010, Ziegler & Nguyen 2010). These records were based on their unpublished reports and examination of voucher specimens. After reviewing the list, we regard one record as questionable. The record of “Homalopsis fasciatus ”, which is based on an unpublished report, is obviously in- correct as this taxon does not exist in current faunal works in this region (Bourret 1936, Nguyen et al. 2009). Besides nine new records based on voucher specimens, the pres- ence of Boiga guangxiensis in the Cat Tien National Park was proven by a photograph taken by W. Van Devender (pers. comm.). Nguyen & Ho (2002) reported the occur- ©ZFMK New records of snakes from southern Vietnam Table 2. List of snake species recorded from Cat Tien National Park (Dong Nai and Lam Dong provinces). Taxon Nguyen Le et al. Nguyen & Ho This study (1988) (1998) (2002) Typhlopidae Ramphotyphlops braminus (Daudin, 1803) x Xx IEBR A.2010.51—A.2010.52 ZFMK 88925 Typhlops siamensis Gunther, 1864 ZFMK 88922 Cylindrophiidae Cylindrophis ruffus (Laurenti, 1768) x x — Pythonidae Python bivittatus Kuhl, 1820 X x x — Python reticulatus (Schneider, 1801) xX x XxX —= Xenopeltidae Xenopeltis unicolor Reinwardt in Boie, 1827 Xx — Colubridae Ahaetulla prasina (Reinhardt, 1827) xX x IEBR A.2010.53 Boiga cyanea (Duméeril, Bibron & Dumeril, 1854) x — Boiga guangxiensis Wen, 1998 Boiga multomaculata (Boie, 1827) ZEMK 88923 Boiga siamensis Nootpand, 1971 x = Calamaria pavimentata Dumeéril, ZFMK 88924 Bibron & Dumeril, 1854 Chrysopelea ornata (Shaw, 1802) x Ke ZFMK 88905-88907 Coelognathus flavolineatus (Schlegel, 1837) ZFMK 88898 Coelognathus radiatus (Boie, 1827) x x x — Dendrelaphis ngansonensis (Bourret, 1935) ZFMK 88913 Dendrelaphis pictus (Gmelin, 1789) x Xx ZFMK 88912 Dryocalamus davisonii (Blanford, 1878) x ZFMK 88929-88930, 91898 IEBR A.2010.46 Gonyosoma oxycephalum (Boie, 1827) x — Lycodon laoensis Ginther, 1864 x ZFMK 88928 Lycodon subcinctus Boie, 1827 IEBR A.2010.42—A.2010.43 ZFMK 91899 Oligodon cinereus (Giinther, 1864) x ZEMK 88921 Oligodon deuvei David, Vogel & van Rooijen, 2008 IEBR A.2010.16—A.2010.17 ZFMK 91226 ZMMU NAP-02811 Oligodon fasciolatus (Gunther, 1864) xe — Oligodon ocellatus (Morice, 1875) IEBR A.2010.54 ZFMK 88919-88920 Ptyas korros (Schlegel, 1837) x x x ZEMK 88915 Ptyas mucosa (Linnaeus, 1758) x x x — Rhabdophis chrysargos (Schlegel, 1837) Xx x TEBR A.2010.45 Rhabdophis subminiatus (Schlegel, 1837) x ZFMK 88908-88909 Sibynophis collaris (Gray, 1853) x — Xenochrophis flavipunctatus (Hallowell, 1861) IEBR A.2010.46, ZFMK 88914 Homalopsidae Enhydris bocourti (Jan, 1865) x x IEBR A. 2010.55—A.2010.56 ZFMK 88926 Enhydris enhydris (Schneider, 1799) x x — Homalopsis buccata (Linnaeus, 1758) xX x ZFMK 88927 Pareatidae Pareas carinatus (Boie, 1828) x ZFMK 88910-88911 Pareas margaritophorus (Jan, 1866) x TEBR A.2010.44 Lamprophiidae (incertae sedis) Psammodynastes pulverulentus (Boie, 1827) x ZFMK 88900-88904 Elapidae Bungarus candidus (Linnaeus, 1758) xe — Bungarus fasciatus (Schneider, 1801) x xX —_— Calliophis maculiceps (Gunther, 1858) x IEBR A.2010.48, ZFMK 91900 Naja kaouthia Lesson, 1831 Ke — Naja siamensis Laurenti, 1768 x x x — Ophiophagus hannah (Cantor, 1836) x x x — Viperidae Calloselasma rhodostoma (Kuhl, 1824) x — Cryptelytrops albolabris (Gray, 1842) Xx x Xx IEBR A.2010.49 ZFMK 91897 Cryptelytrops rubeus Malhotra, Thorpe, Mrinalini & Stuart, 2011 Ke IEBR A.2010.50 (listed as C. macrops by Nguyen & Ho 2002) ZFMK 88916-88918 Viridovipera stejnegeri (Schmidt, 1925) Xx — Bonn zoological Bulletin 60 (1): 9-16 ©ZFMK 16 Peter Geissler et al. rence of Cryptelytrops macrops in the Cat Tien National Park. However, Malhotra et al. (2011) recently described the Ruby-eyed Green Pitviper Cryptelytrops rubeus from southern Vietnam and Cambodia.The distribution of Cryptelytrops macrops is now restricted to Thailand, Cam- bodia, and Laos (Malhotra et al. 2011). Based on our spec- imens with the red eye, we herein confirm the presence of Cryptelytrops rubeus in the Cat Tien National Park. The updated checklist of 51 species of snakes of Cat Tien Na- tional Park is provided in Table 2. For the generic com- position of snakes we follow Pyron et al. (2010). Acknowledgements. We cordially thank Tran Van Thanh (Cat Tien National Park) for his valuable support during the field stud- ies and for issuing respective permits. We gratefully acknowl- edge Le Xuan Canh and Ta Huy Thinh (IEBR) for their support and the loan of specimens. We are grateful to Wayne Van De- vender and Paul Moler for providing scientific material and pho- tographs. We thank Gernot Vogel and Patrick David for their tax- onomic advice. PG thanks Thomas Ziegler, Phung My Trung, Igor Palko, Anton N. Chekanov, Vladimir A. Zryanin, Alexan- der Krohn, Le Duc Khanh, Jendrian Riedel, Bernd Geissler, and Rike Bach for their support during numerous field trips in Cat Tien. We thank Eleanor Sterling and Kevin Koy for providing the map and E. Galoyan for providing photographs. We highly appreciate the comments of Andreas Schmitz and Thomas Ziegler on an earlier version of our manuscript. The field work of PG in Vietnam was partly funded by the Alexander Koenig Gesellschaft and the German Herpetological Society (DGHT). REFERENCES Bourret R (1935) Notes herpétologiques sur |’Indochine fran- caise. XI. Sur quelques serpents recoltés en 1934. Bulletin général de l’instruction publique. 1934/35 (9. Mai): 289-296 Bourret R (1936) Les serpents de I’Indochine, II, Catalogue sys- tématique déscriptif. H. Basuyau, Toulouse Cambden-Main S (1970) A fieldguide to the Snakes of South Vietnam. Division of Reptiles and Amphibians United States National Museum Smithsonian Institut, City of Washington Daltry JC, Wiister W (2002) A new species of Wolf Snake (Ser- pentes: Colubridae: Zycodon) from the Cardamom Mountains, Southwestern Cambodia. Herpetologica 58: 498-504 David P, Vogel G, Pauwels OSG (2008a) A new species of the genus Oligodon Fitzinger, 1826 (Squamata: Colubridae) from Southern Vietnam and Cambodia. Zootaxa (1939): 19-37 David P, Vogel G, van Rooijen J (2008b) A revision of the Oligodon taeniatus (Gunther, 1861) group (Squamata: Colu- bridae), with the description of three new species from the In- dochinese Region. Zootaxa 1965: 1-49 Deuve J (1970) Serpents du Laos. Mémoires de |’ office de la recherche scientifique et technique outre-mer, Paris 39: 1-251 Deuve J (1985) Serpents du Laos. Dessins en couleurs et doc- uments manuscrits. Unpublished manuscript notes, deposit- ed in the library of the ,, Taxonomie-collection- Reptiles & Am- phibiens* unit, Muséum National d’Histoire Naturelle, Paris Geissler P, Nazarov R, Orlov NL, Bohme W, Phung TM, Nguyen QT, Ziegler T (2009) A new species of the Cyrtodactylus ir- Bonn zoological Bulletin 60 (1): 9-16 regularis complex (Squamata: Gekkonidae) from southern Vietnam. Zootaxa 2161: 20-32 Giinther ACLG (1864) The reptiles of British India. Ray Soci- ety, London Le XC, Hoang MK, Le DT, Ho TC, Hoang VT, Nguyen VT, Nguyen TV, Tran VD (1998) Results of zoological survey in Cat Loc Nature Reserve, Cat Tien District, Lam Dong Province. Unpublished report of IEBR, Hanoi Malhotra A, Thorpe RS, Mrinalini, Stuart BL (2011) Two new species of pitviper of the Genus Cryptelytrops Cope 1860 (Squamata: Viperidae: Crotalinae) from Southeast Asia. Zootaxa 2757: 1—23 Nguyen QT (1988) Preliminary list of Reptiles and Amphibians in Nam Cat Tien forest reserve. Garrulax 5: 8-9 Nguyen VS, Ho TC (1996) A checklist of reptiles and amphib- ians of Vietnam. Science and Technology Publishing House, Hanoi (in Vietnamese) Nguyen VS, Ho TC (2002) Species composition of the herpeto- fauna recorded in the Cat Tien National Park. Journal of Bi- ology 24: 2-10 (in Vietnamese) Nguyen VS, Ho TC, Nguyen QT (2009) Herpetofauna of Viet- nam. Edition Chimaira, Frankfurt am Main Orlov NL, Nguyen QT, Nguyen TT, Ananjeva NB, Ho TC (2010) A new species of the genus Calamaria (Squamata: Ophidia: Colubridae) from Thua Thien- Hue Province, Vietnam. Russ- ian Journal of Herpetology 17: 236-242 Polet G, Ling S (2004) Protecting mammal diversity: opportu- nities and contraints for pragmatic conservation management in Cat Tien National Park, Vietnam. Oryx 38: 1-11 Pyron RA, Burbrink FT, Colli GR, Montes de Oca AN, Vitt LJ, Kuczynski CA, Wiens JJ (2010) The phylogeny of advanced snakes (Colubroidea), with discovery of a new subfamily and comparison of support methods for likelihood trees. Molec- ular Phylogenetics and Evolution: doi:10.1016/j.ym- pev.2010.11.006 Smith MA (1943) The fauna of British India, Ceylon and Bur- ma, including the whole of the Indo-Chinese subregion. Rep- tiles and Amphibians, Vol. 3. Serpentes. Taylor and Francis, London Tirant G (1885) Notes sur les reptiles et les batraciens de la Cochinchine et du Cambodge. Imprimérie du Gouvernement, Saigon Wallach V (2001) Typhlops roxanae, a new species of Thai blind snake from the 7: diardii species group, with a synopsis of the Typhlopidae of Thailand (Serpentes: Scolecophidia). Raffles Bulletin of Zoology 49: 39-49 Ziegler T, Hendrix R, Vu NT, Vogt M, Forster B, Dang NK (2007) The diversity of a snake community in a karst forest ecosystem in the central Truong Son, Vietnam, with an iden- tification key. Zootaxa 1493: 1-40 Ziegler T, Nguyen QT (2010) New discoveries of amphibians and reptiles from Vietnam. Bonn zoological Bulletin 57: 137-147 Ziegler T, Vogel G (1999) On the knowledge and specific sta- tus of Dendrelaphis ngansonensis (Bourret, 1935) (Reptilia: Serpentes: Colubridae). Russian Journal of Herpetology 6: 199-208 Received: 28.10.2010 Accepted: 09.03.2011 Corresponding editor: F. Herder ©ZFMK Volume 60 Bonn zoological Bulletin Issue | pp. 17-24 | Bonn, May 2011 Description of a new species of the genus Dendrelaphis Boulenger, 1890 from Myanmar (Squamata: Serpentes: Colubridae) Gernot Vogel!*& Johan van Rooijen? | Society for Southeast Asian Herpetology, Im Sand 3, D-69115 Heidelberg, Germany; E-mail: Gernot. Vogel@t-online.de 2 Netherlands Centre for Biodiversity Naturalis, section Zoological Museum Amsterdam, Mauritskade 61, 1092 AD Amsterdam, The Netherlands 3 Tulpentuin 313, 2272 EH, Voorburg, The Netherlands; E-mail: j1.van.rooijen@hetnet.nl «corresponding author. Abstract. The population systematics of the colubrid snake so far referred to as Dendrelaphis gorei (Wall, 1910) were investigated by carrying out a multivariate analysis of geographic variation. The results reveal the existence of two dis- tinct and apparently disjunct phenotypes. These phenotypes are assumed to represent independent evolutionary lineag- es. The first lineage corresponds with D. gorei which inhabits the Assam Valley and adjoining Himalayan foothills. How- ever, the name Dendrophis gorei Wall, 1910 is actually a junior synonym of Dendrelaphis biloreatus Wall, 1908. Con- sequently, the latter name is adopted for this lineage. The second lineage, described in this paper as a new species, in- habits the South Myanmar lowlands and neighbouring mountain chains. It is distinguished from D. biloreatus on the ba- sis of its higher ventral count (203—212 versus 190-199), its undivided anal shield (divided in D. biloreatus) and its high- er average number of anterior temporal shields (usually 2 versus usually 1). Key words. Dendrelaphis sp. n., Dendrelaphis gorei syn. n., Dendrelaphis biloreatus, Myanmar, analysis of geograph- ic variation. INTRODUCTION The colubrid snakes of the genus Dendrelaphis Boulenger, 1890 are widely distributed, ranging from Pakistan in the West to the northern and eastern coast of Australia in the East and South and to southern China in the North (Ziegler & Vogel 1999). Members of the genus Dendrelaphis are slender, diurnal species that are predominantly arboreal and feed mainly on lizards and amphibians. Boulenger (1894), Wall (1921), Meise & Henning (1932), Mertens (1934) and Smith (1943) have in turn worked on the systematics of this genus. Nevertheless, their cumu- lative effort did not result in an unambiguous and com- plete taxonomy of this genus, a fact that was underlined by the recent descriptions and revalidations of several species (Vogel & Van Rooijen 2007, 2008, 2011; Van Rooyen & Vogel 2008a, 2008b, 2008c, 2009). Dendrelaphis gorei (Wall, 1910) has been reported to oc- cur in Northeast India and Myanmar. It is one of the least known members of its genus and is scarcely represented in museum collections. In recent years however, new ma- terial has become available through the Myanmar Herpeto- logical Survey, a collaborative effort of the California Academy of Sciences, Smithsonian Institution and the Forest Department, Ministry of Forestry, Myanmar (e.g. Wogan et al. 2008). In this paper, we investigate the pop- ulation systematics of D. gorei by performing a multivari- ate analysis of geographic variation based on morpholog- ical data. Furthermore, the name Dendrelaphis biloreatus Wall, 1908 is evaluated. Smith (1943) synonymized D. biloreatus with D. gorei (Wall 1910) as he could not find any differences between the two species apart from the di- vided loreal shield in D. biloreatus. However, according to the rule of priority (Art. 23, ICZN, 1999), the name D. biloreatus should be adopted for this species. MATERIALS AND METHODS Eleven museum specimens were examined for this study. In addition, data on three additional specimens were tak- en from Wall (1908, 1910). For each examined specimen, 23 characters including aspects of colour pattern, body proportions and scalation were recorded (Table 1). Eye- diameter and distance eye-nostril were measured with a slide calliper to the nearest 0.1 mm. These measurements were made on the left and right side and were subsequent- ly averaged. Snout-vent length was measured to the pos- 18 Gernot Vogel & Johan van Rooijen terior margin of the anal plate by marking the length on a piece of string and subsequently measuring the position of the mark to the nearest 0.5 cm. Tail-length was meas- ured to the nearest 0.5 cm by straightening the tail against a ruler. The number of ventrals was counted using DowI- ing’s method (1951). Subcaudals were counted on one side, the terminal scute was excluded. The first sublabial was defined as the scale that starts between the posterior chin shield and the infralabials and that borders the infral- abials (see Peters 1964, fig. 7). The last infralabial was defined as the infralabial still covered completely by the last supralabial. The posterior most temporal scales were defined as the scales of which more than half of the area lies in front of an imaginary line that runs from the apex of the last supralabial to the posterolateral corner of the parietal. Geographic coordinates were taken directly from the field notes or were obtained by translating locations to coordi- nates. Relevant variables were included in a Principal Components Analysis (PCA, e.g. Cramer 2003) in order to reduce the dimensions of the dataset. The resulting PCA-scores were then plotted against longitude and lati- tude in order to visualize the pattern of geographic vari- ation. Confirmatory geographic analyses of the PCA-score were carried out by Analysis of Covariance (ANCOVA; Maxwell & Delaney 1990; Norusis & SPSS 1993) using longitude and latitude as covariates and phenotype as fac- tor. Normality of the PC-scores was first tested by means of Kolmogorov-Smirnovy tests. All statistical analyses were carried out with the software SPSS (2006; SPSS for Windows. Release 14.0.2. Chica- go: SPSS Inc.). Table 1. List of morphometric, scalation and coloration characters used in this study and their abbreviations. Abbreviation Character Morphometrics EYED Horizontal diameter of the eye EYEN Distance from centre of the eye to posterior border of the nostril TAIL Tail-length WSNOUT Width of the snout measured at the position of the nostrils LHEAD Head-length measured from the tip of the snout to the rear of the jaw SVL Snout-vent length Scalation VENT Number of ventrals SUBC Number of subcaudals DORI Number of dorsal scale rows | head-length behind the head DOR2 Number of dorsal scale rows at the position of the middle ventral DOR3 Number of dorsal scale rows | head-length before the tail SUBL Number of infralabials touched by the first sublabial (L+R) SLI Number of supralabials (L+R) SL2 Number of supralabials touching the eyes (L+R) LOR Number of loreals (L+R) INFR Number of infralabials (L+R) ATEMP Number of anterior temporals (L+R) TEMP Number of temporals (L+R) POC Number of postoculars (L+R) VERT Vertebral scales smaller than (0) or larger than (1) scales of the first dorsal row AN Anal shield divided (0) or undivided (1) Coloration TSTRIPE Postocular stripe absent (0), rudimentary (1), present (2) LSTRIPE Ventrolateral stripe absent (0), present (1) The collected data were used to carry out an analysis of geographic variation. The objective of this analysis was to enable differentiation between clinal variation and phe- netic discontinuities, the latter being evidence for lineage separation (e.g. Lenk & Wister 1999; De Queiroz 2007). Bonn zoological Bulletin 60 (1): 17-24 Material examined. S-037, Guwahati, Assam; BMNH 1940.3.4.24, Assam; BMNH 1940.3.4.25, Samagooting, Assam; BMNH 1946.1.10.30, Jaipur, Assam (syntype Dendrophis gorei Wall, 1910); CAS 208429, Myanmar, Bago Division, Bago Yoma, Sein Yay Camp, 18 51 21.636 ©OZFMK A new Dendrelaphis from Myanmar 19 N, 96 10 21.324 E; CAS 211939, Myanmar, Ayeyarwady Division, vic Mwe Hauk Village (16 16 39.2 N, 94 45 32.5 E); CAS 222114-222115, Myanmar: Bago Division, Aok Twin Township, Ka Baung Reserve, Sein Ye Camp, Block 120, 18 51 16.1 N, 96 10 23.6 E; CAS 222339, Myan- mar: Chin State, Min Dat Township, Min Dat Township, Nat Ma Taung National Park, 21 21 14.9 N, 93 56 08.3 E; CAS 234880, Myanmar,: Chin State, Mindat Township, Mindat District, 21 26 43.5 N, 94 00 24.0 E; CAS 244037, Myanmar, Sagaing Division, Leha and Khante township boundary, Nana Sa Laing camp, 26 07 23.1 N, 95 32 24.6 B Museum abbreviations. BMNH: Natural History Muse- um, London, Great Britain. CAS: Collection of the Cali- fornia Academy of Sciences, USA. S: Saibal Senguptas Collection, Guwahati, Assam RESULTS Statistics Visual inspection of the data suggested a striking differ- entiation between specimens from Assam (NE India) and Northwest Myanmar on the one hand and specimens from the remainder of Myanmar on the other hand. One phe- notype was characterized by a high number of ventral counts, an undivided anal shield and usually two anteri- or temporal shields whereas the other was characterized by a low number of ventrals, a divided anal shield and usu- ally one anterior temporal shield. These characters were included in a PCA. The first component explained 92% of the total variance. This demonstrates that these char- acters covary strongly and thus form a suite of characters that sharply differentiates between the two phenotypes. The individual scores on this component were plotted 1 1 : 1 16 ' 0.5 = O oC fi. 2 0.5 2 2 Ae yj 2 2 specie ll | | es eae T aay 5 ee 2 ibe eo aes 20 lon; earl 2 Stud % 98 latitude Fig. 1. PCA-scores based on a PCA of the characters VENT, AN and ATEMP against longitude and latitude. 1 Phenotype 1, 2 phenotype 2, 2b= type D. biloreatus Wall, 1908; 2a = syntype D. gorei Wall, 1910. Bonn zoological Bulletin 60 (1): 17-24 ©OZFMK 20 against latitude and longitude to visualize the pattern of geographic variation. In figure 1, data-points correspon- ding with the type of Dendrelaphis biloreatus and one of the two syntypes of Dendrophis gorei are indicated for a later discussion of the appropriate nomenclature. A phe- netic discontinuity is in evidence, the difference in PCA- scores between the phenotypes being highly significant (ANCOVA, df=1,11; P<0.0001). Subsequent inclusion in the model of longitude and latitude in the form of covari- ates demonstrated that geographic coordinates had no added explanatory power (ANCOVA, df=1,9; P=0.9 and df=1,9; P=0.4 respectively). Thus, there is no evidence of clinal variation, the transition from one phenotype to the other apparently being sudden rather than gradual. The sudden phenetic transition represents strong evidence for lineage separation. Thus, the two distinguished phe- notypes are here interpreted as representing distinct evo- lutionary sister-lineages. An evaluation of the characters separating the two phenotypes supports this view. First, one phenotype exhibits an undivided anal shield where- as the other has a divided anal shield. An undivided anal shield is an exceptional character in the genus Den- drelaphis. The only congeneric species that exhibits this character (in roughly 40% of the specimens) is Den- Gernot Vogel & Johan van Rooien drelaphis proarchos (Wall, 1909) (Vogel & van Rooijen 2011). We examined more than 600 museum specimens of this genus, representing 24 species and subspecies. On- ly a single specimen (beside those identified as D. proar- chos and the material presented in this study) was found to have an undivided anal shield. Second, the two pheno- types differ substantially in the number of ventral scales, the difference being 19 on average (211 in phenotype 1, 192 in phenotype 2). This is a relatively large difference when compared to established interspecific differences. For instance, the difference in ventral count between D. formosus on the one hand and D. kopsteini and D. cyanochloris on the other hand is 5 and 19 respectively (Vogel & Van Rooyen 2007). The difference between D. haasi and D. pictus, D. pictus andamanensis, D. humayu- ni, D. tristis 1s 3, 20, 10 and 14 respectively (Van Rooi- jen & Vogel, 2008a). Finally, the difference between D. chairecacos on the one hand and D. tristis and D. schokari on the other is 10 and 12 respectively (Van Rooijen & Vo- gel 2009). The third difference between the two pheno- types is the number of anterior temporal scales. Pheno- type | usually has two anterior temporal scales whereas phenotype 2 has a single anterior temporal in the major- ity of cases. This character alone already leads to a cor- rect identification in 85% of the cases. Table 2. Comparison between the type of Dendrelaphis biloreatus and six specimens of the species so far referred to as Den- drelaphis gorei. type Dendrelaphis biloreatus Dendrelaphis gorei (n=6) Ventrals Subcaudals Relative tail-length Supralabials Supralabials touching the eye Anterior temporals Total number of temporals Poctoculars Loreals Dorsal rows at midbody Divided anal shield Faint ventrolateral line, not bordered by black lines Postocular stripe that covers the whole temporal region and extends onto the neck Total length vertebrals strongly enlarged Ground color brown Maxillary teeth 192 190-199 147 144-152 0.34 0.34—0.35 9 8-9 3 2-3 1 1-2 8 7-12 2 1-2 2 1 1 13 yes yes yes yes yes yes 70.0 53.5—90.0 yes yes yes yes 21 20 (n=1) Bonn zoological Bulletin 60 (1): 17-24 ©ZFMK A new Dendrelaphis from Myanmar 21 Fig. 2. Dendrelaphis walli sp. n., holotype (CAS 234880). Nomenclature Two available names, Dendrophis gorei Wall, 1910 and Dendrelaphis biloreatus Wall, 1908 are relevant in the context of this study. In figure 1, data pertaining to a syn- type of D. gorei and the type of D. biloreatus are indicat- ed. It is evident that neither of these types represents phe- notype 1 (South Myanmar and neighbouring mountain chains). Thus, we here describe phenotype | as a new species. Phenotype 2 (Assam and Northwest Myanmar) is currently known as D. gorei (Wall, 1910), but we here adopt the older name Dendrelaphis biloreatus Wall, 1908 to represent this lineage. Dendrelaphis biloreatus was de- scribed by Wall on the basis of a single specimen. Smith (1943) synonymized D. biloreatus with D. gorei as he could not find any difference between the two species apart from the divided loreal shield in D. biloreatus. In table 2, a more detailed comparison is provided between Bonn zoological Bulletin 60 (1): 17-24 the morphological characteristics of the type of D. bilore- atus and six specimens of D. gorei. With the exception of the loreal shield, the values pertaining to the type of D. biloreatus all fall within the range of D. gorei. Thus, the results agree with Smith’s view that the type of D. bilore- atus represents an exceptional specimen with a split lo- real shield within D. gorei. In the context of a revision of the taxonomy of this genus, the authors have come across several specimens of Dendrelaphis with either a double or an absent loreal shield. Furthermore D. biloreatus has a rather long and slender postnasal scale which looks like second loreal shield at the first glance. So indirectly the name makes some sense. Consequently, the two names in- deed refer to the same species. According to the rule of priority (ICZN, 1999, art. 23.1), the name Dendrelaphis biloreatus should actually be adopted for this species. OZFMK nN ie) Gernot Vogel & Johan van Rooyen Table 3. Morphological and coloration characters of the types of Dendrelaphis walli sp. n. Collection N° CAS 234880 CAS 208429 CAS 211939 CAS 222114 CAS 222339 CAS 222115 status holotype paratype paratype paratype paratype paratype Sex f - m m m f Snout-vent length (cm) 48.5 ~ 49.0 54.5 61.5 595 Tail-length (cm) 22.5 ~ 25.5 29.5 29.0 31.0 Head-length (mm) 13.5 - 15.5 15.5 17.0 17.0 Eye-diameter (mm) 3.1 — 3.5 35) 3.9 3.5 Snout-width 2.8 ~ 3.2 3.4 i) 3.3 Ventrals 213 211 203 210 212 212 Subcaudals 145 - 148 163 147 159 Dorsal formula 13-13-11 ~ 13-13-11 13-13-11 13-13-9 13-13-11 Temporal formula 2222/2122) 9 222/222 222/212 212/112 2212/222 222/212 Supralabials 8/8 8/8 8/8 8/8 8/8 8/8 Supralabials touching the eye 4,5/4,5 4,5/4,5 4,5/4,5 4,5/4,5 4,5/4,5 4,5/4,5 Infralabials 10/10 - 10/10 2/9 10/10 11/10 Infralabials touched 6,7/6,7 _ 6,7/6,7 6,7/5,6 6,7/6,7 6,7/6,7 by first sublabial Loreals 1/1 1/1 1/1 1/1 1/1 1/1 Postoculars 2/2 3/3 2/2 2/2 2/2 2/2 Number of scales bordering 4 = 5 5 6 5 the parietal scales Vertebrals larger than yes ~ yes yes yes yes dorsals of the first row Anal shield entire yes yes yes yes yes yes Light ventrolateral stripe yes = yes yes yes yes which is faint and is not bordered by black lines Postocular stripe broad, yes — yes yes yes yes covering the whole temporal region and extending onto the neck Taxonomy Dendrelaphis walli sp. n. (Fig. 2) Dendrelaphis gorei (Wall, 1921) (part.) Holotype. CAS 234880, adult female from Myanmar (Chin State: Mindat township, Mindat District, 21 26 43.5 N, 94 00 24.0 E, 3582 ft), collected by A.K. Shein and T. Nyo, 23 Aug 2005. Paratypes. CAS 208429, 222114222115, CAS 222339. CAS. 211939, CAS Bonn zoological Bulletin 60 (1): 17-24 Diagnosis. A species of Dendrelaphis, characterized by the combination of: 1) 13 dorsal scale rows at midbody; 2) strongly enlarged vertebral scales; 3) 203—213 ventrals; 4) 145-163 subcaudals; 5) 8 supralabials; 6) supralabials 4 and 5 bordering the eye; 7) 2 anterior temporals in the majority of specimens; 8) a short sublabial that touches 2 infralabials; 9) an undivided anal shield; 10) a black pos- tocular stripe that covers the majority of the temporal re- gion and extends onto the neck; 11) black, oblique bars on the neck region; 12) a pale ventrolateral line, not bor- dered by black lines. ©ZFMK A new Dendrelaphis from Myanmar 23 Table 4. Diagnostic differences between Dendrelaphis walli sp. n. and Dendrelaphis biloreatus Wall, 1908. Dendrelaphis walli n. sp. (n=7) Dendrelaphis biloreatus (n=7) Ventrals 211 (203-213) Anterior temporals anal shield entire usually 2 (92% of specimens) 192 (190-199) usually 1 (86% of specimens) divided Description of the holotype. Adult female; body very slender; snout-vent length 48.5 cm; tail-length 22.5 cm;head distinct from neck; head-length 13.5 mm; snout- width 2.8 mm; pupil round; eye-diameter 3.1 mm; distance eye-nostril 3.1 mm; 213 ventrals; 145 subcaudals; dorsal scales in 13-13-11 rows; 8 supralabials, 4'* and 5‘ bor- der the eye; 10 infralabials, infralabials 1—5 touch the first chinshield, infralabials 5 and 6 touch the second chin- shield; | preocular; 2 postoculars; | loreal; temporal for- mula 2:2:2:1:2 (L), 2:1:2:2 (R); first sublabial touches in- fralabials 6 and 7; vertebrals strongly enlarged, with straight posterior margin, width of the vertebral scale at the position of the middle ventral scale 2.4 mm; anal en- tire; parietal scales bordered posteriorly by 4 scales; ground color brownish; supralabials and throat white; some black spots on supralabials 2-4, the loreal and the preocular; a black postocular stripe starts behind the eye, covers the majority of the temporal region, and extends onto the neck where it breaks up into narrow, black oblique bars which fade away further posteriorly; a faint ventro- lateral line is present, covering the first dorsal row and lower half of the second dorsal row; the ventrolateral line is not bordered by black lines; belly whitish. Bassein RANGOON ». Maui ee iY THAILAND | Lf De a | Fig. 3. Currently known distribution of D. biloreatus (white b) and D. walli sp. n. (black W). Bonn zoological Bulletin 60 (1): 17-24 Variation. Table 3 provides data regarding the types of D. walli sp. n. Comparison with congeneric species. Dendrelaphis wal- li sp. n. differs from all congeners, except Dendrelaphis biloreatus, Dendrelaphis caudolineatus and Dendrelaphis caudolineolatus, by its dorsal formula of 13-13-11/9. It dif- fers from Dendrelaphis caudolineatus in its strongly en- larged vertebral scales (not enlarged in D. caudolineatus), its much more slender body (stout in D. caudolineatus), its undivided anal shield, the number of supralabials (9 vs. 8 in D. walli sp. n.) and in its coloration. It differs from Dendrelaphis caudolineolatus in the number of ventral scales (149-175 vs. 203-213 in D. walli sp. n.) and sub- caudal scales (111-129 vs. 145-163 in D. walli sp. n.) and its undivided anal shield. Differences between D. walli sp. n. and D. biloreatus are given in Table 4. Sexual dimorphism. Females have a wider snout than males (ANCOVA, df=1,5, P=0.004). In addition, there is some evidence that females have a larger eye than males (ANCOVA, df=1,5, P=0.08). No evidence of other sexu- al dimorphisms was found though this may be due to lack of statistical power (e.g. Streiner 1990). Distribution. According to currently known locality records, D. biloreatus inhabits the Assam Valley and ad- joining Himalayan foothills whereas D. wailli sp. n. inhab- its the South Myanmar lowlands and neighbouring mountain chains. Thus, the two species appear to exhib- it a disjunct distribution (figure 3). However, this may be due to collecting gaps. In reality, one or both of these species may inhabit the intermediate area. On the other hand, the Myanmar Herpetological Survey did collect in the central and northern parts of Myanmar, which did not yield a specimen of either species, with the exception of one D. biloreatus (CAS 244037) near the border with As- sam, within the known range of this species. If these species indeed occur allopatrically, the central dry zone of Myanmar and adjoining mountain ranges may be the geographic barrier that separates these species and that may have enabled the independent evolution of the two. Etymology. This species is dedicated to Major Frank Wall (1868-1950), in recognition of his outstanding work on the genus Dendrelaphis. ©ZFMK 24 Gernot Vogel & Johan van Rooijen Acknowledgements. We thank J. V. Vindum (CAS, San Fran- cisco, USA), C. J. McCarthy (BMNH, London, United King- dom), R. Vonk and H. Praagman (ZMA, Amsterdam, The Netherlands), Saibal Sengupta and Jayaditya Purkayastha (In- dian collections) for letting us examine preserved specimens. REFERENCES Boulenger GA (1894) Catalogue of the Snakes in the British Mu- seum (Natural History). Volume II., Containing the Conclu- sion of the Colubridae Aglyphae. London: Taylor & Francis Cramer D (2003) Advanced quantitative data analysis. Philadel- phia: Open University Press De Queiroz K (2007) Species concepts and species delimitation. Systematic Biology 56: 879-886 Dowling HG (1951) A proposed standard system of counting ventrals in snakes. British Journal of Herpetology 1: 97-99 ICZN — International Commission on Zoological Nomenclature (1999) International Code of Zoological Nomenclature, Fourth Edition adopted by the International Union of Biolog- ical Sciences. The International Trust for Zoological Nomen- clature, London Lenk P, Witister W (1999) A multivariate approach to the sys- tematics of Italian Ratsnakes of the Elaphe longissima com- plex (Reptilia, Colubridae): revalidation of Camerano’s Cal- lopeltis longissimus var. lineata. Herpetological Journal 9: 153-162 Maxwell SE, Delaney HD (1990) Designing experiments and analyzing data. California: Wadsworth Inc. Meise W, Henning W (1932) Die Schlangengattung Dendrophis. Zoologischer Anzeiger 99: 273-297 Mertens R (1934) Die Schlangengattung Dendrelaphis Boulenger in systematischer und zoogeographischer Beziehung. Archiv ftir Naturgeschichte, Berlin (N. F.) 3(2): 187-204 Norusis & SPSS Inc. (1993) SPSS Advanced Statistics, SPSS, Chicago Peters JA (1964) Dictionary of herpetology: a brief and mean- ingful definition of words and terms used in herpetology. New York: Hafner Publ. Co. Smith MA (1943) The fauna of British India, Ceylon and Bur- ma. Reptilia and Amphibia. Vol. 3 Serpentes. London: Tay- lor & Francis Streiner DL (1990) Sample size and power in psychiatric re- search. Canadian Journal of Psychiatry 35: 616-620 Bonn zoological Bulletin 60 (1): 17-24 Van Rooijen J, Vogel G (2008a) A review of the Dendrelaphis pictus complex (Serpentes: Colubridae) — I: Description of a sympatric species. Amphibia-Reptilia 29: 101-115 Van Rooijen J, Vogel G (2008b) A new species of Dendrelaphis (Serpentes: Colubridae) from Java, Indonesia. The Raffles Bul- letin of Zoology 56: 189-197 Van Rooijen J, Vogel G (2008c) An investigation into the tax- onomy of Dendrelaphis tristis (Daudin, 1803): revalidation of Dipsas schokari (Kuhl, 1820) (Serpentes, Colubridae). Con- tributions to Zoology 77: 33-43 Van Rooyen J, Vogel G (2009) A multivariate investigation in- to the population systematics of Dendrelaphis tristis (Daudin, 1803) and Dendrelaphis schokari (Kuhl, 1820): revalidation of Dendrophis chairecacos Boie, 1827 (Serpentes: Colubri- dae). The Herpetological Journal 19: 193-200 Vogel G, Van Rooijen J (2007) A new species of Dendrelaphis (Serpentes: Colubridae) from Southeast Asia. Zootaxa 1394: 25-45 Vogel G, Van Rooijen J (2008) A review of the Dendrelaphis pictus complex (Serpentes: Colubridae) — 2: the eastern forms. Herpetozoa 21: 3-29 Vogel G, Van Rooijen J (2011, in press) Contributions to a re- view of the Dendrelaphis pictus (Gmelin, 1789) complex (Ser- pentes: Colubridae) — 3. the Indian forms, with the descrip- tion of a new species from the Western Ghats. Journal of Her- petology 45 Wall F (1908) Two new snakes from Assam. Journal of the Bom- bay natural History Society 18: 272-274 Wall F (1910) Notes on snakes collected in Upper Assam. Part 2. Journal of the Bombay Natural History Society 19: 825-845 Wall F (1921) Remarks on the Indian Species of Dendrophis and Dendrelaphis. Records of the Indian Museum 22: 151-162 Wogan GOU, Vindum JV, Wilkinson JA, Koo MS, Slowinsky JB, Win H, Thin T, Kyi SW, Oo SL, Lwin KS, Shein AK (2008) New country records and range extensions for Myan- mar amphibians and reptiles. Hamadryad 33: 83-96 Ziegler T, Vogel G (1999) On the knowledge and specific sta- tus of Dendrelaphis ngansonensis (Bourret, 1935) (Reptilia: Serpentes: Colubridae). Russian Journal of Herpetology 6: 199-208 Received: 20.08.2010 Accepted: 20.02.2011 Corresponding editor: P. Wagner ©ZFMK Bonn zoological Bulletin Volume 60 Issue | pp. 25-34 Bonn, May 2011 Studies on African Agama IX. New insights into Agama finchi Bohme et al., 2005 (Sauria: Agamidae), with the description of a new subspecies Philipp Wagner!2*, Wolfram Freund!, David Modry34, Andreas Schmitz5 & Wolfgang Bohme! | Zoologisches Forschungsmuseum Alexander Koenig, Adenauerallee 160, D-53113 Bonn, Germany 2 Department of Biology, Villanova University, Villanova, Pennsylvania 19085, USA 3 Dept. of Parasitology, Faculty of Veterinary Medicine, University of Veterinary and Pharmaceutical Sciences Brno, Palackeho 1-3, 612 42 Brno, Czech Republic + Biology Center, Institute of Parasitology, Academy of Sciences of the Czech Republic, BraniSovska 31, 370 05 Ceské Budéjovice, Czech Republic 5 Department of Herpetology and Ichthyology, Muséum d’histoire naturelle, C.P. 6434, CH-1211 Geneva 6, Switzerland “corresponding author: philipp.wagner.zfmk@uni-bonn.de. Abstract. We present new information on the distribution and morphology of Agama finchi from eastern Africa. For the first time, material from three different populations (including the type locality) was available and the question of a pos- sible subspecies in Uganda was positively answered. Based on the distribution pattern of Agama species groups, a gen- eral distribution pattern is discussed. Key words. Reptilia, Sauria, Agamidae, Agama finchi, Agama finchi ssp. n., Africa, Kenya, Uganda, DR Congo, Ethiopia. INTRODUCTION Cursorily, the taxonomy and relationships between east- ern African Agama species were thought to be well known, but after intensive research on these species, it became ob- vious that species limits were underestimated and sever- al taxa were described, revalidated or regarded as syn- onyms (Bohme et al. 2005; Wagner 2007; Wagner et al. 2008a, 2008b; Wagner 2010). Most Agama species occur- ring in eastern Africa seem to be part of a monophyletic group (Wagner et al. unpubl. data). However, the only member of the Agama agama species group occurring in eastern Africa is still 4. finchi (Leaché et al. 2009). This species was described by Bohme et al. (2005) from Mal- aba in western Kenya close to the Ugandan border. These authors also mentioned a population from Murchison Falls in Uganda, but specimens from this population were on- ly known from photographs. They differed from topotyp- ical Agama finchi in the colouration of the forelimbs and until recently, it was not possible to confirm these photo- graphs with voucher specimens. Additionally, further in- formation given by Bohme et. al. (2005) show the occur- rence of A. finchi at the “Yale River in Uganda’, but it was not possible to trace this locality. Probably the locality refers to the western Kenyan Yala River, which was in the ‘Uganda’ province of East Africa before 1926 (pers. comm. Stephen Spawls, 25.1.2011). A. finchi is a small lizard and is characterized by its bi- coloured red and black tails, scarlet-red heads and fore- limbs and the velvet-black bodies of adult males. The throat colouration is uniform pale reddish, with a pattern of longitudinal dark reddish lines. The females are differ- ent in colouration from other Agama species as they pos- sess a uniform brown body with reddish to yellow dorso- lateral bands. In both sexes, a white to yellowish supral- abial line is usually obvious. Nevertheless, differences in colouration between the specimens from Uganda and topo- typical A. finchi are obvious, since the former possess body-like coloured forelimbs. However, without vouch- er specimens, it was so far not possible to assess the sta- tus of this population. Currently, only few Agama species are known from Ugan- da. Agama lionotus elgonis Lonnberg, 1922 is known to occur in the Mt. Elgon region, and most probably Agama finchi, described from an area close to the border to Ugan- da, should occur there. Spawls et al. (2002) also mentioned a wide distribution of Agama agama in Uganda, but these records should be referred to Agama lionotus since A. a. agama is restricted to Central Africa (Wagner et al. 2009). 26 Philipp Wagner et al. A Agama f. leucerythrolaema ssp. n. Uganda: Murchison Falls ZFMK 88809, holotype B Agama f. finchi Kenya: Malaba NMK L/2716, topotype Ec Agama lebretoni Cameroon: Mamfe ZFMK 87689, holotype D Agama paragama Cameroon: Waza ZFMK 15244 E Agama spec.CAR-1 Central African Republic: Koumbala ZFMK 40252 F Agama spec. CHAD Chad: Lake Chad MNHN uncatalogued G Agama sp. n. DR Congo: Nagero ZFMK 51578 Fig. 1. Throat and left head side of male Agama species. Bonn zoological Bulletin 60 (1): 25-34 OZFMK Studies on African Agama IX. New insights into Agama finchi ah Since the description of A. finchi, specimens from Ugan- da became available and it was possible to recognize the species from other countries by images and specimens. Therefore, the aim of this publication is to summarize and discuss the distribution of A. finchi and to assess the sta- tus of the populations possessing body-like coloured fore- limbs. MATERIAL AND METHODS Material and morphological sampling. 21 specimens of Agama finchi, including type material, were examined. Specimens from collections of the following institutions (Institutional abbreviations in parenthesis) were used: Muséum d’histoire naturelle, Geneve (MHNG); Nation- al Museums of Kenya (NMK); Zoologisches Forschungsmuseum Alexander Koenig, Bonn (ZFMK). Type material of the new species is deposited in the Zo- ologisches Forschungsmuseum A. Koenig, Bonn. For each specimen, external characters were recorded: snout-vent-length (SVL), length of tail (TL), head width between the anterior margins of the ear openings (HW), head length from the tip of the snout to the anterior mar- gin of the ear opening on the left side (HL), head height at the highest point of the head (HH), number of scales beneath the 4th toe on the left side (SD), number of scales around mid-body (MBS), number of scales between gu- lar- and inguinal fold (V; ventrals), and the number of pre- cloacal pores (PP). Measurements were taken with a di- al calliper to the nearest 0.1 mm. Measurements and scale counts were done according to Grandison (1968) and Moody & Bohme (1984). Colour patterns in living speci- mens were described using field notes and life photo- graphs. Material examined. Agama f. finchi Kenya. Malaba: ZFMK 82091-82094, NMK L/2533/3, L/2534/1,3,6. Aga- ma f. ssp. n. Ethiopia. Gambela: ZFMK 8709-8711, 66271. Uganda. Murchison Falls: ZFMK 88808-814, 88829. Agama lebretoni. Cameroon. Douala, Foyer du Marin: MHNG 2713.31; Fako (Mt. Cameroon), Limbe (Victoria): ZFMK 18891—894; Korup, Mundemba: ZFMK 61243; Magba: ZFMK 51686, 54906-907; Makum: ZMB 55709, 37061; Mamfe: ZFMK 87694—699, 87700; Mamfe, Mukwecha, Amebisu: ZFMK 87694—-699; Met- chum, Wum: ZFMK 15194-15200; Mt. Nlonako, Nguen- gue: ZFMK 69017, Rumpi Hills, Mofako Balue: MHNG 2713.29; Rumpi Hills, Big Massaka: MHNG 2713.30, 2713.32. Equatorial Guinea. Bioko Island, San Carlos: ZFMK 9353-359. Gabon. Fougamou: ZFMK 73239-245; Ngouassa: IRSNB 15686—-687. Agama paragama. Cameroon. Logone et Chari, Waza: ZFMK 15242-256. Niger. Agadez: ZFMK 36599; Tessaoua: ZFMK Bonn zoological Bulletin 60 (1): 25-34 33749-750. Agama cf. paragama. Cameroon. Benoue, Boki: ZFMK 15227-241. Agama cf. sylvanus. Cameroon. Benoué: ZFMK 33751—754. Agama spec. | CAR-1. Cen- tral African Republic. Koumbala: ZFMK 40251—260. Agama spec. | CAR-2. Mélé: ZFMK 33766; Ndeéle: ZFMK 33755-7654; Sibut: ZFMK 33765. Agama spec. | Chad Chad. Bol (near Lake Chad): 29171-29221 (will be inventoried at the collection of the MNHN). Agama sp. n. Democratic Republic of the Congo. Nagero: ZFMK 51576—-587; Isiro: ZFMK 51588. RESULTS Agama finchi leucerythrolaema ssp. n. Holotype. ZFMK 88809, adult male from Murchison Falls, Uganda; collected by W. Freund, July 2009. Paratypes. ZFMK 88810, adult male from Murchison Falls, Uganda; collected by W. Freund, July 2009. ZFMK 88811, 88829, adult females from Murchison Falls, Uganda; collected by W. Freund, July 2009. ZFMK 88808, 88812—814, juveniles from Murchison Falls, Uganda; collected by W. Freund, July 2009. Diagnosis. This is a medium-sized lizard of the genus Agama (total length of adult males up to 275 mm), which is characterized by a large gular fold, a reticulated throat and a bright nuptial coloration of adult males. The throat colouration (Fig. 1A) is a reticular pattern of red lines, which thus far is only known from A. paragama (Fig. 1D), A. sylvanus (no true specimen available), A. /ebretoni (Fig. 1C) and from recently unidentified material from the Cen- tral African Republic (formerly identified as A. sy/vanus; Fig. 1E) Lake Chad (Fig. IF) and DR Congo (Fig. 1G). Females are similar to those of the nominate form. Differentiated diagnosis. A. finchi leucerythrolaema ssp. n. not only differs from the nominotypic form by its larg- er size, but additionally males of the new taxon are dis- tinct as they possess a large gular fold, a reticulated colour pattern of the throat (for both see Fig. 1A) and body-like coloured forelimbs. Differences in pholidosis to the nom- inate form are only marginal but the new subspecies has a lower count of body scales (Table 1). Because of the reticulated throat, the new subspecies is similar to some other Agama species. From the two Cen- tral Africa species, A. paragama and A. lebretoni, the new subspecies differs as follows: Agama paragama possesses a yellow-whitish to chalk white head and has a higher number of scale rows around the midbody (Tab. 1). Additionally, adult males of A. ©OZFMK ii) Co Philipp Wagner et al. nnes ywiaz — Fig. 2. Male holotype (ZFMK 88809) of Agama finchi leucerythrolaema ssp. n. from Murchison Falls, Uganda. Bonn zoological Bulletin 60 (1): 25-34 ©ZFMK Studies on African Agama IX. New insights into Agama finchi 29 paragama show a black instead of a dark blue tail tip. Sim- ilar to the herein described new species, Grandison (1968) described the colouration of the throat of A. paragama as ‘a dark network on a cream ground which takes the form of isolated, round, cream spots’ (Fig. 1D), which is pres- ent in both sexes, and therefore similar to A. finchi leucery- throlaema ssp. n. as the females also have a striated throat pattern. A. lebretoni differs from the new subspecies by having a pale vertebral band and scattered white body scales. Ad- ditionally, the average snout-vent-length in this species is larger than in the new subspecies (A. /ebretoni: 117.8 mm; A. f. leucerythrolaema ssp. n.: 102.4 mm). The new subspecies differs from the West African, A. sy/- vanus, in having a higher mid-body scale count. Accord- ing to MacDonald (1981), A. sylvanus has 59 to 66 scale rows, whereas A. finchi leucerythrolaema ssp. n. has 71 to 78 rows. The same author described the pattern of the throat as ‘marked with an irregular pattern of longitudi- nal whitish lines and (more rarely) small spots’, which is similar to the herein described subspecies. A. finchi leucerythrolaema ssp. n. is clearly dissimilar to the Central African A. a. agama, since the latter possess- es a uniform to striated red throat, a tri-coloured tail and a yellow head. However, according to Grandison (1968), her specimens of A. agama from Nigeria (currently not traceable if it is A. agama sensu stricto or sensu lato) has 59 to 77 rows, which is similar to A. f- leucerythrolaema ssp. n. (71 to 78 rows). A. finchi leucerythrolaema ssp. n. differs from other East African Agama species as follows: — from A. lionotus in having a reticulated throat, a tri- coloured tail and a dark blue body colouration, in hav- ing lower scale counts around the midbody (A. /. liono- tus: 67-91 | 75.4; A. 1. elgonis: 79-87 | 81.8; A. finchi leucerythrolaema ssp. n.: 73.9) and a large gular fold; — from A. turuensis in having a reticulated throat, a gular fold and lower count of scale rows around the midbody (A. turuensis: 71-85 | 77.6; A. finchi leucerythrolaema ssp. n.: 73.9); — from A. kaimosae in having a reticulated throat, a gu- lar fold, a blue body colouration, lower scale counts around the midbody (A. kaimosae: 79-82 | 80.0; A. finchi leucerythrolaema ssp. n.: 73.9) and a tricoloured tail; — from A. mwanzae in having a reticulated throat, a gular fold, a blue body colouration, lower scale counts around the midbody (A. mwanzae: 67-82 | 75.2; A. finchi leucerythrolaema ssp. n.: 73.9) and a tri-coloured tail; Bonn zoological Bulletin 60 (1): 25-34 — from A. caudospinosa in having a reticulated throat, a gular fold, a blue body colouration, lower scale counts around the midbody (4. caudospinosa: 74-116 | 93.3; A. finchi leucerythrolaema ssp. n.: 73.9) and a tri- coloured tail; — from A. montana, A. mossambica and the much small- er A. armata in not possessing a heterogeneous body scalation. Description of the holotype (ZFMK 88809, Fig. 2) Habitus stout, snout-vent length 118.2 mm, tail length 150.1 mm, head length 30.3 mm, head width 19.4 mm, head height 12.8 mm. Large triangular nasal scale slightly above the canthus ros- tralis and pierced with the nostril in the posterior part, di- rected and supplied obliquely upwards. Between the nasal scales, a single narrow longitudinal smooth scale is visi- ble, followed by one smooth scale, larger than the other head scales. Ten supralabial and sublabial scales are on both sides. Head scales between the eyes are smooth, di- rected sideward from a midline of two rows of feebly keeled scale; head scales between posterior end of the eyes and neck smooth to feebly keeled, directed forwards; head scales of the temporal region smooth to feebly keeled, not directed to one side; free anterior margins of head scales with sensory pits; supraocular scales smooth. Parietal shield large and more or less pentagonal, pineal organ vis- ible, pierced more or less in the middle of the shield. Ear hole large, about the same size as the eye, margin being composed by spiny scales, surrounded by four tufts of more or less spiny, mucronate scales; tympanum superfi- cial. Nuchal crest low, consisting of 13 lanceolate scales. Gular scales flat, smooth, juxtaposed and becoming small- er towards the large gular fold. Dorsal body scales strong- ly keeled, but becoming feebly keeled at the vertebral re- gion, mucronate, equal in size, in 56 scales from midpoint of pectoral region to midpoint of the pelvic region. Ven- tral body scales smooth, slightly imbricate at their poste- rior margins, in 69 scales from midpoint of pectoral re- gion to midpoint of pelvic region. There are 72 scales rows around the midbody. Ten precloacal scales stringed in one row only. Tail scales strongly keeled and mucronate. Scales on the upper side of the forelimb strongly keeled, smooth on the underside, on the upper arm scales twice as large as the dorsal body scales, becoming smaller to- wards the underside and the manus. 4" finger longest, dig- ital length decreasing 3-2-5-1, subdigital lamellae keeled and mucronate. Scales on the upper side of the hindlimb strongly keeled, becoming smooth on the underside, on the upper tights slightly larger in size than the dorsal body scales, becoming larger towards the lower tights. 4" toe longest, digital length decreasing 3-2-5-1. ©ZFMK 30 Philipp Wagner et al. Fig. 3. Living specimens from the type localities (A) male and (B) female of Agama finchi leucerythrolaema ssp. n. from Murchi- son Falls, Uganda (photos by David Modry); (C) male and (D) female of Agama finchi finchi from Malaba, Kenya (Photos: Bri- an Finch). Colouration. (in alcohol after three months of preserva- tion). Head and neck red with a light red vertebral band extending to the back. Limbs, body and anterior third of the tail dark blue. Tail tri-coloured, at the base dark blue, followed by red and dark blue. Belly and underside of the limbs blue, tail anterior whitish, posterior bluish. Throat and the large gular fold with a reticulated pattern of red stripes on a white background. Colouration in life. Males. Head, neck and parts of the shoulders red, a broad vertebral red band extends on the back from head to about half way to hindlimbs. Body, limbs and anterior third of the tail dark blue. Tail tri-band- ed: dark blue (extending from the body), followed by red and dark blue at the tip (Figs 3a, 4a). Belly and underside of limbs blue; throat with a reticulate pattern of red stripes on a white background. Females. Head yellowish brown, with fine yellow stripes and dots; stripes and dots usually extending to the shoul- ders. Body and tail light brownish with broad lateral yel- low bands (Fig. 3b). Underside whitish, with fine dark lon- gitudinal stripes on the throat. Bonn zoological Bulletin 60 (1): 25-34 Juveniles. Similar to females but with more distinct yel- low dots and stripes on the head on the shoulders. Yellow lateral bands sometimes extending to the back. Underside whitish, with a fine dark reticulated pattern on the throat. Variation. Variations in morphology are shown in the ap- pendix and compared to other agamid species in Table 1. Colouration of males is not variable and all show the typ- ical colour pattern of throat and forelimb. Etymology. The new species is named after its remark- ably characteristic red and white vermiculated throat as compared to the nominate form. The name is derived from the Greek words ‘leukos’ for white, ‘erythros’ for red and ‘laema’ for the throat. Distribution. Beside the type localities of the two sub- species, A. finchi was also identified from Gambela in Ethiopia, represented by material from the collection of the Zoologisches Forschungsmuseum A. Koenig (ZFMK 8709-8711; ZFMK 66639, 66271—72). In contrast to the specimens of A. f. finchi from Kenya and DR Congo (see below and figs 3, 5), the adult males of this population ©ZFMK Studies on African Agama IX. New insights into Agama finchi 3] Fig. 4. (A) Living male and (B) habitat of Agama finchi leucerythrolaema ssp. n. from Murchison Falls, Uganda (Pho- tos: Wolfram Freund). show body-like coloured forelimbs and therefore repre- sent the new subspecies. The colouration of the throat is not distinguishable, because of the long preservation time. However, specimens from Gambela are similar in the av- erage of their snout-vent-length to topotypical material of the nominal subspecies (Ethiopia: 86.2 mm | Kenya: 85.7 mm). A. f/ leucerythrolaema ssp. n. is also cryptically men- tioned in Largen & Spawls (2010). Within the species chapter on Agama agama (sic., as Agama agama in nei- ther occurs in Ethiopia nor Eritrea), specimens from Gam- bela are figured (Largen & Spawls 2010, fig. 147) show- ing the herein described subspecies. According to Stephen Spawls (pers. comm. 07.X.2010), the western population of the former ‘Agama agama’ in Ethiopia and Eritrea be- longs to A. finchi, whereas the southern ones belong to Agama lionotus. Quite recently, the new subspecies was also recorded from the Lorionotom Range, Ilemi Triangle (approx coordi- nates: 4°53738.53”N, 35°31°59.44”E) by Miroslav Jirku (specimens will be inventoried in the collection of the Na- tional Museums of Kenya, Nairobi), which fills the gap between the Ethiopian and Ugandan localities. Habitat. Specimens at Murchison Falls were collected in open grassland (Fig. 4b), sitting on old houses (Fig. 4a) Bonn zoological Bulletin 60 (1): 25-34 and on palm trees. The habitat of the Ethiopian popula- tions is unknown, but is likely to be similar. Relationships. According to Wagner et al. (unpubl. da- ta) Agama finchi sensu lato is differentiated in two clades. Individuals from the type locality of the nominate form are distinct in colouration of forelimbs, throat, and in body size to specimens from Uganda and Ethiopia, but there are few other morphological differences (see Appendix). Ge- netic analysis indicates (Wagner et al. unpubl. data) that the new taxon is closely related to A. finchi and both taxa are members of the Agama agama species group. Agama f. finchi: new distribution record. Democratic Republic of the Congo. The nominate species was docu- mented by photographs from the Ituri forest (Fig. 5) where it occurs on an inselberg surrounded by rainforest is not otherwise connected to savannah areas. Ecologically, this inselberg is an arid area, as rainwater flows off immedi- ately, and the unshaded dark rocks are heated up by sun- light during the day. Therefore, fauna and flora (with e.g. rock hyrax and aloe plants; pers. comm. Reto Kuster) of this inselberg are very different from those of the surround- ing rainforest. The status of these forests as a true rain- forest is supported by the occurrence of Lepidothyris hinkeli Wagner et al., 2009, which is a character species of equatorial rainforests (Wagner et al. 2009). As can be seen from the figures 4 C—D, the specimens are identical in colouration to the specimens from the type locality of A. f. finchi in western Kenya. However, the throat coloura- tion is still unknown. The distribution of both taxa is shown in figure 6. DISCUSSION Preliminary genetic analyses (unpubl. data) separated Aga- ma finchi sensu lato into two distinct clades: this was sup- ported by differences in the colouration of the forelimbs of adult males. However, there are more morphological differences between the Ugandan and Kenyan populations than between the Ugandan and Ethiopian. The Ugandan population is distinct from the Kenyan population because of the reticulated throat and the much larger size of adult males. Also, the Ugandan population is larger than the Ethiopian population. Furthermore, only small series were collected from all populations, and body-sizes are prob- ably underestimated. Because the Ethiopian specimens were preserved a long time ago, the throat colouration is not identifiable anymore. Little information is available about the species limits within the genus Agama. In contrast to genetic distances in-between other Agama species, the distances within the A. finchi sensu lato populations are comparatively small. ©ZFMK 32 Philipp Wagner et al. Fig. 5. However, identification of the subspecies is possible and demonstrated by pholidosis, differences in colouration of adult males, and the colour pattern of the throat. There- fore, these populations should be recognized as a devel- oping species and according to article 45.6 of the Inter- national Code of Zoological Nomenclature (ICZN 1999), they can be recognized as the taxonomic rank of a sub- species. Distribution patterns and relationships are poorly known in Agama lizards. For decades, the genus was not in the focus of herpetological interests and most of the taxa were recognized as subspecies of Agama agama which, because of this original taxonomic classification, was supposed to be distributed nearly everywhere in sub-Saharan Africa. Therefore, in the older literature, A. agama is supposed to have a distribution range with includes nearly every Afrotropical country. Additionally, many specimens are misidentified or simply inaccurately labelled as A. aga- ma in museum collections. Wagner et al. (2009) described a neotype and restricted A. agama s. str. to northern Cameroon. The whole A. agama species complex is now Bonn zoological Bulletin 60 (1): 25-34 Agama f. finchi occurrence on an inselberg within the Ituri Forest, DR Congo (A) View to the inselberg within the for- est. (B) Habitat on the inselberg. (C) Living male of 4. f. finchi from the same locality. (D) Living female of 4. f finchi from the same locality (Photos: Reto Kuster). distributed along the southern border of the Sahara desert from western to eastern Africa and along the Atlantic coast to Namibia. Other than that, former East African sub- species of A. agama are now recognized as a distinct species (BoOhme et al. 2005), forming the A. /ionotus species group, which is distributed from Ethiopia through Kenya to southern Tanzania. However, the distribution pat- terns of the 4. agama and A. lionotus species groups gen- erally follow the theory of African arid corridors and 1s comparable to distributions of other reptile species groups, such as Varanus exanthematicus/ albigularis (see Wagner 2010). The only representative of the 4. agama species complex in eastern Africa (including Ethiopia) is A. finchi, as none of the other A. agama species complex taxa are present in Ethiopia and Uganda (pers. comm. Steven Spawls). To date, both subspecies of A. finchi have only been found west of the eastern branch of the East African Rift, but it is not possible to recognize this geological barrier as a dis- tribution limit since this species 1s only known in four lo- calities. ©ZFMK Studies on African Agama IX. New insights into Agama finchi 33 Fig. 6. Distribution of Agama finchi. Red dots refer to A. f finchi, whereas blue dots refer to A. f- leucerythrolaema ssp. n. (1) Kenya: road to Busia near Malaba (type locality); (2) DR Congo (photo documention); (3) Uganda: Murchison Falls (type locality); (4) Ethiopia: Gambela. Other Ethiopian localities ob- tained from Largen & Spawls 2010; (5) Kenya: Lorionotom Range, Ilemi Triangle (approx coordinates: 4°53’38.53”N, 35°31°59.44”E). Distribution was also influenced by the dispersal and re- traction of the equatorial rain forest in the past. The pop- ulation in the Ituri Forest is a relict population found on an inselberg surrounded by rainforest, usually a non-suit- able habitat for Agama species. The status of the Guineo- Congolian rainforest is supported by the occurrence of Lepidothyris hinkeli, which is a character species of this forest type (Wagner et al. 2009). Therefore, the rainforests in this area seem to be relatively young as A. finchi was enclosed on its inselberg during an extension of the rain- forest. As a result, the species must have been widely dis- tributed within the area during the time when the rainfor- est last receded. Acknowledgements. We are grateful to Reto Kuster for permission to publish his images of Agama finchi from DR Congo and his comments on the occurrence of this species in the DR Congo. Moreover, we are grateful to Stephen Spawls and Miroslav Jirku for their information about A. finchi in Ethiopia and Kenya. We thank the two referees for their comments which have improved the manuscript, and especially to Shelley Barts-Pankow who did the English review. Bonn zoological Bulletin 60 (1): 25-34 REFERENCES Bohme W, Wagner P, Malonza P, Lotters $, Kohler J (2005) A new species of the Agama agama group (Squamata: Agami- dae) from western Kenya, East Africa, with comments on Aga- ma lionotus Boulenger, 1896. Russian Journal of Herpetology 12: 83-90 Grandison AGC (1968) Nigerian lizards of the genus Agama (Sauria: Agamidae). Bulletin of the British Museum of Nat- ural History Zoology 17: 67-90 ICZN (1999) International Code of Zoological Nomenclature. International Trust for Zoological Nomenclature, Fourth Edi- tion: 306 pp. Largen M, Spawls S (2010) The amphibians and reptiles of Ethiopia and Eritrea. Edition Chimaira, 693 pp. Macdonald MA (1981) A new species of agamid lizard from Ghana. Journal of Zoology 193: 191-199 Moody SM, Bohme W (1984) Merkmalsvariationen und taxo- nomische Stellung von Agama doriae Boulenger, 1885 und Agama benueensis Monard, 1951 (Reptilia: Agamidae) aus dem Sudangiirtel Afrikas. Bonn. zool. Beitrage 35: 107-128 Spawls S, Howell K, Drewes RC, Ashe J (2002) Field Guide to the Reptiles of East Africa: All the Reptiles of Kenya, Tanza- nia, Uganda, Rwanda and Burundi. Princeton Field Guides: 543 pp. Wagner P (2007) Studies in African Agama I — On the taxonom- ic status of Agama lionotus usambarae Barbour & Loveridge, 1928 (Squamata: Agamidae). Herpetozoa 20: 69-73 Wagner P, Burmann A, Bohme W (2008a) Studies on African Agama IU. Resurrection of Agama agama kaimosae Loveridge, 1935 (Squamata: Agamidae) from synonymy and its elevation to species rank. Russian Journal of Herpetology 15: 1-7 Wagner P, Krause P, Bohme W (2008b) Studies on African Aga- ma III. Resurrection of Agama agama turuensis Loveridge, 1932 (Squamata: Agamidae) from synonymy and its elevation to species rank. Salamandra 44: 35—42 Wagner P, Wilms TM, Bauer A, Bohme W (2009) Studies on African Agama V. On the origin of Lacerta agama Linnaeus, 1758 (Squamata: Agamidae). Bonner Zoologische Beitrage 56: 215-223 Wagner P (2010) Diversity and distribution of African reptiles. Unpublished PhD thesis, University of Bonn: 374 pp. Received: 25.11.2010 Accepted: 25.02.2011 Corresponding editor: F. Herder OZFMK Philipp Wagner et al. 34 “SqUII] UdaMjoq YISUI] =Tq'T ‘seiod [eovopsaid Jo saquiNU =GOd £90} pp YO] Youdg sejjowe] =o], “IOBuLy wh Yo] you -9q se][oule] =I ‘Apogprur punose sayeog =es ‘el[esiog =YOR ‘eyeNUsA =NAA ‘ysusy peoy =TH Spm peoy =H WYs1oy peoy =H ‘yISugy [ej0} =ToY ‘ySugy prey =Tp ‘yBue]-yuaa-jnous =T AS qyeulay = | Ep = fii Ol @2 &) —s3 vl TZ ST «OST «LOL LOTT) «6VHEL) «= 9B STP YOINY :epuesq = HZ88 MINAZ = Pwavjo.ysCiaonay fy ayeuraz = Lp = 6 Mo 7 @ ST €T TW VST $6 BEET VMEPI T06 Sled Younpy :epuesEy 11888 MANAZ Yuanjosyjsduaonay fy qeu 709 ?@l td 61 SL OL 18 vl OT 98Z OOT OPI PH8LT TI TIT ses YoNpl epues O18 MNAZ MMavjosysdiaonaj fp qe = 09 I 6 8 “re 09) 08 91 TT 96C OT EI = — P8Il sey yom sepuedssy,) = 60888 MINAZ PManjosyjduaonay f jaw Ci Gl i Gl Sh to 77 I €% 67 6ST TOT ie =. 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'Faculty of Veterinary Science, Gunn Building B19, University of Sydney, NSW 2006, Australia; E-mail: jaime.gongora@sydney.edu.au 2Departamento de Ecologia, Instituto de Biociéncias, Universidade Estadual Paulista, Avenida 24-A 1515, Rio Claro, SP, 13506-900, Brazil 3Department of Biological Science, Purdue University, West Lafayette, IN 47907 USA 4Center for International Forestry Research (CIFOR), P.O. Box 0113 BOCBD, Bogor 16000, Indonesia SWildlife Conservation Society, Brazil, Rua Jardim Botanico 674, 22461-000, Rio de Janeiro, RJ, Brazil 6Environmental Studies Center, University of Redlands, 1200 East Colton Avenue, P.O. Box 3080, Redlands, California, 92373, USA 7Genetics Division, Instituto Nacional de Cancer, Rua André Cavalcanti 37, 4° andar. Bairro de Fatima, 20231-050, Rio de Janeiro, RJ, Brazil 8Departamento de Genética e Biologia Evolutiva, Instituto de Biociéncias, Universidade de Sao Paulo, Rua do Matao 277, Sao Paulo, SP 05508090, Brazil °Wildlife Conservation Society, 2300 Southern Boulevard Bronx, New York 10460 10Pepartament de Sanitat 1 d’ Anatomia Animals, Facultat de Veterinaria, Universitat Autonoma de Barcelona, Edifici V, E-08193 Bellaterra, Barcelona, Spain ‘Genetica de la Conservacion, Instituto de Investigaciones Biologicas Clemente Estable, Facultad de Ciencias, Universidad de la Republica Oriental del Uruguay, Av. Italia 3318-11600. Montevideo, Uruguay Abstract. Three extant species of peccaries (Tayassuidae) are currently recognized and are distributed in the Americas: Pecari tajacu, Tayassu pecari and Catagonus wagneri. The recent claim for a new peccary species, Pecari maximus, rais- es questions about whether there is sufficient molecular or other evidence to support it as a new species. Here, we revis- it that evidence using 91 published and 50 novel DNA sequences mainly from P. tajacu, as well as comment on the mor- phological and ecological evidence used to support such a claim. Phylogenetic analyses show that the single specimen of P. maximus used clustered within the South American lineage of P. tajacu closely related to Brazilian specimens. In addition, the morphological and ecological data used to claim the species status of P maximus appears to be deficient and inconclusive, in the light of available literature. Key words. Collared peccary, control region, mitochondrial DNA, Pecari maximus, Pecari tajacu, Tayassuidae. INTRODUCTION Three extant species of peccaries (Tayassuidae) are cur- rently recognized: Pecari tajacu (Collared peccary), Tayassu pecari (White-lipped peccary) and Catagonus wagneri (Chacoan peccary) (Wetzel et al. 1975; Wood- burne 1968; Wright 1989, 1993). These species have been identified according to morphological and cytogenetic pa- rameters and more recently using DNA sequences. Clas- sification has generally relied on biological concepts of reproductive and genetic isolation. One confounding fac- tor is that P. tajacu and T. pecari are known to hybridize in captivity (Sowls 1997) and in the wild with the latter being reproductively sterile (Andrea et al. 2001). Cyto- genetic studies have confirmed the existence of three species, showing significant chromosomal differences in number and structure: P tajacu (2n=30), 7) pecari (2n=26) and C. wagneri (2n=20) (Benirschke & Kumamo- 96 Jaime Gongora et al. to 1989). Although chromosome painting across species (Bosma et al. 2004) has provided some chromosomal evo- lutionary insights into this family, a full understanding of the species relationships from a cytogenetic perspective remains hampered by a lack of information on C. wag- neri. However, phylogenetic analyses of mitochondrial and nuclear DNA sequences have revealed that T. pecari and C. wagneri grouped in separate clades from P. tajacu (Gongora & Moran 2005; Theimer & Keim 1998). These conclusions obtained from molecular markers contrast with cladistic analyses of morphological data which pro- posed two different scenarios of relationships: First, P. ta- jacu and C. wagneri are more closely related whereas 7: pecari 1s considered to be a member of a separate clade (Wright 1989, 1993, 1998); and second, P. tajacu and T. pecari are more closely related to each other than to the C. wagneri (Wetzel et al. 1975; Wetzel 1977). The new species Pecari maximus. The recent claim of a new peccary species by van Roosmalen et al. (2007) rais- es questions about whether there is sufficient molecular and/or other evidence to recognise P. maximus as a new species. The morphological and genetic evidence used by van Roosmalen et al. (2007) to support the species status for P. maximus appears to be controversial. They analysed a small number of specimens from deceased animals (five skins and three skulls, two of which were from adults), and the body weight (40—50 kg) and size of the animals (total body length 127 cm; ear length 13 cm; shoulder height 85 cm) were estimated from reports and photo- graphs by local hunters. Morphometric analyses were al- so estimated from skin measurements, and they relied on pelage colour to point the differences with other species. In this study, the mitochondrial control region and two nu- clear SINE PRE-1 sequences of a single individual were used to assess the phylogenetic position of P. maximus among recognised species of peccaries. In addition, van Roosmalen et al. (2007) used morphological and behav- ioural data to support a separate status for those pecca- ries. These authors argue that in contrast with the other peccary species, which they suggest roam semi-nomadi- cally ina highly variable landscape in a noisy herd, P.: max- imus appears to walk silently through its habitat in small family groups that contain a single adult pair with or with- out 1-2 offspring. They also claim that P. maximus per- forms little or no uprooting and has been seen feeding mainly on freshly fallen fruits and seeds exposed on the forest floor, which appears to be result of direct observa- tions. The claim by van Roosmalen et al. (2007) under- lines the urgent need to complete an ongoing modern tax- onomic assessment of the family Tayassuidae. A revision of the peccaries is imperative to orient conservation man- agement and planning as well as to increase understand- ing of the basic biology, ecology and evolution of this group (Taber 1993). Bonn zoological Bulletin 60 (1): 95-101 Differentiation within P tajacu. Highly relevant to the claim by van Roosmalen et al. for a fourth species in the genus Pecari 1s that previous phylogenetic mitochondri- al DNA studies have shown genetic variation within Col- lared peccary to be higher than that observed between C. wageneri and T. pecari, and was also higher than that ob- served between recognised species of the family Suidae (Gongora & Moran 2005; Gongora et al. 2006). Pheno- typic, morphological, chromosomal and DNA data have been used to assess the levels of differentiation within P tajacu. This lineage has a broad distribution in the sub- tropical and tropical ecosystems of the Americas compris- ing a wide variety of environments (rainforest, semi-arid thorn forest, coastal forest, cloud forest, deserts, islands, rangelands, scrublands, savannas, and freshwater wetland) between the south-western United States of America and northern Argentina (Grubb & Groves 1993). Its ancestry also seems to have an early divergence from the other pec- cary lineages in the Americas (Gongora et al. 2006). Ear- ly morphological studies provided the first evidence of cra- nial and dental variation between P. tajacu from through- out the Americas (Kiltie 1985; Woodburne 1968), although specimens were ultimately grouped into a single species. Variations in size and pelage colour, coupled with distri- bution data, have been the basis for proposing the exis- tence of 14 subspecies of P. tajacu (Grubb & Groves 1993; Hall 1981). However, the inheritance of these traits has not been tested or substantiated by other methods. Conventional cytogenetic (Builes et al. 2004; Gongora et al. 2000; Vasart et al. 1994) and cross-species chromosome painting (Adega et al. 2006; Bosma et al. 2004) studies showed variation in the structure of two chromosomes be- tween P. tajacu specimens from Arizona, Colombia and Brazil, providing additional evidence for differentiation within this species. A single captive specimen from Colombia, heterozygous for a balanced translocation may represent a hybrid between the two different P. tajacu lin- eages (Builes et al. 2004). This may contrast with other cytogenetic studies (Adega et al. 2006) which propose a ‘significant barrier’ for hybridization between major Col- lared peccary lineages/species. However, germ line cyto- genetic and/or reproductive studies are required to assess whether this possible hybrid is fertile before any conclu- sion can be addressed. In this paper we revisit the genetic data from P. maximus (van Roosmalen et al. 2007) using a DNA dataset of the three peccary species generated by Gongora et al. (2005, 2006) and including 50 new DNA P. tajacu sequences from United States, Mexico, Colombia and Brazil. We al- so analyse the morphological and ecological evidence used to support the species status of P. maximus in the light of available literature. ©ZFMK Revisiting the status of Pecari maximus 97 MATERIAL AND METHODS Sampling and DNA extraction. Blood, tissue and hair samples from 50 captive and wild P. tajacu from south- eastern and central-western Brazil (BRAI—BRAS; n=5); southern and northern Colombia (COL11—COL40; n=30);: central and southern-central Mexico (MEX41—MEX48; n=8); and Texas in the United States (USA6—USA 12; n=7) were used in this study (Appendix I). DNA was extract- ed using either a standard proteinase K and phenol-chlo- roform protocol (Sambrook et al. 1989) or QIAamp DNA mini kits (Qiagen Pty Ltd). PCR and Sequencing. We used previously published primers to amplify approximately 1,200 bp of the com- plete mitochondrial control region (Kim et al. 2002) at the University of Sydney, Australia. PCR was carried out us- ing methods outlined in Gongora et al. (2005; 2006) ex- cept for sequencing of forward and reverse strands which were directly sequenced rather than clone inserts. Control region sequences (450 bp) from the Texan specimens (n=7) were independently generated at Purdue Universi- ty. Primers, PCR and sequencing conditions were per- formed as described in Cooper et al. (2010). Although there is a difference in length between the above subsets of sequences, the corresponding regions are informative for assessing the major clades of P. tajacu described in pre- vious studies (Gongora et al. 2006). Data analysis. Forward and reverse sequences were over- lapped in order to obtain a single double-stranded se- quence for each animal using CodonCode Aligner v3.5.5 (CodonCode Corporation). After excluding flanking primer regions, the novel sequences (~1,120 bp, GenBank accession numbers HM102371 through HM102419) were aligned using the program Muscle (version 3.6; Edgar, 2004) along with those P tajacu sequences (AY546539—AY 546569) published by Gongora et al. (2005; 2006) and the single sequence (DQ009006) from P. maximus published by van Roosmalen et al. (2007). The control region from 7. pecari (AY546516—AY 546519) and C. wagneri (AY 546520, AY546521) were included as an outgroup. The alignment used for analyses described con- sisted of 1,140 bp after excluding the tandem repeat mo- tifs, except for a single motif, due to heteroplasmy accord- ing to Gongora et al. (2006). Because few gaps were ob- served they were kept in the alignment. A separate dataset of concatenated nuclear PRE-1 642 and PRE-1 27 sequences was generated and aligned as de- scribed above. It consisted of available sequences from GenBank (P. maximus, DQ016372 and DQ016371; P. ta- Jacu, DQ190931—DQ190883, AY569340—AY 569339, and AY568052—AY568047; 7. pecari, AY546331 and AY546528; and C. wagneri, AY546530 and AY546527). Bonn zoological Bulletin 60 (1): 95—101 The best fit DNA substitution model for the control region and PRE-1 alignments were selected by the program Mod- elgenerator (version 0.85; Keane et al. 2006) and imple- mented in Maximum Likelihood (ML) to reconstruct a phylogenetic tree using the software PhyML (version 3.0; Guindon & Gascuel 2003). The tree topology space was searched by using the best of Nearest Neighbour Inter- change and Subtree Pruning and Regrafting starting from five random starting trees generated by BioNJ ( Guindon & Gascuel 2003; Guindon et al. 2010). Branch support was calculated using the approximate likelihood ratio test (aLRT) with SH-like interpretation, as it is as conserva- tive and accurate test compared with bootstrapping but less computationally intensive (Anisimova & Gascuel 2006; Guindon et al. 2010). RESULTS AND DISCUSSION Clustering of P. maximus within recognised major clades of P. tajacu. Novel and published Collared peccary mito- chondrial control region sequences representing 44 hap- lotypes clustered in two major clades: North/Central American clade and South American clade (Fig. 1) as de- scribed by Gongora et al. (2006). The novel P. tajacu se- quences from Brazil clustered within the South American clade closely related to two sequences from Argentina, while those from Texas clustered within the North/Cen- tral American clade closely related to specimens from Ari- zona, Mexico and Colombia. Sequences from Colombian specimens split between the two major clades consistent with previous studies of being a paraphyletic group (Gongora et al. 2006). The current results corroborate that the Gongora et al. (2006) dataset is suitable for assessing major subcontinental geographical origins of captive and wild specimens. Unexpectedly for a sequence from what is considered as a different species, Pecari maximus clustered within the major South American clade closely related to central- western Brazilian and northern Argentinean sequences supported by high aLRT values (Fig. 1) rather than in a separate new clade. Consistent with control region analy- ses, ML tree of PRE-1 sequences shows that those from P. maximus cluster within the P. tajacu closely related to sequences from Bolivia, Colombia and the United States (data not shown). The clustering of Pecari maximus with- in a previously recognized South American clade of P. ta- Jacu (Gongora et al. 2006) do not support the new species status of those specimens studied by van Roosmalen et al. (2007). Although there is no agreement as to which par- ticular sequences accurately and universally reflect the species’ boundaries (Vogler & Monaghan 2007), other mo- lecular markers, such as cytochrome b or cytochrome c oxidase / are usually recommended as being more inform- ©OZFMK 98 COL10 0.92 0.86 Fig. 1. Jaime Gongora et al. North/Central America P. tajacu South America Outgrup Maximum Likelihood tree of the mitochondrial control region sequence representing forty-four haplotypes observed in eighty P. tajacu specimens, using both C. wagneri (CHP) and 7! pecari (WLP) sequences as outgroups. Brackets indicate the two major clades found in P. tajacu with Colombian (COL) specimens showing paraphyletic relationships. Pecari maximus clustered within P. tajacu closely related to those specimens from Brazil (BRA) and Argentina (ARG). Details of samples used in this study are described in Appendix 1. Numbers close to branches are aLRT values. For clarity, only aLTR values for the main internal groups higher than 0.80 are shown. ative to assess genetic species distinction (Blaxter 2004; Blaxter et al. 2005; Bradley & Baker 2001). Phylogenet- ic conclusions based on a single specimen are insufficient and uncertain. Under the current DNA evidence it is like- ly that single specimens studied by van Roosmalen et al. (2007) correspond to one of the P. tajacu lineages iden- tified by Gongora et al. (2006). Species status has come and gone for some putative taxa within the distantly re- lated suid family. For instance, the possible extinct S. buc- Bonn zoological Bulletin 60 (1): 95-101 culentus (Indochinese or Vietnam Warty pig) was initial- ly considered as a separate species within the genus Sus (Groves 1997), but after DNA analyses showed that its se- quences clustered closely with S. scrofa (domestic pig and wild boar), it has been suggested that it could correspond to a geographically restricted variant of S. scrofa rather than a distinct species (Mona et al. 2007; Robins et al. 2006). ©OZFMK Revisiting the status of Pecari maximus 99 Comments on the morphological, behavioural and eco- logical evidence of P. maximus. The body weights report- ed for PR. maximus (van Roosmalen et al. 2007) are with- in the range reported for P. tajacu, albeit at the large end. For instance, Sowls (1997) reports exceptional weights of wild Collared peccaries in Arizona in excess of 42 kg, and Bodmer (1989) reports weights close to 40 kg in Peru. Bodmer et al. (pers. obs.) have observed considerable vari- ation in the skull sizes of both P. tajacu and T. pecari from the Tahuayo, Yavari and Tamshiyacu regions of north-east- ern Peru, well within the range reported by van Roosmalen et al. (2007). These preliminary observations are based on the collection held at the zoology museum of the Univer- sidad Nacional de la Amazonia Peruana (UNAP), which consists of several thousand skulls collected by local sub- sistence hunters over a 10 year period. In addition, Bod- mer et al. (per. obs.) have recorded body weights from both P. tajacu and T. pecari hunted by local people in the Yavari river valley. Again, there is considerable variation, and the body weights are well within the range reported by van Roosmalen (Bodmer pers. obs.). Comparing 7. pecari with the proposed new species, the former may sim- ilarly reach head-body lengths over 120 cm and weights as high as 40 kg in the Brazilian Pantanal (Keuroghlian et al. 2006) and 50 kg in the northern Amazon (Fragoso 1998). It would be desirable that skull measurements of the proposed P. maximus be statistically compared with those from extremely large P. tajacu found elsewhere in their range. Further, using skin pelage characteristics to distinguish species can be problematic since bristle colour may differ substantially within peccary species even in the same area (Gongora et al. 2006). In Collared peccaries, their lighter-haired collar pattern may vary from distinct in some individuals to barely noticeable in others. The most compelling morphological lines of evidence present- ed by van Roosmalen et al. (2007) are three photographs from three individuals of exceptionally long-legged pec- caries which appear different from peccaries we are fa- muiliar with, but these are not backed up by measurements. Ecological and behavioural differences have also been used to characterize peccary species (Sowls 1997). van Roosmalen et al. (2007) have attributed a number of dif- ferentiating traits for their peccary in line with this rea- soning. However these claims are not based on intensive field observations, nor do they draw on recent literature concerning the ecology and behaviour of both P. tajacu and 7. pecari to inform contrasts. For instance, while they claim that the new peccary uniquely lives in adult pair groups, sometimes with young, P. tajacu is also known to forage alone or in small groups, even while belonging to herds typically of six to twelve individuals (Keurogh- lian et al. 2004; Sowls 1997; Taber et al. 1994), although larger groups have been reported from the Amazon (Fragoso 1994). Furthermore, the number of individuals Bonn zoological Bulletin 60 (1): 95-101 seen together may depend on the time of day. Radio telemetry studies have shown that 7. pecari has distinct seasonal movements that are related to habitat and fruit availability within their large home ranges, and P. tajacu has relatively small stable home ranges (Altrichter et al. 2001; Fragoso 1998; Keuroghlian et al. 2004; Keurogh- lian & Eaton 2008; Sowls 1997). Neither species can be described as roaming semi-nomadically as van Roosmalen et al. (2007) has suggested. Instead they commonly for- age on freshly fallen fruits (Altrichter et al. 2000, 2001; Beck 2005; Keuroghlian & Eaton 2008). Therefore, these are not unique behaviours of P- maximus. Whether pec- caries need to dig with their noses during foraging depends on the type of habitat where fruits are available, i.e. buried within the sediments of a swamp or exposed on the for- est floor (Keuroghhan & Eaton 2008). With only three skulls attributed to the new species, sample size is too low to comment on tooth wear patterns. Also, the lack of scent from the new peccary skins may be related to the skill of the hunters (that remove scent glands during butchering; Keuroghlhian pers. obs.). We note that subjectively there seems to be a considerable range between species and in- dividuals with Chacoan peccaries having the least intense scent which may contribute to them being harder for the dogs of local hunters to track (Taber et al. 1993). Final- ly, observations are too limited, and overly dependent on local hunter hearsay, to draw conclusions that this possi- ble new species has abandoned social groupings, group defence and territorial scent marking. The possible discovery of a new peccary species from the Amazon basin is very exciting, and plausible, consider- ing recent discoveries of new mammal species in this re- gion mainly by van Roosmalen et al. (1998, 2000, 2002, 2003). However, based on the scant information report- ed by van Roosmalen et al. (2007), and the results of our own genetic analysis, we conclude that there is not enough evidence to support the claim of a new species, and that the specimens studied by van Roosmalen are most likely P. tajacu. We also consider that further DNA, cytogenet- ic and morphological research is necessary to better un- derstand peccary taxonomy. Additional studies using oth- er genetic molecular markers, cytogenetic, and morpho- logical analyses incorporating new samples from all across the Amazon basin, will provide the basic information for understanding the genetic and morphology variation un- der the extant recognized species. Acknowledgements. This study was funded by the University of Sydney International Program Development Fund to J. Gongora to support collaboration between the University of Syd- ney (Australia) and the Universidade de Sao Paulo (Brazil). Brazilian peccary samples were sourced under permit IBAMA 13601 from Instituto Brasileiro do Meio Ambiente e dos Recur- sos Naturais Renovaveis and imported into Australia under per- mits AQIS IP07011173 and CITES, 09BR002827/DF. Other pec- ©ZFMK 100 Jaime Gongora et al. cary samples were sourced from wild and captive specimens col- lected for other studies from J. Gongora and J.D. Cooper. We thank Peter Waser and J. Andrew DeWoody for their support to J.D. Cooper to access the Collared peccary sequences from Texas. We also would like to thank all the institutions that col- laborated in obtaining samples listed in Appendix I. REFERENCES Adega F, Chaves R, Kofler A, Krausman PR, Masabanda J, Wienberg J, Guedes-Pinto H (2006) High-resolution compar- ative chromosome painting in the Arizona Collared peccary (Pecari tajacu, Tayassuidae): a comparison with the karyotype of pig and sheep. Chromosome Research 14: 243-251 Altrichter M, Saenz J, Carrillo E, Fuller T (2000) Dieta estacional de Tayassu pecari (Artiodactyla: Tayassuidae) en el Parque Nacional Corcovado, Costa Rica. Revista de Biologia Trop- ical 48: 689-702 Altrichter M, Carrillo E, Saenz J, Fuller T (2001) White-lipped peccary (Zayvassu pecari, Artiodactyla: Tayassuidae) diet and fruit availability in a Costa Rican rain forest. 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Pp. 389-400 in Evolution of Ter- tiary Mammals of North America (C. M. Janis, K. M. Scott, and L. L. Jacobs, eds). Cambridge University Press, Cam- bridge, United Kingdom Appendix I. Origin of Collared peccary samples Argentina: North, Ciudad de Rio Cuarto ARG1-2; Ola- varria, ARG3-4; Bolivia: North, La Paz, Mallasa Zoo, BOL 1-2. Brazil: State of Amazonas, PMAX; South East, State of Sao Paulo, BRA1—4; Central West, State of Ma- to Grosso do Sul, BRAS. Colombia: Central, Santa Cruz Zoo, COL1; Jaime Duque Zoo, COL2; North Central, Ma- tecana Zoo, COL4, COL20—22; Santafe Zoo, COL5; North West, Barranquilla Zoo, COL6, COL30—-40; South Central, La Lagartija Zoo, COL7; La Lagartija Zoo, COL8; Cali Zoo, COL9; South East, Macagual Farm, COL10, COL23-29; Central, Santa Cruz Zoo, COL11-19; Mexico: Central, Guadalajara Zoo, MEX1-—2; North, Puebla Zoo, MEX3-4, MEX41-42; South Central, Me- xico DF Zoo, MEX5—8, MEX43-48. USA: Arizona, USAI1-3; Texas, USA4—-12. Unknown: Adelaide Zoo, UNK1-2. Received: 23.10.2010 Accepted: 25.01.2011 Corresponding editor: R. Hutterer ©OZFMK Ge Bonn zoological Bulletin Volume 60 Issue | pp. 103-107 Bonn, May 2011 Two new species and a new record of the genus Neolindus Scheerpeltz, 1933 (Coleoptera: Staphylinidae: Paederinae) Ulrich Irmler Institute of Ecosystem Research, Dept. Applied Ecology, University, Olshausenstrasse 40, D-24098 Kiel, Ger- many; E-mail: uirmler@ecology.uni-kiel.de; phone: +49 431 880 4311. Abstract. The two new species Neolindus verhaaghi from Peru and Neolindus pastazae from Ecuador are described. A new record from central Amazonia of Neolindus densus Herman, 1991, is presented. Key words: Paederinae, new species, Ecuador, Peru, new record, central Amazon. Resumen: Las dos especies nuevas Neolindus verhaaghi de Peru y Neolindus pastazae de Ecuador estan descrito. Uno lugar nuevo de Amazonica Central de Neolindus densus Herman, 1991, esta presentado. Palabras clave: Paederinae, especie nueva, Ecuador, Peru, lugar nuevo, Amazonica Central. INTRODUCTION The genus Neolindus is restricted to Central and South America and distributed from Bolivia and southern Brazil in the south to Costa Rica in the north. Ecuador is the most species-rich country with 10 species. The first species was described by Sharp (1876) as Lindus religans Sharp, 1876. Later, Scheerpeltz (1933) renamed the genus Neolindus, since the name Lindus was already preoccupied. Irmler (1981) added five new species from Brazil and Peru, and Herman (1991) published a review of the genus, added 27 species, and published a cladistic analysis of the genus to- gether with Cylindroxystus Bierig, 1943. Thus, the genus contained 33 species in the Neotropical region. In 2009, I collected a new species of the genus in the Rio Pastaza basin, Ecuador. Earlier, I had found a further new species in the collection of my colleague Manfred Ver- haagh, Natural History Museum in Karlsruhe, Germany. Inferred from the low number of specimens found, all species seem to be very rarely collected. Most species have been described by only one specimen. The two new species are also known by only one specimen. Overall, 72 specimens have been collected in the Neotropics. There- fore, a new record of NV. densus Herman, 1991 found near Manaus in the central Amazon basin is also published, here. MATERIAL AND METHODS The new material is deposited in my private collection (UIC). For the photographs, a Makroskop M 420 (Wild, Her- brugg) was used in combination with a digital camera (Nikon D100). Head length was measured from anterior edge of clypeus to posterior edge of head disc, pronotum along the midline from anterior edge to posterior edge, ely- tra from anterior edge of shoulders to posterior edge; width was measured at the widest part of tagmata. In the meas- urements of total length, the abdominal inter-segmental space is subtracted. RESULTS Neolindus verhaaghi, new species (Figs 1 A—H) Holotype. Male; Peru; Huanuco, Panguana, 150 km SW Pucallpa, tropical rain forest, pitfall trap, 2—16.7.1984, leg. M. Verhaagh, #pWA65 (UIC). Diagnosis. The species is attributed to the group of N. agi- lis Herman, 1991 and N. densus Herman, 1991. The cen- tral emargination of the sth sternite [Fig. | F] is similar 104 Ulrich Irmler Fig. 1. Neolindus verhaaghi; antennae (A), aedeagus in lateral (B) and ventral (C) aspect, 7“ and 8¢" visible tergites (D), 6! vis- ible tergite (E) and sternite (F), dorsal aspect of head and pronotum (G) and elytra (H) showing punctation and surface (scale bar 0.5 mm). Bonn zoological Bulletin 60 (1): 103-107 ©ZFMK New Neolindus from South America 105 to N. densus Herman, 1991 [Fig. 131 in Herman 1991] and N. cephalochymus Herman, 1991 [Fig. 139 in Herman 1991]. Antennomere 2 [Fig. | A] is slightly shorter than 3, whereas it is slightly longer in N. densus and N. cephalochymus. Furthermore, the penultimate anten- nomeres in N. densus are wider than they are long, but more or less quadrate in N. verhaaghi. As in N. agilis [Fig. 147 in Herman 1991], N. verhaaghi has no dense patch of setae on the 8" sternite [Fig. | F]. The apical emargina- tion on the 8" sternite is deeper in N. verhaaghi than in N. agilis, and the aedeagus has an apical cavity [Fig. | B, C] which is absent in all related species. Description. Length: 4 mm. Colour: red, legs, antennae and posterior edge of tergites yellow. Head: 0.50 mm long, 0.55 mm wide; eyes as long as temples, temples rounded in smooth curve without forming angles; extremely fine micro-punctation and without microsculpture; surface shiny; one trichobothrium and one setiferous puncture at front edge of eyes; two setiferous punctures on disc bet- ween eyes [Fig. | G]; distance between these punctures slightly wider than between upper edge of eye and adja- cent puncture; transverse row of punctures at posterior edge of vertex, and two diagonal rows of punctures be- tween eyes and posterior edge of vertex. Antennae as long as head and pronotum; 3" antennomere slightly longer than 274 antennomere [Fig. | A]; 4** antennomere slight- ly longer than wide; following antennomeres more or less quadrate, pubescent and with few apical setae. Pronotum: 0.65 mm long, 0.60 mm wide; with longitudinal row of 11-12 punctures on each side of smooth midline [Fig. 1 G]; several punctures on laterad to paramedial row of punctures; distance between these punctures at least as wide as diameter of punctures; surface without mi- crosculpture, polished and shiny. Elytra: 0.75 mm long, 0.65 mm wide; with sutural row of 11—12 punctures and three more rows on disc [Fig. | H]; two rows of irregu- lar punctures laterad to disc. Abdomen densely punctate; anterior segments distinctly more densely punctate than posterior segments; 8'" tergite apically rounded [Fig. 1 E]; 8th sternite [Fig. 1 F] with triangular emargination, cen- tral impunctate stripe, and without clusters of setiferous punctures. Aedeagus with apical cavity and lateral trun- cate prominences behind cavity [Fig. 1 B, C]. Etymology. The specific name refers to the collector of the species, Manfred Verhaagh, from the Natural History Museum in Karlsruhe, Germany. Neolindus pastazae, new species (Figs 2 A—I) Holotype. Male; Ecuador, Tungurahua, 10 km W of Banos, valley of Rio Pastaza, path near waterfall “Man- Bonn zoological Bulletin 60 (1): 103-107 to de la Novia” (78°20.16W, 1°24.12S), sifting litter, 28.7.2009, leg. U. Irmler (UIC). Diagnosis. Neolindus pastazae is certainly closely relat- ed to N. punctogularis Herman, 1991, as determined by the transverse cluster of numerous setae at the apical edge of the gula. It can be easily differentiated from N. punc- togularis by the darker colour of head and pronotum and the different punctation of pronotum [Fig. 2 H]. The last abdominal tergites and sternites [Fig. 2 E, F, G] are very similar to N. punctogularis Herman, 1991 [Figs 205—207 in Herman 1991]. Besides the development of the trans- verse row of numerous setae at the anterior edge of the gula, both species can be differentiated from N. schubar- ti Irmler, 1981, N. religans Sharp, 1876, and N. bidens Herman, 1991, since the latter are carinate on 8th stern- ites or have a ridge at the posterior edge [Figs 211, 220, 223 in Herman 1991]. Description. Length: 10.5 mm. Colour: brown, antennae, palpae and legs yellow. Head: 0.90 mm long, 1.5 mm wide; with eyes slightly prominent [Fig. 2 H]; temples 1/4 as long as eyes; temples obtusely angled to posterior edge, outer part of posterior edge transversely narrowed to neck; labrum with pair of apically rounded denticles near the middle, with five setae at anterior edge; disc with fine mi- cro-punctation; surface polished; two large punctures on each side of vertex between eyes; distance between these punctures wider than between each puncture and adjacent eye; few setiferous supraocular punctures; central punc- ture with trichobothrium; row of setiferous punctures along posterior edge; supraocular punctures and posteri- or row of punctures smaller than two large punctures be- tween eyes, but much larger than micro-punctures; gula with transverse row of numerous setae near anterior mar- gin. Antennae with first antennomere elongate and as long as antennomeres 2 and 3 combined [Fig. 2 A]; 2™4 anten- nomere short, only 1/3 as long as 3" antennomere; fol- lowing antennomeres elongate and decreasing in length; antennomeres 3 to 11 pubescent. Maxillary palp at 2™4 seg- ment with several long setae at inner side. Pronotum: 1.35 mm long, 1.65 mm wide; with sides more or less paral- lel in anterior half [Fig. 2 H]; in posterior half, smoothly rounded to posterior edge without forming angle; margin continuing from posterior edge to anterior edge and cov- ered by anterior angles in dorsal aspect; disc polished and shiny; irregular row of 11 to 12 punctures on each side of smooth midline; two transverse rows laterad to parame- dial rows of punctures and few scattered punctures later- ad. Elytra: 1.95 mm long, 1.85 mm wide; surface polished and shiny; with irregular rows of dense punctures [Fig. 2 I]; average distance between punctures distinctly narrow- er than half diameter of punctures; partly coriaceous. Ab- domen densely and deeply punctate; anterior tergites more densely punctate than posterior tergites; 8" tergite trilobed ©OZFMK 106 Ulrich Irmler D ea ‘i 7 AI ry J Fig. 2. | Neolindus pastazae; antennae (A), aedeagus in lateral (B) and ventral (C) aspect, 5‘ visible sternite in ventral aspect (D), 78 and 8th visible tergites (E), 6'" visible tergite (F) and sternite (G), dorsal aspect of head and pronotum (H) and elytra (I) showing punctation and surface (scale bar 0.5 mm). Bonn zoological Bulletin 60 (1): 103-107 OZFMK New Neolindus from South America 107 with triangular central prominence and with striate struc- tures at posterior edge [Fig. 2 F]; 7" sternite with semi- circular emargination at posterior edge [Fig. 2 D]; 8" ster- nite with deep central emargination and glabrous central stripe [Fig. 2 E]. Aedeagus slightly asymmetric, with two long bifurcate prominences apically and one pair of hooks in apical half [Fig. 2 B, C]; ventral surface with deep cav- ity in paramedial position. Etymology. The specific name derives from the location in the valley of the Rio Pastaza where it was found. Neolindus densus Herman, 1991 New record. Brazil, Amazonas, 15 km SW Manaus, on Ilha de Marchanteria (59°55.21 W, 3°13.59S), inundation forest in Varzea, tree eclector #50E, 1 male, 1 female, 18.2.1982, leg. J. Adis (UIC). DISCUSSION Together with the two new species the total number of Ne- olindus species increased to 35 species with Ecuador (11) and Peru (7) as countries with high numbers of species. Only Brazil, which has a much larger area than these two countries, has a similarly high number with nine species. The species have been very rarely collected. As N. den- sus Herman, 1991 shows, the distribution can neverthe- less cover a wide area. This species seems to occur along the Amazon valley from its mouth near Belém to the An- dean foot hills near Leticia (Columbia). A more detailed analysis of the ecology is difficult. According to the in- formation given by Herman (1991) many species have been found in leaf litter of rain forests. In some labelled information the habitat was described as “under felled tree”. The habitat of N. pastazae was also in litter layer under a felled tree. The collection of N. densus Herman, 1991 by a tree eclector in the central Amazon basin sug- gests that the tree habitat might be a more important habi- tat than can be derived from the labelled information. Thus, soil litter layer might be only an accidental habitat of the normally inhabited tree trunks, which are rarely in- vestigated. Bonn zoological Bulletin 60 (1): 103-107 According to Herman (1991), the Neolindus species can be separated more or less in three species groups which are closer related. N. verhaaghi can be attributed to clade (17) with N. punctventris, N. densus, N. agilis, N. cephalochymus, N. bullus, N. hamatus, N. procarinatus, and N. retusus. This group is characterised by the pres- ence of parallel carinae at the base of sternum VII. The species of this group are distributed over the whole equa- torial region, from the lowland rain forest of the Amazon basin to the western rainforest in Ecuador. Neolindus pas- tazae seems to be related to clade (24) with N. punctogu- laris, N, hangarthi, N. schubarti, N. bidens, and N. reli- gans. In this group, species are characterised by a prono- tum wider than it is long and tergum VIII being trilobed. Whereas N. religans and N. schubarti are distributed in eastern Brazil, N. bidens, H. hanagarthi, and the new N. pastazae represent a western branch of this group. Acknowledgements. Dr. Manfred Verhaagh (Natural History Museum in Karlsruhe, Germany) and Prof. Joachim Adis (for- merly Max-Planck Institute, Dep. Tropical Ecology, Plén, Ger- many) gratefully gave me their Neolindus material for the study and for the deposition in my collection. REFERENCES Herman L (1991) Revision of the subtribe Cylindoxystina (Coleoptera: Staphylinidae: Paederinae). Bulletin of the American Museum of Natural History 203: 1-83 Irmler U (1981) Neue Arten der neotropischen Gattung Neolin- dus Scheerpeltz (Coleoptera, Staphylinidae). Studies on Neotropical Fauna and Environment 16: 209-215 Scheerpeltz O (1933) Coleopterorum Catalogus. Staphylinidae VI Supplementum I. W. Junk, Berlin, 989-1500 Sharp D (1876) Contributions to an insect fauna of the Amazon valley. Coleoptera — Staphylindae. Transactions of the Ento- mological Society of London 1876: 27-424 Received: 03.05.2010 Accepted: 20.10.2010 Corresponding editor: D. Ahrens ©ZFMK Bonn zoological Bulletin Volume 60 | Issue | | pp. 109-111 | Bonn, May 2011 A remarkable record of Phaneroptera falcata (Poda, 1761) (Saltatoria: Phaneropteridae) from north-eastern Poland Wolfgang Bohme*, Peter Geissler & Philipp Wagner Zoologisches Forschungsmuseum Alexander Koenig, Adenauerallee 160, D-53113 Bonn, Germany “ Corresponding author: E-mail: w.boehme.zfmk@uni-bonn.de. Abstract. A record of Phaneroptera falcata is made from the northeastern Poland, extending the known distribution in this country for about 300 km to the north and linking it with a newly discovered occurrence in southern Lithuania. The northern margin of the species’ range in eastern Central Europe and in East Europe is briefly reviewed. Phaneroptera falcata (Poda, 1761) belongs to those fau- nal elements which underwent a rapid northward-direct- ed range extension on the northwestern edge of their dis- tribution areas, particularly in NW Germany (Northrhine- Westphalia) and the Benelux countries (Detzel 1998). A similar trend was observed in Thuringia (Kohler, 2010), and even the federal states of Lower Saxony and Bran- denburg have already been reached (Grein 2000, 2007, Landeck et al. 2005). In the eastern part of its Central Eu- ropean range, however, P. falcata was believed, despite some expansional trend in the Czech Republic (Koéarek et al. 2008), to be restricted to the southern half of Poland, as indicated by e.g. the sketch maps in Detzel (1998) and Maas et al. (2002). However, a recent record in southern- most Lithuania (Lazdijai) close to the Polish northeastern border (Ivinskis & Rimsaite 2008) suggests a much more northern distribution also in Poland. Southern Poland (Galicia) had already been mentioned in the classic work by Harz (1957) as that part of this coun- try which is inhabited by P. falcata. Even much earlier, Zacher (1917) had pointed on a doubtful voucher speci- men of the Wroclaw (formerly Breslau) Museum from “Schlesien” (= Silesia), but he claimed that Pylnov (1913) had already recorded this species more northernly of Wro- claw, viz. from “Nova Alexandria, Russisch Polen” (= “Russian Poland”). This place, today Pulawy (51.26N; 21.59E) can be found on older maps as situated between Radom and Lublin, on the Visla river south of Deblin: 51.34N; 21.50E. Several new Polish references give a more detailed and also extended picture of the distribu- tion range of P. falcata in Poland (Bazyluk & Liana 2000, Koéarek 2000, Orzechowski 2009). The locality data pub- lished by these authors document this species to be more widespread in the central part of Poland, the most north- western records (Lubuskie Province: Orzechowski 2009) being adjacent to the relatively new records from Bran- denburg (Landeck et al. 2005). The two northern Polish regions Pojezierze Pomorskie and Mazurskie which to- gether roughly comprise the northern third of the coun- try, were so far lacking Phaneroptera falcata records. The new and unexpected find of this species in southern Lithuania (Ivinskis & Rimsaite 2008), however, made the occurrence of P. falcata likely also in northern Poland . On August 13, 2010 two of us (WB & PW) passed through northeastern Poland towards Lithuania. On road no. 16 east of Wigierski National Park, between Serski Las vil- lage and Sejny, at 53.55N and 23.09E, when searching for lizards on a spruce forest clearing, we happened to find an adult female of P. falcata (Fig. 1) which in view of what is said above seems to be a remarkable record, as it ex- Fig. 1. The voucher specimen of Phaneroptera falcata from E of Wigierski National Park, Northeast Poland. Photograph: P. Geissler. 110 Wolfgang Bohme et al. Fig. 2. Habitat east of Wigierski National Park where our P. falcata record was found. Photograph: W. Bohme. tends the known range within Poland for about (appr.) 300 km northeastwards and immediately links it with the first Lithuanian record from 2008. The locality (Fig. 2) 1s sit- uated less than 20 km from the Lithuanian border and closely corresponds to the new and single Lithuanian lo- cality of this thermophilous species which is situated in the Lazdiyai district at 54.12N and 23.50E (Ivinskis & Rimsaite 2008) (Fig. 3). Our specimen is deposited in the Orthoptera collection of the Zoologisches Forschungsmu- seum A. Koenig (ZFMK) in Bonn. According to Zuna-Kratky et al. (2009), adults in Austria appear about on mid-July but start their main adulthood season from mid-August. This agrees with our female voucher specimen, which was also adult, despite its much more northernly situated locality. Road no. 16 from Augustow via Serski Las and Sejny runs in parallel to the main road (no. 8, via Suwalki to Mar- jampole in Lithuania) and has much less traffic than the latter. This alone makes it unlikely that the specimen of P. falcata could have been passively displaced by human transportation. Rather, this thermophilous species demon- strated its potential for a natural, northeastward-directed range extension not only in Germany and the Benelux countries with a predominantly oceanic climate, but also in NE Poland, under much more continental climatic con- ditions and even reached Lithuania. The several individ- uals registered there underline the existence of a popula- tion in this area rather than displaced single individuals. Intensive faunistic search is necessary to assess the dis- tribution range and its dynamics of Phaneroptera falca- ta in this region. The eastward continuation of the northern borderline of P. falcata through Belarus and the European part of the Russian Federation also needs more faunistic research ef- forts. According to the map in Willemse (2007), P. falca- Bonn zoological Bulletin 60 (1): 109-111 ta is not yet known from Belarus. In Ukraine, the exact northern borderline has yet to be assessed but runs, accord- ing to Kotenko (in litt., November 2010), between 52 and 53N. In the European part of the Russian Federation, it is marked by localities situated approximately on a similar latitude as the formerly known Polish findings, e.g. Sevsk (Fig. 3: 12), Kursk (Fig. 3: 13), and Lipetsk (Fig. 3: 14) in the southern part of the Ryazan area (Bey-Bienko 1954). A newly collected voucher specimen from the southern part of the Bryansk area close to the Ukrainian border (Chukrai village, district of Suzemka, 52.19N; 34.05E, collected by one of us (PG) in July, 2010 and also deposit- ed in ZFMK’s Orthoptera collection), roughly fits this dis- tributional pattern (Fig. 3: 11), which ranges between 51 and 51.40N in the Asiatic part of Russia (Bey-Bienko 1954, 1964). It can be assumed, however, that P. falcata will extend its northern distribution borderline also in these parts of its range. Fig. 3. | Map showing the northern part of the distribution range of P. falcata in the eastern Central and East European realm, our new record (2) and the one from Lithuania (1) being the northernmost sites. 1. Lazdijai District (LIT); 2. Wigierski National Park (PL); 3. Treuenbrietzen (D); 4. Berkenbriick (D); 5. Lubuski Province (PL); 6. Wroclaw (PL); 7. Nizina San- domierska (PL); 8. Roztocze National Park; 9. Pulawy (PL); 10. Polesie National Park (PL); 11. Chukrai (RUS); 12. Sevsk (RUS); 13. Kursk (RUS); 14. Lipezk (RUS). Map: P. Wagner. Acknowledgements. The authors are indebted to Prof. Dr. Zbig- niew Szyndlar and to Dr. Elsbieta Warschowska (both from Krakow), as well as to Ryszard Orzechowski (Zielona Gora), and to Tatiana Kotenko (Kiev), for valuable advice and literature ref- erences. PG thanks Igor Palko (Moscow) for the invitation to Chukrai village. OZFMK A remarkable record of Phaneroptera falcata (Poda, 1761) from north-eastern Poland 111 REFERENCES Bazyluk, W & Liana A (2000) Prostoskrzydle Orthoptera. Kat- alogu fauny Polski 58: 1-156 Bey-Bienko GY (1954) Fauna SSR. Pryamokrylye, Moskva, Leningrad (Izd. Akad,. Nauk SSSR), vol. 2, no. 2: Kuznechikovye, podsem. Listobye kuznechiki (Phaneropteri- nae): 394 pp. Bey-Bienko GY (1964) Opredelityel nasekomykh Evropeskey chasti SSSR. Tom 1. Moskva, Leningrad (Nauka): 935 pp. Detzel P (1998) Die Heuschrecken Baden-Wiirttembergs. Stutt- gart (Ulmer): 580 pp. + 20 pls Grein G (2000) Zur Verbreitung der Heuschrecken (Saltatoria) in Niedersachsen und Bremen., Stand 10.4.2000. Informati- onsdienst Naturschutz Niedersachsen 20: 74-112 Grein G (2007) Zur Ausbreitung von Phaneroptera falcata (Po- da, 1761) und Conocephalus fuscus (Fabricius, 1793) in Nie- dersachsen. Articulata 22: 91—98 Harz K (1957) Die Geradfltigler Mitteleuropas. Jena, G. Fischer: 494 pp. Ivinskis P & Rimsaite J (2008) Phaneroptera falcata (Poda, 1761) (Orthoptera, Phaneropteridae) in Lithuania. Acta Zoo- logica Lituanica 18: 270-272 Koéarek P (2000) Orthopteroid insects (Orthoptera, Blattaria, Dermaptera) of the Polesie National Park and its surroundings. Parki i Rezerwaty Przyrody 19: 89-97 Koéarek P, Holusa J, VIk R, Marhoul P & Zuna-Kratky T (2008) Recent expansion of the bush-crickets Phaneroptera falcata and Phaneroptera nana (Orthoptera: Tettigoniidae) in the Czech Republic. Articulata 23: 67—75 Bonn zoological Bulletin 60 (1): 109-111 Kohler G (2010) Fauna der Heuschrecken (Ensifera et Caelife- ra) des Freistaates Thiiringen. Naturschutzreport Jena 17: 378 pp. Maas S, Detzel P & Staudt A (2002) Gefahrdungsanalyse der Heuschrecken Deutschlands. Verbreitungsatlas, Gefahrdungs- einstufung und Schutzkonzepte. Bonn (BfN): 401 + xvi pp. Orzechowski R (2009) Obserwacje wybranych gatunkow pro- stoskrzydlych (Orthoptera) w poludniowe} czesci woje- wodztwa Lubuskiego. Przeglad Przyrodnicyyzy 20: 45-50 Pylnov E (1913) Materialy po faunye pryamokrylych (Orthoptera saltatoria) russkoj Polshi. Russkoye entomologichnoye obozryenie, St. Peterburg 13: 85—98 Willemse F (2007) Fauna Europaea: Ensifera. In: Heller KG (ed.): Fauna Europaea, version ileshs http://www. faunaeur.org.2007 Zacher F (1917) Die Geradfliigler und ihre Verbreitung. Jena, G. Fischer: 287 pp. Zuna-Kratky T, Karner-Ranner E, Lederer E, Braun B, Berg, H- M, Denner M, Bieringer G, Ranner A, Zechner L (2009): Ver- breitungsatlas der Heuschrecken Ostésterreichs. Wien (Natur- historisches Museum): 303 pp. Received: 17.01.2011 Accepted: 23.02.2011 Corresponding editor: F. Herder ©ZFMK i oo et i Sy , Bratt i} my _ : 7 TE oC Thi te ae ’ aa i ' ion ‘ « i i 1 ‘Ay i 1 1 Pah F fT G I ; ( ; we : io ee Bon zoological Bulletin Volume 60 | Issue | p. 113 Bonn, May 2011 Erratum to: Sonnenberg & Busch (2010) Description of Callopanchax sidibei (Nothobranchiidae: Epiplateinae), a new species of killifish from southwestern Guinea, West Africa Rainer Sonnenberg !.2 & Eckhard Busch 3 ' Zoologisches Forschungsmuseum Alexander Koenig, Adenauerallee 160, D-53113 Bonn, Germany; Corresponding author; E-mail: r.sonnenberg.zfmk@uni-bonn.de 2 Max-Planck-Institut fiir Evolutionsbiologie, August-Thienemann-Strasse 2, D-24306 Plén, Germany 3 Diederichsstrasse 45, D-42855 Remscheid, Germany Erroneously the name of the new species was formed as a noun in genitive case for a single male person (ICZN, Article 31.1.2), however, in the text it refers to: Sex. The new species is named after Mr. Samba Sidibe and his family.... (Sonnenberg & Busch 2010, p. 11, bold in- troduced here for clarification). Therefore the species name should be formed in plural with the ending -orum. In accordance with the International Code of Zoological Nomenclature (4th Edition, 1999), Article 32, we propose as correct species name Callopanchax sidibeorum. Acknowledgements. We thank Wolfgang Béhme (ZFMK, Bonn) for alerting us on this mistake in the publication and his advice for this Erratum. REFERENCES International Code of Zoological Nomenclature, 4‘ edition, 1999. International Trust for Zoological Nomenclature, Lon- don Sonnenberg R, Busch E (2010) Description of Callopanchax sidibei (Nothobranchiidae: Epiplateinae), a new species of kil- lifish from southwestern Guinea, West Africa. Bonn zoolog- ical Bulletin 57: 3-14 114 Book Review Brandt T, Jiilch C, Wasmer K, Moning C, Wagner C (2011) The Top 50 sites for birdwatching in Germany. (Die 50 besten Vogelbeobachtungsplatze in Deutschland). Der Falke. Aula-Verlag, Wiesbaden. ISBN 978-3-89104-746- QO: 19:95'€: This book is published as a special issue of one of the lead- ing German birdwatcher’s magazines, “Der Falke”, and large parts of its contents have already been published in single articles of the journal. It presents the 50 best sites for birdwatching in Germany. Each single chapter gives an overview about typical habitats and interesting bird species and some logistical advices of the where and when of birdwatching at the spot. Sometimes, interesting addi- tional information on other cultural or natural study ac- tivities in the region is given. Boxes show “How to get there” and give useful local contact addresses. In contrast, explicitly stated GPS coordinates are often not very help- ful as they regularly do not show the particular spots from where to find the birds (e.g. a hide) but the center of the next village. A register at the end of the volume shows which rare species occur at each site. A Top 50 list of sites can always be debated. However, it is a pity that some species which are much sought after are not covered by the selected sites (e.g. Rock Bunting) or are not mentioned at suitable sites (e.g. Cirl Bunting). Nice photos and maps make it a pleasure to browse in the well illustrated book and to think about the next short trip. However, if you are seriously interested in birdwatching in Germany, the three volume book “Vogel beobachten in Nord-, Siid- und Ost- deutschland” published by Kosmos and written by mem- bers of the same author team might be a better and more comprehensive (but more expensive) choice. Germany still lacks a concise English “Where to watch birds” guide, and authors and editors sadly failed to fill this gap. Instead, recycled material was used to write a nice although not really essential book for birdwatchers. 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Does the abstract summarize only the significant findings? . Is the length of the paper appropriate? . Are all (colour) figures and tables necessary and appropriate? Nn Oo on DW SMITHSONIAN INS Contents MIMO Van Rooijen, Johan, Chan Kin Onn, L. Lee Grismer & Norhayati Ahmad: Estimating the herpetofaunal species richness of Pangkor Island, Peninsular Malaysia Hi i 3 9088 01617 336 Geissler, Peter, Truong Quang Nguyen, Nikolay A. Poyarkov & Wolfgang Bohme: New records of snakes from Cat Tien National Park, Dong Nai and Lam Dong provinces, southern Vietnam Vogel, Gernot & Johan van Rooijen: Description of a new species of the genus Dendrelaphis Boulenger, 1890 from Myanmar (Squamata: Serpentes: Colubridae) Wagner, Philipp, Wolfram Freund, David Modry, Andreas Schmitz & Wolfgang Bohme: Studies on African Agama IX. New insights into Agama finchi Bohme et al., 2005 (Sauria: Agamidae) with the description of a new subspecies Bohme, Wolfgang, Mark-Oliver Rodel, Christian Brede & Philipp Wagner: The reptiles (Testudines, Squamata, Crocodylia) of the forested southeast of the Republic Guinea (Guinée forestiére), wit a country-wide checklist IE 1 Ahmadzadeh, Faraham, Fatemeh Khanjani, Aref Shadkam & Wolfgang Bohme: A new record of the Persian Brook Salamander, Paradactylodon persicus (Eiselt & Steiner, 1970) (Amphibia: Caudata: Hynobiidae) in northern Iran Bohme, Wolfgang: Type list of amphibians and reptiles in the Zoologisches Forschungsmuseum A. Koenig, Bonn: corrections and additions Van der Zee, Jouke R. & Rainer Sonnenberg: Aphyosemion musafirii (Cyprinodontiformes: Nothobranchiidae), a new species from the Tshopo Province in the Democratic Republic of Congo, with some notes on the Aphyosemion of the Congo basin Valdesalici, Stefano & Kiril Kardashev: Nothobranchius seegersi (Cyprinodontiformes: Nothobranchiidae), a new annual killifish from the Malagarasi River drainage, Tanzania Gongora, Jaime, Cibele Biondo, Jennifer D. Cooper, Andrew Taber, Alexine Keuroghlian, Mariana Altrichter, Fabricia Ferreira do Nascimento, Amanda Y. Chong, Cristina Yumi Miyaki, Richard Bodmer, Pedro Mayor & Susana Gonzalez: Revisiting the species status of Pecari maximus van Roosmalen et al., 2007 (Mammalia) from the Brazilian Amazon Irmler, Ulrich: Two new species and a new record of the genus Neolindus Scheerpeltz, 1933 (Coleoptera: Staphylinidae: Paederinae) Bohme, Wolfgang, Peter Geissler & Philipp Wagner: A remarkable record of Phaneroptera falcata (Poda, 1761) (Saltatoria: Phaneropteridae) from north-eastern Poland Sonnenberg, Rainer & Eckhard Busch: Erratum to: Sonnenberg & Busch (2010) Description of Callopanchax sidibei (Nothobranchiidae: Epiplateinae), a new species of killifish from southwestern Guinea, West Africa Cover illustration: Naja sumatrana Miller, 1890 from Pangkor Island, Peninsular Malaysia (this volume, pp. 3-8) Ministerium fiir Innovation, Wissenschaft und Forschung des Landes Nordrhein-Westfalen P, vay /)% tf Leibniz S. Gemeinschaft IN TITUTION LIBRAR Ss 3 17 25 35 63 67 73 89 95 103 109 113