Bonn Volume 5/7 zoological ae Bulletin tormerly: Bonner zoologische Beitrage 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, 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 0006-7172 Produced by Eva-Maria Levermann, Kaiserstr. 151, 53113 Bonn, Germany; emlevermann@netcologne.de. Printed and bound by DCM, Werner-von-Siemens-Str. 13, 53340 Meckenheim, Germany. Bonn zoological Bulletin Editor-in-Chief Fabian Herder, Zoologisches Forschungsmuseum Alexander Koenig (ZFMK), Ichthyology Section, Adenauerallee 160, 53113 Bonn, Germany, tel. +49 228-9122-255, fax: +49 228-9122-212; f-herder.zfmk@uni-bonn.de Editorial Board Dirk Ahrens, Insects: Coleoptera, ZFMK, tel. +49 228-9122-286, fax: +49 228-9122-332; d.ahrens.zfmk@uni-bonn.de Wolfgang Bohme, Amphibians and Reptiles, ZFMK, tel. +49 228-9122—250, fax: +49 228-9122-212; w.boehme.zfmk@uni-bonn.de Netta Dorchin, Insects: Diptera, ZFMK, tel. +49 228-9122-292, fax: +49 228-9122-212; n.dorchin.zfmk@uni-bonn.de Renate van den Elzen, Birds, ZFMK, tel. +49 228-9122—231, fax: +49 228-9122-212; r.elzen.zfmk@uni-bonn.de Bernhard Huber, Invertebrates except Insects, ZFMK, tel. +49 228-9122-294, fax: +49 228-9122-212; b.huber.zfmk(@uni-bonn.de Rainer Hutterer, Mammals, ZFMK, tel. +49 228-9122-261, fax: +49 228-9122-212: r.hutterer.zfmk@uni-bonn.de Gustav Peters, Mammals, ZFMK, tel. +49 2289122262, fax: +49 2289122212: g.peters.zfmk@uni-bonn.de Bradley Sinclair, Canadian National Collection of Insects, Ottawa Plant Laboratory — Entomology, CFIA, K.W. Neat- by Bldg., C.E.F., 960 Carling Ave., Ottawa, ON, Canada KIA 0C6, tel. + 1 613-759-1787, fax: + 1 613-759-1927; bradley.sinclair@inspection.gc.ca Dieter Sttining, Insects except Coleoptera and Diptera, ZFMK, tel. +49 228-9122-—220, fax: +49 228-9122-212; d.stuening.zfmk@uni-bonn.de Advisory Board Theo C. M. Bakker, Rheinische Friedrich-Wilhelms-Univer- sitat, Institut fiir Evolutionsbiologie & Okologie, 53113 Bonn, Germany, tel. +49 228—73—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 22628-7304, fax: +48 22-629-6302; wieslawb@miuiz.waw.pl Matthias Glaubrecht, Museum fiir Naturkunde Berlin, Leib- niz-Institut fiir Evolutions- und Biodiversitatsforschung an der Humboldt-Universitat zu Berlin, Invalidenstrasse 43, 10115 Berlin, Germany, tel. +49 30-2093—8504/ 8400, fax: +49 030-2093-8565; matthias.glaubrecht@mfn-berlin.de Jurgen Haffer, Tommesweg 60, 45149 Essen, Germany, tel. +49 201—710426; j.haffer@web.de Jeremy D. Holloway, The Natural History Museum, Depart- ment of Entomology, Cromwell Road, London, SW7 5BD, U.K.; j.holloway@nhm.ac.uk Boris Krystufek, Slovenian Museum of Natural History, P. O. Box 290, Ljubljana, Slovenia; boris.krystufek@zrs.upr.si Wolfgang Schawaller, Staatliches Museum ftir Naturkunde, Rosenstein 1, 70191 Stuttgart, Germany, tel. +49 711-8936—221, fax: +49 711—8936-100; schawaller.smns@naturkundemuseum-bw.de Ulrich K. Schliewen, Department of Ichthyology, Bavarian State Collection of Zoology, Miinchhausenstr. 21, 81247 Munchen, Germany, tel. + 49 89-8107—110; schliewen@zsm.mwn.de Michael Schmitt, Ernst-Moritz-Arndt-Universitat, Allge- meine & Systematische Zoologie, Anklamer Str. 20, 7489 Greifswald, Germany, tel. +49 3834-86-4242, fax: +49 3834864098; michael.schmitt@uni-greifswald.de W. David Sissom, Dept. of Life, Earth and Environmental Sciences, W. Texas A. & M. University, WTAMU Box 60808, Canyon, Texas 79016, USA, tel. +1 806-651-2578, fax: +1 806-651-2928; dsissom@mail.wtamu.edu Miguel Vences, Technical University of Braunschweig, Zoo- logical Institute, Spielmannstr. 8, 38106 Braunschweig, Germany, tel. + 49 531-391-3237, fax: + 49 531-391-8198; m.vences@tu-bs.de Erich Weber, Eberhard-Karls-Universitat, Zoologische Schausammlung, Sigwartstr. 3, 72076 Tiibingen, tel. +49 7071—2972616, fax +49 7071—295170; erich.weber@uni-tuebingen.de Editorial Welcome to the first issue of BzB in 2010. The journal has been subject to major recent changes, including title and scope, editor, boards, cover design, and free availabil- ity of all contributions (open access). The title “Bonner zoologische Beitrage” changed to “Bonn zoological Bulletin”. This follows the fact that the jour- nal’s contributions, like elsewhere in biological sciences, are nowadays nearly exclusively published in English. The new title shall also contribute to increasing the journal’s publicity while keeping the well-known acronym “BzB”. The journal focuses now explicitly on three fields, sharp- ening its traditional profile as a museum journal: (1) tax- onomy, (2) systematics and evolution, and (3) biodiver- sity and biogeography, all with respect to terrestrial ani- mals (including fresh or brackish waters). Publication in BzB continues to be free of charge, includ- ing colour figures, open access, retention of full rights with the authors, and production of a widely distributed print version. Manuscripts will be published online ahead of print directly after acceptance, allowing immediate pub- lic access to the latest contents. The BzB webpage (see interior cover for address) is still under revision, but con- tains already all published issues back to 1993. Our goal for the near future is to make the contents of all past vol- umes available online. In summary, the concept of BzB will guarantee fast and easy availability of the published contents. Some of the editional responsibilities changed include the editor-in-chief and the boards. Michael (“Theo’’) Schmitt, responsible for BzB since 2003, left Museum Koenig for Greifswald University last year. Likewise, the boards have been subject to some rearrangement. On behalf of the “BzB-team’’, I thank Theo and the former editorial and ad- visory board members for their invaluable engagement. Theo will however stay with BzB as a member of the Ad- visory Board. A warm “welcome on Board” to those col- leagues joining or staying with the advisory or editorial team of Bonn zoological Bulletin, I am looking forwards to working with you. JUL 07 2010 LIBRARES Despite of the changes outlined above, the journal will re- tain its core strengths as a museum journal. The board members cover a wide array of expertise in zoology, and share common interests related to the journal’s focus. [am convinced that this is a sound basis for the journal’s fu- ture, building on the history of “Bonner zoologische Beitrage” since 1950. Contributions published in the present issue of BzB re- flect a wide array of approaches and methods applied in organismal zoology. Besides a checklist and contributions dealing with morphology, autecology or host-parasite in- teraction, the contents of the present issue include formal descriptions of six species new to science, including a chameleon, a gecko, a killifish (see front cover), two new species of moths, and a new fossil aquatic dance fly from Baltic amber. Taxonomic papers like these contribute to the exploration of our planet’s still only partly known bio- ta, and are increasingly not in the focus of interest of many present zoological journals. Publishing sound taxonomic work remains a core task for museum journals like BzB. Finaily, some words regarding cover design and text for- mat. In 2003, the BzB front cover was changed from com- pletely yellow to the design known until 2009 when the journal switched to a larger format. At this time, the ad- ditional cover space was contrasted in light grey. The pres- ent design by Uwe Vaartjes takes the next step, replacing the yellow colour completely by grey and neutral ele- ments, and incorporates Regina Liitzenkirchen’s idea of including Bonn’s heraldic animal, the “King (Koenig in German) of Animals”. I am grateful to both for their work. Some readers and authors will have to get used to the more simplified literature style, which is a substantial reduction of the workload in production for both, authors and edi- tors. I hope that the current revision of BzB in terms of both, focus and style, will meet the approval of the jour- nal’s audience. With best wishes to all readers, authors and ‘““BzB-Team” members, Tibi lect Fabian Herder (Editor-in-Chief; Fish Curator at ZFMK) Bonn, April 2010 Bonn zoological Bulletin Volume 57 Issue | pp. 3-14 Bonn, April 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-53223 Bonn, Germany. Corresponding author. E-mail: r.sonnenberg.zfmk@uni-bonn.de 2 Max-Planck-Institut fiir Evolutionsbiologie, August-Thienemann-Strasse 2, D-24306 Plon, Germany 3 Diederichsstrasse 45, D-42855 Remscheid, Germany Abstract. A new species of the annual killifish genus Callopanchax (Nothobranchiidae: Epiplateinae) is described from coastal Guinea, West Africa. It is distinguished from its congeners by morphological and colour pattern characters in- cluding up to three irregular and interrupted horizontal stripes on blue flanks in males. A preliminary DNA study indi- cates that it is closely related with C. occidentalis and C. toddi, which is supported by the following apomorphic char- acters compared to the more distantly related C. monroviae: a dark red or black postopercular blotch and an origin of the dorsal fin anterior to or above the first ray of the anal fin. The species first turned up in commercial imports of ornamen- tal fish from Guinea. The local collectors, the Sidibe family, are acknowledged for sharing their knowledge on this species and their support to the second author during his collections by naming it Callopanchax sidibei, new species. Currently it is only known from the type locality and is endemic to Guinea. Résumé. Une nouvelle espéce annuelle du genre Callopanchax (Nothobranchiidae: Epiplateinae) est décrite des cdtes maritimes de Guinée, Afrique de l|’Ouest. Elle se distingue de ses congeneres par certains caracteres morphologiques et un motif coloré comptant jusqu’a trois bandes horizontales irréguliéres et interrompues sur les flancs bleus chez les ma- les. Une étude ADN préliminaire indique que ce taxon est proche de C. occidentalis et C. toddi. Ceci est également sup- porté par les caractéres apomorphes suivants — comparés a C. monroviae, de parenté plus éloignée: tache post-opercu- laire rouge sombre ou noire, et origine de la nageoire dorsale au-dessus ou antérieure au premier rayon de la nageoire anale. L’espéce a été détectée en premier dans les importations commerciales d’espéces ornementales en provenance de Guinée. Callopanchax sidibei sp. nov. est dédiée a la famille Sidibe, engagée dans les collectes locales, pour avoir par- tagé ses connaissances et avoir aidé le second auteur durant sa mission de collecte. Cette espece endémique de Guinée n’est actuellement connue que de la localité type. Key words. Cyprinodontiformes, Callopanchax sidibei, new species, taxonomy, endemic, Upper Guinea. INTRODUCTION The nothobranchiid genus Callopanchax Myers, 1933 consists of four described species of which three are cur- rently considered valid (Costa 2008; Romand 1985, 1992; Seegers 1997; Wildekamp 1996; Wildekamp & Van der Zee 2003). They are relatively large annual killifishes, reaching up to 90.0 mm in total length according to Wildekamp & Van der Zee (2003), living in the coastal plains from southwestern Guinea and Sierra Leone to western Liberia. Callopanchax occidentalis (Clausen, 1966) and C. toddi (Clausen, 1966) were described from Sierra Leone, the latter as a subspecies of C. occidental- is. Some years later two further species, C. monroviae (Roloff & Ladiges, 1972) from Liberia, and C. huwaldi (Berkenkamp & Etzel, 1980) from Sierra Leone, were for- mally described. The latter species is currently considered a synonym of C. occidentalis by most authors (Costa 2008; Lazara 2001; Romand 1985; Wildekamp 1996; Wildekamp et al. 1986). Callopanchax species are easily separated from all other species groups within the Epiplateinae by the large num- ber of anal and dorsal fin rays and an origin of the dorsal fin anterior or slightly posterior to origin of the anal fin; in the remaining genera the dorsal fin inserts distinctly posterior to the anal fin origin. They share, with the re- maining Epiplateinae, the frontal and preopercular neu- romast pattern (Clausen 1966, 1967; Grimm 1974; Van der Zee 2002; Van der Zee & Wildekamp 1994), a red pig- mentation pattern on the side of the head (Van der Zee 2002), and with Archiaphyosemion guineense (Daget, Rainer Sonnenberg & Eckhard Busch Table 1. List of specimens used for DNA sequencing with voucher sample number, collection locality, country, and GenBank ac- cession numbers. Accession numbers with * are published in Murphy & Collier (1997), with ** in Sonnenberg & Busch (2009). Aquarium strain means that the origin in nature for this sample is unknown. Sample no. _ species Collection locality Country GenBank acc. no. OCC GB Callopanchax occidentalis aquarium strain Sierra Leone U73242* OCC1530 Callopanchax occidentalis SL 03/11, Malai Sierra Leone GU553007 OCC1531 Callopanchax occidentalis SL 89, Teme Yellah Sierra Leone GU553008 OCC1532 Callopanchax occidentalis SL 03/19, Magbenta Sierra Leone FJ872034** TOD 1534 Callopanchax toddi GM 97/26, Takhori Guinea GUS553009 TODI535 Callopanchax toddi GM 97/24, Mola Guinea GU553010 cTOD1758 ~~ Callopanchax cf. toddi GM 97/3, Mangata Badjana Guinea GU553011 SID1536 Callopanchax sidibei GM 97/2, Bombokore Guinea GU553012 MON1533 ~~ Callopanchax monroviae L 97, Paynesville Liberia GU553013 GER1381 Scriptaphyosemion geryi SL 03/9, Robis 1 Sierra Leone FJ872033** GUI1404 Archiaphyosemion guineense SL 93/37, Lenghe Curoh Sierra Leone FJ872031** 50 km Fig. 1. Tay, Map of Callopanchax collection localities in coastal West Africa. The letter T indicates the respective type localities, D indicates origin of Callopanchax DNA samples used in this study. Black square = C. occidentalis, white square = C. toddi, black circle = C. cf. toddi, white circle = C. huwaldi, black triangle = C. monroviae, white triangle = C. sidibei. Bonn zoological Bulletin 57 (1): 3-14 ©OZFMK A new Callopanchax from Guinea 5 Table 2. Uncorrected p-distances for the mitochondrial LSU sequences with pairwise exclusion of missing data among nine sam- ples of Callopanchax, one of Archiaphyosemion, and one of Scriptaphyosemion. The upper triangle contains the linear geographic distances between the different collection localities of the sequenced Callopanchax specimens, except for OCC GB, which is ac- cording to Murphy et al. (1999) an aquarium strain of unknown origin. sample OCEGBROCES (OKELE OCC TOD TOD cIOD SID MON GER 1530 S31 N37 1534 3S) 1758 1536 1533 1381 OCC 1530 LEV — 204km 40km 196km 64km 106km 95km 370km— —- O€Een53)1 1S od! = 185km 396km 245km 301 km 289km 175km = — OC@1532 I) IES 1S — 230km 103km 144km 132km 340km— — TOD 1534 6.6 5.8 6.8 6.6 — 154km 97km 110km 565km— — TOD 1535 5) Se 6.2 6.0 ed = S57km 43km 418km — cTOD 1758 6.8 5.6 6.6 6.4 Del 1S = 14km 472km— — SID 1536 7.6 6.7 eS Toll HES 6.5 6.2 = 458km — MON 1533 14.2 W23) 12.8 Salk WS LES IES HSA = = GER 1381 9.6 Jai) 8.7 9.3) OFS) OF) 10.1 10.6 14.2 — GUI 1404 IN 10.8 11.3 10.8 10.8 10.8 11.6 12.4 14.8 9.4 1954) and Scriptaphyosemion Radda & Piirzl,1987 a dis- tinct constriction of the subdistal portion of the posterior arm of the parasphenoid (Costa 2009). The phylogenetic placement of Callopanchax within the Epiplateinae and its sister group relationship with Scriptaphyosemion 1s sup- ported by molecular data (Murphy & Collier 1997; Mur- phy et al. 1999; Sonnenberg & Busch 2009). In 1992 a scientifically unknown Callopanchax species turned up in a commercial import of ornamental fish from Guinea to the Netherlands. Its distinctness was first rec- ognized by the unique colour pattern of this species (Busch 1993; Wildekamp 1996; Wildekamp & Van der Zee 2003) and was later supported by a study based on molecular da- ta (Murphy et al. 1999). During several research trips in Guinea, the second author was able to collect specimens of this undescribed Callopanchax, which is herein de- scribed as Callopanchax sidibei, new species. MATERIAL AND METHODS The specimens on which the following description is based were deposited in the ichthyological collections of the Musée Royal de |’Afrique Centrale (MRAC), Tervuren, Belgium, and the Zoologisches Forschungsmuseum Alexander Koenig, Bonn, Germany (ZFMK). Compara- tive material of the other species is listed below. Measurements and counts are made as indicated in Son- nenberg & Busch (2009), transverse scale count was made between the anterior insertion of the pelvic fin and dor- sal fin; the very small scales on the belly were not count- Bonn zoological Bulletin 57 (1): 3-14 ed. Digital X-ray images of the types and some specimens of the comparative material used for unpaired fin ray and vertebrate counts were made with a Faxitron LX-60 Dig- ital Specimen Radiography System at the ZFMK. Caudal fin ray counts were only made from X-ray images and in- clude all visible rays. We prepared a preliminary DNA dataset of a fragment of the mitochondrial large ribosomal subunit (16S rDNA) for a small number of specimens, including three of C. occi- dentalis from Sierra Leone, one sequence of C. occiden- talis from GenBank (accession number U73242, Murphy & Collier 1997), two specimens of C. toddi and one of C. cf. toddi from Guinea, one specimen of each C. monrovi- ae and C. sidibei, new species, and, as an outgroup, Archi- aphyosemion guineense and Scriptaphyosemion geryi (Lambert, 1958). The latter two are the type species of the respective genera. Specimens are listed with locality in- formation and GenBank accession number in Table 1, methods for DNA extraction, amplification and sequenc- ing of the 16S rDNA fragment are provided in Sonnen- berg & Busch (2009), where also some of the sequences were published. Resulting sequences were aligned with ClustalX 1.8 (Thompson et al. 1997) and checked by eye in BioEdit 7.0.5.3 (Hall 1999). Sequences were tested for the anti-G bias characteristic of mitochondrial DNA (Zhang & Hewitt 1996). Uncorrected p-distances with the pairwise deletion of missing data were calculated in Mega 4.1 beta 3 (Tamura et al. 2007) and are given in Table 2. Analyses of sequence data were made with PAUP 4.0b10 (Swofford 1998) by maximum parsimony and with Mr- Bayes 3.1.2 (Huelsenbeck & Ronquist 2001) by a ©ZFMK 6 Rainer Sonnenberg & Eckhard Busch Fig. 2. Bayesian approach. Maximum parsimony analyses were done with the exclusion of uninformative positions, gaps were coded as a fifth character state, and heuristic tree searches were performed with random addition of se- quences for 1,000 replicates. Bootstrapping was performed with the random addition of sequences for 100 replicates and 10,000 bootstrap replicates. Bayesian analyses were performed with the number of generations set to 10,000,000, with sampling every 1,000 generations, and the results of the first 1,000,000 generations were discard- ed as a burn-in time after checking that the runs had reached the stationary phase at this point of the analysis. We performed two runs for this dataset, one with two pa- rameter models (Nst 2) and one with six parameter mod- els (Nst 6), settings for rate=gamma and ngammacat=4. We chose to perform these two runs to check if there are serious differences in the results due to potential under- or Overparametrization, especially because of the small dataset with large genetic divergences (Table 2). We con- sider only nodes with 2 75 % bootstrap support and 2 95 % posterior probabilities as supported by the data. Additional information for the collection localities in Table 1 can be found in the comparative material list. Collec- tion points in Figure | are from museum collection data Bonn zoological Bulletin 57 (1): 3-14 A) Holotype of Callopanchax sidibei, male, ZFMK 41613, 58.5 mm SL. B) Paratype of Callopanchax sidibei, male, ZFMK 41614, 47.0 mm SL. C) Callopanchax sidibei, male with red dots in the anal and caudal fin, not preserved. D) Paratype of Callopanchax sidibei, female, ZFMK 41615, 55.4 mm SL. as accessed via Fishbase (Froese & Pauly 2009), from Hu- ber (2007), and collection data of the second author. We adopted, as a species concept, the definition by Moritz et al. (2000) which ,,...recognize as species geographically bounded sets of populations that are distinct for morpho- logical traits or are reproductively isolated from congeners, with or without corresponding molecular divergence...“. RESULTS Callopanchax sidibei, new species (Fig. 2A—D, Table 3) Roloffia sp. ,,auinea, CI 92°: Busch (1993), p. 64. Fundulopanchax species aff. toddi: Wildekamp (1996), pp. 212-213. Callopanchax sp. Guinea: Wildekamp & Van der Zee (2003), pp. 381-383. Callopanchax huwaldi (non Berkenkamp & Etzel, 1980): Neumann (2005), misidentification in fig. caption p. 48. Holotype. ZFMK 41613, male, 58.5 mm SL, Guinea, Guinée maritime, Kindia region, small river and adjacent pools and ditches in a secondary forest (9°32.71° N, ©ZFMK A new Callopanchax from Guinea 7 Table 3. Morphometric and meristic data of Callopanchax sidibei. All measurements in percentages of standard length except stan- dard length in mm. Fin ray counts were made on X-rays and include all visible rays. SD = standard deviation. Holotype &@ Types @Q Types all Types all Types all Types SD mean mean mean range Standard length 58.5 48.4 50.7 49.7 39.0—59.9 6.8 Total length 124.8 124.6 122.7 WBE 117.1-132.5 4.5 Head length 28.4 29.2 DD), 2) Il 27.7-30.5 0.9 Predorsal fin distance S10 // 57.6 60.1 59.0 55.4—63.2 1S, Prepelvic fin distance 54.1 52.0 Sal 52.4 49.1-56.6 2.0 Preanal fin distance 62.9 62.5 64.2 63.5 60.1—68.9 be Greatest body depth 24.3 D3 Dre BI} 20.0—25.8 1.6 Depth of caudal peduncle 1333 I WI 12.6 10.7-14.6 1.0 Length of caudal peduncle 23m 21.6 Japa) AN) IE8=2551 lad Base of dorsal fin 28.3 28.2 Soi 26.8 23.5—-30.7 1:9 Base of anal fin D3) 23rl 18.2 20.3 17.6—24.6 2.6 Eye diameter VP 6.8 6.8 6.8 6.1—7.6 0.4 Interorbital distance IBo7/ 211 Wed IES, 10.3-13.7 0.8 Dorsal fin rays 19 19.8 19.8 19.8 19-21 0.8 Anal fin rays 7) eo 18.1 18.0 17-19 0.6 Dorsal / anal fin position -2 -2.8 -3.2 -3.0 -2— -4 0.7 Caudal fin rays 33 32.0 31.8 Shes) 29-34 72 Total number of vertebrae 33 Saal 32.6 32.8 32-34 0.7 Vertebrae with pleural ribs 15 1522 14.8 15.0 14-16 OW Vertebrae with haemal spines 18 We 17.8 17.8 WS 0.6 Lateral line scales 3444 Aros) seollasio) SI SollaeSioy Ses V4 riesies) 0.9+0.6 Transverse row of scales WP eZ 10.8 11.0 10-13 0.8 Scales around caudal peduncle 16 NS) 15.8 LY) 15=1'6 0.4 13°14.51° W), close to the small village Bombokoré, about 3 km from the closest larger village Fandi, collection lo- cality GM 08/2, E. Busch and B. Wiese, 25.10.2008, Fig- ure 2A. Paratypes. ZFMK 41614-41615, 1 male, 47.0 mm SL, 1 female, 55.4 mm SL, paratopotypes, collected with holo- type, Figure 2B & D. ZFEMK 41616-41628, 5 males, 8 females, 39.0—58.9 mm SL, paratopotypes, collected with holotype. MRAC 2010—02—P—1—5, 2 males, 3 females, 46.3—59.9 mm SL, paratopotypes, collected with holotype. Diagnosis. Callopanchax sidibei (Fig. 2A—D) is, togeth- er with C. toddi (Fig. 3A—F) and C. occidentalis (Fig. 4A & B), distinguished from C. monroviae (Fig. SA—C) by insertion of first dorsal fin ray slightly anterior to first anal fin ray versus a posterior insertion. The former three species are also distinguished from C. monroviae by a dark red to black postopercular blotch above insertion of pec- toral fin versus absence and a dark or black blotch on the Bonn zoological Bulletin 57 (1): 3-14 anterior dorsal fin rays instead. The postopercular blotch is present in females of C. sidibei, C. occidentalis, and C. toddi but often only faint; the dark blotch on anterior dor- sal fin in C. monroviae may not be visible in females. Cal- lopanchax sidibei, C. occidentalis, and C. toddi males have an anal fin with a white or bluish-white margin and usu- ally only a small number of red dots or blotches on fin center; no horizontal red band or only traces of red under the eye from lower lip to posterior border of opercle; postopercular red pigmentation on flank formed by up to three irregular red horizontal bands interrupted into blotch- es. Males of C. monroviae have a dark or dark-red anal fin margin, proximal red dots, blotches or broad vertical streaks, and streaks between fin rays on distal part of anal fin; a more or less complete red band from snout under the eye and often extending on side to origin of pelvic fin; postopercular red pigmentation consists of incomplete rows of small red dots on anterior side and less and irreg- ularly distributed red dots or a pattern of incomplete and irregular vertical stripes on posterior side. Males of C. sidibei can also be distinguished from C. monroviae by ©ZFMK Rainer Sonnenberg & Eckhard Busch Fig. 3. A) Male of Callopanchax cf. toddi, Mangata Badjana, Guinea, collection locality GM 08/3 (= GM 97/3). B) Female of Callopanchax cf. toddi, Mangata Badjana, Guinea, collection locality GM 08/3 (= GM 97/3). C) Male of Callopanchax toddi, Mo- la, Guinea, collection locality GM 09/17 (= GM 08/4 & GM 97/24). D) Female of Callopanchax toddi, Mola, Guinea, collection locality GM 09/17 (= GM 08/4 & GM 97/24). E) Male of Callopanchax toddi, Lamayah, Guinea, collection locality GM 08/11. F) Male of Callopanchax toddi, Takhori, Guinea, collection locality GM 09/12 (= GM 08/9 & GM 97/26). body colouration which is mainly blue to blue-green on sides and dorsally often a golden sheen to the anterior, ver- sus anteriorly pale blue to blue-green and dorsal and pos- terior half of body reddish, reddish-brown or orange, or in the blue form body completely pale to grey-blue (Fig. S5A-C). Bonn zoological Bulletin 57 (1): 3-14 Callopanchax sidibei can be distinguished from males of C. occidentalis and C. toddi by up to three irregular and interrupted horizontal stripes on flanks, an absence of ver- tical dark red or reddish-brown stripes or broad blotches on sides and a mainly blue to blue-green body colour ver- sus dark brown, brown or red vertical stripes or blotches ©ZFMK Anew Callopanchax from Guinea 9 Fig. 4. A) Male of Callopanchax occidentalis, Malai, Sierra Leone, collection locality SL 03/11 (= SL 93/54). B) Female of Callopanchax occidentalis, Malai, Sierra Leone, collection locality SL 03/11 (= SL 93/54). Fig. 5. A) Male of Callopanchax monroviae, aquarium strain from Paynesville, Liberia, collection locality L 97, red form. B) Male of Callopanchax monroviae, aquarium strain from Pay- nesville, Liberia, collection locality L 97, blue form. C) Female of Callopanchax monroviae, aquarium strain from Paynesville, Liberia, collection locality L 97. Bonn zoological Bulletin 57 (1): 3-14 and a more reddish brown or copper to golden body colour with blue only on ventral side. Exceptions are populations with rare blue males in C. occidentalis and, if valid, C. huwaldi, which show the same pattern as the ‘typical’ form but lack orange and yellow pigments, but not the red pig- mentation and the dark vertical stripes or blotches on pos- terior side (e.g. Berkenkamp & Etzel 1980; Eberl 1999, pictures of C. occidentalis from Mabeimah on p. 65; Roloff 1976). Large males of C. sidibei show only a slightly rugged anal and dorsal fin border and rounded or slightly pointed fin tips versus more pronounced rugged fin borders in most C. occidentalis and C. toddi, especially the posterior tip of the anal fin, which can show filamentous extensions. Females of C. sidibei can be distinguished from C. occi- dentalis and C. toddi females by mid-laterally blue to blue- green coloured scales and the absence of the dark brown vertical stripes or blotches, similar as in males versus blue to blue-green coloured scales only on anterior side, often with short vertical dark blotches dorsally on side. They are distinguished from females of C. monroviae by their pigmentation pattern of red horizontal stripes with broad blotches versus more regular horizontal rows of small red dots and the absence of metallic blue to blue-green coloured scales on side in C. monroviae. Description. See Fig. 2A—D for general appearance and Table 3 for morphometric and meristic data of the types. Callopanchax sidibei (Fig. 2A—D) shares with its con- geners the following combination of diagnostic characters within the Epiplateinae: large and robust species, dorsal and anal fins with a high number of fin rays, origin of dor- sal fin close to origin of anal fin. A species with strong sexual dimorphism, adult males usually larger, with larg- er fins and more colourful than females. Maximum ob- ©OZFMK 10 Rainer Sonnenberg & Eckhard Busch served SL in males 59.9 mm, in females 59.0 mm. Body cylindrical, outline from tip of snout to posterior end of dorsal and anal fin convex, on caudal peduncle dorsally and ventrally slightly concave to nearly straight. Greatest body depth approximately at half distance between ori- gins of pectoral and pelvic fins. Snout slightly pointed, mouth directed upwards, lower jaw longer than upper jaw; many irregularly distributed un1- cuspid, slightly curved teeth of different size, a small num- ber of larger ones on the outer row of upper and lower jaw. Closed frontal neuromast system in one groove, preoper- cular neuromast system with 5 pores. Total number of vertebrae 32-34, number of vertebrae with pleural ribs 14-16, with haemal spines 17—19. The X-ray images indicate that in about half of the studied specimens the hypural plates are fused, as no gap or in- dication of overlap 1s visible. Fins in males slightly larger than in females, base of dor- sal mean value 28.2 % of standard length (SL) in males versus 25.7 % SL, base of anal in males 23.1 % SL ver- sus 18.2 % SL in females. Dorsal and anal fin rounded or slightly pointed, no long filamentous extension, fin inser- tion in both sexes posterior to mid-body. In large adult males, dorsal and anal fin rays sometimes protrude from the fin membran, giving the impression of a rugged fin border. Papillae on anal fin rays in males. First 2—4 dor- sal fin rays anterior of origin of anal fin. Dorsal fin rays 19-21, anal fin rays 17-19. Caudal fin rounded or slight- ly truncated, 29-34 fin rays. The single specimen with 29 caudal fin rays has a small gap between the anterior dor- sally situated rays explaining the comparatively low val- ue. Pectoral fin origin on ventral half of side, posterior of opercle, rounded, extending posteriorly, not reaching ori- gin of pelvic fin. Pelvic fin small and slightly pointed, not or just reaching anal fin. Scales cycloid, body and head entirely scaled except throat, scales dorso-anteriorly larger then ventrally and posteriorly, frontal squamation of G-type, scales on lat- eral line 32-34 + 3-5 on caudal fin base, not all scales along lateral line with small grooves in center. Transverse row of scales above pelvic fin 10-13, circumpeduncular scale row 15-16. Live colouration. Males. (Fig. 2A—C) Dorsal from head to dorsal fin brown, some scales on back with red dots or scale borders, forming an irregular reticulated pattern to the end of the caudal peduncle; flanks blue to blue-green, dorsolaterally with a metallic copper or golden sheen on scales, belly blue to bluish-grey. Bonn zoological Bulletin 57 (1): 3-14 Red pigmentation pattern on head posterior of the eye typ- ical for Epiplateinae (Van der Zee 2002) with two poste- rior directed horizontal red stripes, originating from one point posterior of the eye to posterior border of opercle, and two more or less complete vertical stripes dorsally. On flanks three interrupted and irregular red stripes, up- per and lower stripes only extending up to the posterior insertion of dorsal fin, only the middle stripe, positioned slightly above mid-line of body, extends to the end of the caudal peduncle, in some males the dorsolateral stripe on- ly consists of a small number of red dots or red scale bor- ders. Posterior of the opercle and above insertion of pectoral fin usually 2—3 scales in two neighbouring rows with broad black or dark red margin, forming a black or dark red blotch. Unpaired fins blue to blue-green, with white or, rarely, yel- low-white margins and red submarginal stripe. Red pig- mentation pattern on unpaired fins variable. Dorsal fin with red dots or short streaks between fin rays, below red submarginal band a whitish zone. Anal fin without or on- ly with little red pigmentation beside submarginal red stripe, in some males a proximal row with a small num- ber of red dots (Fig. 2C). Caudal fin in some males with red dots or blotches, lower white margin broader then up- per and in some specimens slightly yellow-white. In males with a more rounded caudal fin shape marginal and sub- marginal stripes can be closed on posterior end of fin, in males with a more truncate fin shape they end on the pos- terior fin border (Fig. 2A—C). Pelvic fins bluish, with red submarginal and white marginal stripe, pectoral hyaline with a white or blue-white margin. Females. (Fig. 2D) Body brown to light brown, dorsally darker then ventrally, sometimes with a golden colour sheen on anterior side. Scales dorsally and laterally with dark border, forming a reticulate pattern. Lateral red pig- mentation pattern similar to males, usually less pro- nounced and paler. Broad dark brown to brownish-grey lateral stripe from posterior border of operculum above origin of pectoral fin to posterior end of caudal peduncle on upper half. This stripe is not always visible and espe- cially pronounced in stressed specimens. On upper half of opercle, posterior of the eye, extending along the dark stripe on the side irregularly distributed scales with metal- lic blue or blue-green colour, with the highest density an- teriorly. Fins except pectoral light brown and slightly hya- line, unpaired fins with a whitish margin, on dorsal and caudal with pale red submarginal line. Dorsal and caudal fin with red dots, on dorsal sometimes restricted to fin base. Pectoral fin transparent with whitish to light blue margin. ©ZFMK Anew Callopanchax from Guinea 1] Preserved in ethanol. Males. Head dark grey, body dark brown to grey, some traces of red dots and blotches on side, belly whitish, beige, or greyish, a ventral dark stripe posterior to level of pectoral fin insertion to anterior ori- gin of anal fin. Fins pale grey with a dark submargin and small whitish margin, except caudal fin with a ventral broad and dorsal thin whitish margin, posterior border dark brown to reddish. Females. Head laterally and dorsally dark grey, chin with a dark blotch on ventral inner side of lower lip, body dor- sally darker grey or brown, a dark stripe from posterior border of opercle to origin of caudal fin, dark stripe dor- sally with a sharp border, ventrally diffuse; below this stripe brown, anterior ventral side and belly whitish to light brown, on side traces of red spots and blotches. Fins light grey with a dark border, sometimes with a small white margin, in dorsal and caudal fin traces of red spots, very rare with proximal single row of red dots in anal. Etymology. The new species is named after Mr. Samba Sidibe and his family, who first collected this fish and made specimens available for the ornamental fish trade. It is named in acknowledgement of their high interest in this species, their effort for a sustainable use and protec- tion of the habitat, and providing important information for the description. Distribution and habitat. Currently C. sidibei is only known from the type locality close to the small village Bombokoré, despite some effort to find further occur- rences of this species in the vicinity. Bombokoré is the spelling in the Sousou language, it might have a different spelling on maps or gazetteers. The habitat is a depression with small water bodies, ditch- es and connected pools in a secondary forest adjacent to a small creek. The water in the creek is only slowly flow- ing and stagnant in pools and ditches. During the first col- lection in 1997, the second author measured an air tem- perature of 36°C, the water temperature in the creek was 27°C and 29°C in ditches, pH was about 5.2, conductiv- ity 10uS/cm. Water in the ditches was about 20-30 cm deep and clear, the bottom consists of of sand and some stones with a layer of decaying leaves and wood with no aquatic vegetation. Herbaceous plants were found along the edges. Callopanchax sidibei and Epiplatys fasciola- tus (Gunther, 1866) were found in ditches and pools with standing water. In places with only grass and no leaf cov- er, only Scriptaphyosemion geryi was recorded. During the collection in 2008, the depression was flood- ed. The stagnant or slowly flowing water was shaded by herbaceous vegetation and young trees on edges and some aquatic vegetation, especially Nymphea. Air temperature Bonn zoological Bulletin 57 (1): 3-14 was 31°C, water temperature 27°C, water depth about 60-80 cm maximum. Interestingly the specimens caught in 2008 were of different size, from juveniles with about 2.5 cm total length to full grown adults, and in good con- dition, whereas in 1997 all were fully grown and showed first signs of ageing. During the last visits in 2008 and 2009 only a little human impact on the forest could be de- tected. Callopanchax sidibei seems to prefer shaded areas of the water bodies with slow flowing or standing water. How- ever, the life cycle of this species depends on the flowing water of the creek, because there is a seasonal connection between the creek and the pools and ditches. During the rain season the creek floods the depression and provides habitat, fresh water and food sources for the growing ju- veniles. Later, the inflow is reduced and isolated or par- tially connected pools and ditches are formed, which dry out during the dry season. Within these water bodies re- production takes place and adult fish most likely die dur- ing the dry season. Callopanchax sidibei 1s, like its con- geners (e.g. Scheel 1968; Wildekamp 1996), an annual species with eggs showing a prolonged development of 2—9 month, apparently an adaptation to temporary water bodies. However, some adults might move during the dry season into the flowing creek, which is supported by the observation of the local people. DNA analyses. The complete alignment of the mitochon- drial large ribosomal subunit for 11 specimens contains 530 bp including gaps, the sequence lengths without gaps ranges between 518-526 bp; the GenBank sequence (U73242) is 46 base pairs shorter, missing positions were filled with N. Of the 530 bp 133 are variable and 61 were parsimony informative, calculated without gaps. All se- quences show an anti-G bias typical for mitochondrial se- quences (Zhang & Hewitt 1996). Calculated intraspecific p-distances for the samples of C. occidentalis as well as C. toddi and C. cf. toddi (Table 2) are higher than for the corresponding fragment in Nim- bapanchax (Sonnenberg & Busch 2009), ranging from 1.3—2.7, which is in the range of the interspecific p-dis- tances within the latter genus. Interspecific p-distances within Callopanchax range from 5.2—7.6 between C. oc- cidentalis, C. toddi, and C. sidibei, and up to 14.2 with the inclusion of C. monroviae. Interestingly the intraspe- cific distances, even between geographically close collec- tion localities, are comparatively high (Table 2). In all resulting phylogenies, C. monroviae splits off first with respect to the remaining three taxa, which form an unresolved trichotomy (Fig. 6); this is congruent with the observed differences in colouration and morphology. As in Murphy et al. (1999), the distinct position of C. sidibei ©OZFMK is supported. However, these results should be seen as a first preliminary insight into the species relationships in Callopanchax as the dataset is currently too small for de- veloping well supported, stable phylogenetic hypotheses, and should be expanded both in number of specimens and genes in further analyses, especially with regard to the large genetic distances between the species. DISCUSSION The distinctness of Callopanchax sidibei with regard to its congeneric species was recognized very early after the first commercial import to the Netherlands (Busch 1993; Wildekamp 1996), but nothing was known about its ori- gin, distribution, and habitat. The second author was able to collect this species in 1997 with the help of the Sidibe family, who are engaged in the ornamental fish trade and were probably the first to export this species from Guinea. Callopanchax sidibei is currently only known from one locality; other Callopanchax collections in this area con- tain either C. toddi or C. cf toddi (Fig. 1). Interestingly in wild males two different caudal fin shapes exist: a more elongate and truncate form (Fig. 2A) and a shorter and rounded one (Fig. 2C), of which only the lat- ter turns up in aquarium bred specimens. In general the red pigmentation pattern on the body and fins is quite vari- able (Fig. 2A—C). As for C. toddi, currently no other colour morphs are known from C. sidibei, unlike the orange (or red) and blue phenotypes of C. occidentalis and C. monroviae (Fig. 5A-C, see also figures e.g. in Eberl 1999; Seegers 1997; Van den Nieuwenhuizen 1974; Wildekamp 1996). These colour morphs are usually sympatric with the more com- mon form which is the blue morph in C. monroviae and the red in C. occidentalis (Berkenkamp & Etzel 2003; Wildekamp 1996; Wildekamp & Van der Zee 2003 ). One species, C. huwaldi, which 1s considered as junior syn- onym to C. occidentalis, differs from the latter species in external appearance only by the absence of yellow and or- ange pigments (see colour figure in Roloff 1976, p.523). There is still some discussion about the validity of the species; protein electrophoretic data and cross-breeding experiments by Berkenkamp & Etzel (2003) and Etzel & Berkenkamp (1981) show differences between the two species, but protein electrophoretic data by Romand (1985) do not differ. DNA data by Murphy et al. (1999) place C. hwwaldi close to C. occidentalis, but the data set was to limited for further conclusions. Up to now, several other collections from an area close to the type locality of C. huwaldi were made e.g. by the second author, which contained small numbers of blue Bonn zoological Bulletin 57 (1): 3-14 12 Rainer Sonnenberg & Eckhard Busch males between red ones. Roloff (1974) reported a sym- patric occurrence of C. occidentalis and C. toddi, collect- ed by Chaytor close to Njala, which is also within this area, and is most probably an occurrence of sympatric red and blue C. occidentalis males and not a sympatry of the re- ported species. The second author was able to breed the red and the blue phenotype from Largo (SL 89) and Mabeimah (SL 93/46) several times in the aquarium and it seems that the blue phenotype is recessive with regard to the red, which is sup- ported by the observation that the blue phenotype is usu- ally rare in nature (pers. observation E. Busch). The fe- males of both forms show no differences. In case of C. sidibei it can be excluded that it is just a colour form of C. toddi or C. occidentalis. Beside differ- ences in body colouration the red pigmentation pattern al- so differs from all populations of C. toddi and C. occiden- talis, whereas in the cases of C. monroviae and C. occi- dentalis, including C. huwaldi, only the yellow and orange pigments are lost but the red pigmentation pattern is sta- ble. This is also supported by DNA data, in which C. sidibei is placed as separate lineage related to C. occiden- talis and C. toddi (Fig. 6 and Murphy et al. 1999). The recent collections of the second author in Guinea pro- vide additional information about the distribution of Cal- lopanchax, and extends the areal of the genus further northwards along the coast (Fig. 1). It should be noted that several of the populations collected so far show combi- nations of characters, which do not agree with the current diagnoses of C. toddi and C. occidentalis (Clausen 1966; Wildekamp & Van der Zee 2003). Further studies will show whether these species are more variable than pre- viously known, or if the divergent populations represent distinct units which might be accepted as separate species. These fishes are currently listed here as C. cf. tod- di (see Fig. 1 and as example Fig. 3A & B). COMPARATIVE MATERIAL Callopanchax huwaldi: ZFMK 12663-12667, Sierra Leone, 20 km von Moyamba, Ngabu (8°10’ N, 12°28’ W). Callopanchax monroviae: ZFMK 41629-41630, Liberia, Paynesville, L 97 (6°17? N, 10°43’ W); ZFMK 41631-41634, aquarium bred, strain from Liberia, Pay- nesville, L 97 (6°17’ N, 10°43’ W); RS1533, DNA sam- ple, aquarium bred, strain from Liberia, Paynesville, L 97 (6°17’ N, 10°43’ W). Callopanchax occidentalis: ZFMK 41635-41643, first generation after collection, Sierra Leone, SL 89 Largo ©ZFMK A new Callopanchax from Guinea 13 TODI534 TODI535 eTODI758 OCC1532 OCC1530 SID1536 MONI533 GERI381 GUI1404 — 5 Fig. 6. B 99 cTOD1758 TOD1I534 TODIS535 OCC1530 OCC1532 98/96 | 100 MONI533 GERI381 GUI1404 —_—i 0.02 Results of the maximum parsimony (A) and Bayesian (B) analyses of the mitochondrial large ribosomal subunit (16S) fragment. Numbers on nodes are bootstrap values in A and posterior probabilities in B. In the latter, the first values are from the analysis with Nst=2, the second values from the analysis with Nst=6, if both have the same value, only one value is given. Only for nodes with 2 75 % bootstrap support and = 95 % posterior probabilities values are shown. (8°16’ N, 12°09’ W); ZFMK 41644-41647, Sierra Leone, SL 89 Magbenta (8°24 N, 12°56’ W); ZFMK 41648-41654, aquarium bred strain from Sierra Leone, SL 89 Ngabu (8°15’ N, 12°22’ W); ZFMK 41655-41657, Sierra Leone, SL 89 Romeni (8°53’ N, 12°43’ W); ZFMK 41658-41671, Sierra Leone, SL 89 Teme Yellah (7°41 N, 11°25’ W); RS1531, DNA sample, Sierra Leone, SL 89 Teme Yellah (7°41’ N, 11°25’ W); ZFMK 41672-41674, Sierra Leone, SL 89 Waanje River (7°50’ N, 11°22’ W); ZFMK 41675, Sierra Leone, Ma Barie, SL 93/39 (8°25’ N, 11°45’ W); ZFMK 41676-41681, Sierra Leone, Mog- bomoh Village, Saspoor Water, SL 99/12 (8°20 N, 12°49’ W); ZFMK 41682, Sierra Leone, Brama School, SL 03/7 (8°22’ N, 12°58’ W); ZFMK 41683-41688, Sierra Leone, Malai, SL 03/11 (8°45’ N, 12°56’ W); ZFMK 41689-41697, Sierra Leone, Rowala, SL 03/12 (8°47’ N, 11°54’ W); ZFMK 41698-41704, Sierra Leone, Magben- ta, SL 03/19 (8°24’ N, 12°56’ W). Callopanchax toddi: ZFMK 41705-41715, Guinea, Mo- la, GM 08/4 (9°20’ N, 12°55’ W); ZFMK 41716-41723, Guinea, Takhori, GM 08/9 (10°09’ N, 14°01’ W); ZFMK 41724-41728, Guinea, Katamara, GM 08/10 (10°11? N, 14°03’ W); ZFMK 41729-41735, Guinea, Lamayah, GM 08/11 (10°09’ N, 13°44’ W); ZFMK 41736-41739, Sier- ra Leone, Sentai, SL 93/15 (9°04’ N, 12°53’ W). Callopanchax cf. toddi: ZFMK 41740-41744, Guinea, Gbeteyah Moribayah, GM 08/1 (9°36’ N, 13°20’ W); ZFMK 41745-41751, Guinea, Mangata Badjana, GM 08/3 (9°38’ N, 13°19’ W); ZFMK 41752-41755, Guinea, Waralneen GV 08/8 (0237. IN, 13.197 W)3 ZEMIK 41756-41757, Guinea, Kabak, GM 08/12 (9°21’ N, 13°23’ W); ZFMK 41758-41763, Guinea, Kabak, GM 08/13 (ONAN G13 237 W)): Bonn zoological Bulletin 57 (1): 3-14 Acknowledgements. The authors thank Kouroma Christine Sag- no, Direction National, Ministere du Development, and Bakary Keita, Ingenieur des Eaux et Forets, Chef de la Division Faune, Conakry, Guinea, for providing research permissions and sup- port, Samba, Ousmane and Seybou Sidibe, Conakry, Guinea, for help with organisation of the research trip and support in the field; Barbara Wiese, Wilhelm Kugelmann for support in the field, Fabian Herder (ZFMK, Bonn) for loan of specimens and access to the digital X-ray machine; Diethard Tautz for support and laboratory facilities, Floyd A. Reed and Anja C. Schunke (all MPI, Plén) and two anonymous reviewers for valuable com- ments and suggestions on earlier versions of the manuscript, and Gunther Fleck for the French translation of the abstract. REFERENCES Berkenkamp HO, Etzel V (1980) Description provisoire de Ro- loffia huwaldi spec. nov. 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Sinauer Associates, Sunderland, Massachusetts Tamura K, Dudley J, Nei M, Kumar S (2007) MEGA4: Molec- ular Evolutionary Genetics Analysis (MEGA) software ver- sion 4.0. Molecular Biology and Evolution 24: 1596-1599 Thompson JD, Gibson TJ, Plewniak F, Jeanmougin F, Higgins DG (1997) ClustalX windows interface: flexible strategies for multiple sequence alignment aided by quality analysis tools. Nucleic Acids Research 24: 4876-4882 Van den Nieuwenhuizen A (1974) Leuchtende Edelsteine vom Schwarzen Erdteil. Die Aquarien- und Terrarienzeitschrift (DATZ) 27: 46-50 Van der Zee JR (2002) Neue Einsichten in die Verwandtschaft der Gattung Episemion. Deutsche Killifisch Gemeinschaft Journal 34 : 135-143 Van der Zee JR, Wildekamp RH (1994) Description of a new Fundulopanchax species (Cyprinodontiformes: Aplocheilidae) from the Niger delta, with a redefinition of the genus Fundu- lopanchax. Journal of African Zoology 108: 417-434 Wildekamp RH (1996) A world of killies. Atlas of the oviparous cyprinodontiform fishes of the World. Volume 3, American Killifish Association Publications, Mishawaka, Indiana Wildekamp RH, Romand R, Scheel JJ (1986) Cyprinodontidae. Pp. 165-276 in: Daget J, Gosse J-P, Thys van den Audenaerde DFE (eds.) Check-list of the freshwater fishes of Africa CLOF- FA, Volume 2. ISNB Bruxelles, MRAC Tervuren, ORSTOM Paris Wildekamp RH, Van der Zee JR (2003) Cyprinodontiformes. Pp. 298-442 in: Paugy D, Lévéque C, Teugels GG (eds.) The Fresh and Brackish Water Fishes of West Africa. Volume II. IRD Editions, Paris Zhang D-X, Hewitt GM (1996) Nuclear integrations: challenges for mitochondrial DNA markers. Trends in Ecology and Evo- lution 11: 247-251 Received: 18.01.2010 Accepted: 05.02.2010 Corresponding editor: F. Herder ©ZFMK Bonn zoological Bulletin Volume 57 pp. 15-17 | Bonn, April 2010 Gekko ulikovskii Darevsky & Orlov, 1994: a junior synonym of Gekko badenii Szczerbak & Nekrasova, 1994 Quang Truong Nguyen!3, Andreas Schmitz? & Wolfgang B6hme?3 ! Institute of Ecology and Biological Resources, 18 Hoang Quoc Viet, Hanoi, Vietnam; E-mail: nqt2@yahoo.com 2 Muséum d’histoire naturelle, C.P. 6434, CH-1211 Geneva 6, Switzerland; E-mail: andreas.schmitz@ville-ge.ch 3 Zoologisches Forschungsmuseum Alexander Koenig, Adenauerallee 160, D-53113 Bonn, Germany; E-mail: w.boehme.zfmk@uni-bonn.de In Vietnam, a total of seven species of Gekko are current- ly recognized: G. badenii, G. chinensis, G. gecko, G. grossmanni, G. palmatus, G. scientiadventura, and G. ulikovskii (Rosler et al. 2004; Nguyen et al. 2009). Both Gekko badenii and G. ulikovskii were simultaneously de- scribed in 1994 based on the type series collected from Ba Den Mountain in Tay Ninh Province and “Gilai-Con Tum Province” (now Kon Tum Province) near the border with Laos and Cambodia, respectively (Darevsky & Orlov 1994; Szezerbak & Nekrasova 1994). The diagnostic char- acters of G. badenii and G. ulikovskii obtained from the original descriptions are presented in Table 1. In the key to the species of Gekko from Vietnam, Résler et al. (2004) distinguished these two species by the num- ber of interorbital scales (30—37 in badenii versus 40-46 in ulikovskii). Based on the original descriptions, G. badenii further differs from G. ulikovskii by the follow- ing characters: smaller SVL (51.5—76.5 mm in G. badenii vs. 87-108 mm in G. uwlikovskii), smaller ratio of SVL/TaL (0.82 in G. badenii vs. 0.94—-1.24 in G. ulikovskii), and larger number of precloacal pores (14-18 in G. badenii vs. 10-15 in G. ulikovskii). During recent field work in southern Vietnam, a series of G. badenii was collected in October 2004 by Nguyen Huu Dat from the type locality on Ba Den Mountain, Tay Ninh Province, and was subsequently deposited in the Institute of Ecology and Biological Resources, Hanoi, Vietnam (IEBR 1555, 1562-1564, 1566). Our own examination of this new col- lection compared with two paratypes of G. ulikovskii de- posited in the collection of the Zoologisches Forschungsmuseum Alexander Koenig, Bonn, Germany (ZFMK 55110-55111) revealed that there are no distinct differences in morphology between these two species. All Fig. 1. Gekko badenii (IEBR 1562) from Ba Den Moutain, Tay Ninh Province, Vietnam (alive animal) 16 Quang Truong Nguyen et al. Fig. 2. Gekko badenii (IEBR 1562) from Tay Ninh Province (a) and Gekko ulikovskii (Paratype: ZFMK 55110) from Kon Tum Province (b) (preserved specimens). diagnostic characters obtained from the original descrip- tions and examined specimens resemble each other or rather overlap: SVL (51.5—-106.0 in G. badenii and 87.0-108.0 in G. ulikovskii), ratio of SVL/TaL (0.81—1.28 in G. badenii and 0.94—1.24 in G. ulikovskii), interorbital scales (30-39 in G. badenii and 35—46 in G. ulikovskii), precloacal pores (12-18 in G. badenii and 10-15 in G. ulikovskii), dorsum dark grey with 6—8 light bands in both species, for further comparison see Table 1. It is noted that our scalation counts for two paratypes of G. ulikovskii (ZFMK 55110-55111) are a bit different from the origi- nal description: interorbitals (number of scales between middle orbitals, not including granular scales on upper eyelids) 35-38 versus 40-41 in the original description; scales around midbody (number of scales around body at the position between axilla and groin): 114-118 versus 131—136 in the original description. Moreover, the char- acter “scales around midbody” was erroneously noted as “dorsals around midbody—DoM” for 11 paratypes [Table 1, p. 74] by Darevsky & Orlov (1994). Although these au- thors did not mention their method to count the number of dorsals around midbody, we supposed that this is the number of scales between lateral folds at midbody. Our examination of ZFMK 55110-55111 showed that the Bonn zoological Bulletin 57 (1): 15-17 scales between lateral folds at midbody only range from 84-91 (131-136 scales in the original description). Therefore, this morphological character should be correct- ed as “scales around midbody” as it was mentioned for the holotype (132 scales around midbody). In terms of priority, the formal description of Gekko badenii was published in Vestnik Zoologii on 15 April 1994, whereas the description of G. ulikovskii was pub- lished in the journal Salamandra on 15 May 1994. There- fore, based on the evidence mentioned above and accord- ing to the International Commission on Zoological Nomenclature (ICZN, 1999: Articles 23.1 and 23.3) Gekko ulikovskii Darevsky & Orlov, 1994 must be regarded as a junior synonym of Gekko badenii Szczerbak & Nekraso- va, 1994. Acknowledgements. NQT acknowledges Nguyen Huu Dat (Ho Chi Minh City) and Nguyen Van Duc (IEBR) for providing spec- imens from Tay Ninh Province. We thank Ho Thu Cuc (IEBR) for translation of the article in Russian. We are grateful to L. Lee Grismer (La Sierra University) and Thomas Ziegler (Cologne Zoo) for their comments on the manuscript and providing the map. This research was partially funded by the German Acad- emic Exchange Service (DAAD). Vietnam Gulf of Tonkin Cambodia v Gulfof ° Thailand Fig. 3. Map showing the type localities of Gekko badenii (Tay Ninh Province) and Gekko ulikovskii (Kon Tum Province) in Vietnam. ©OZFMK Gekko ulikovskii Darevsky & Orlov, 1994: a junior synonym of Gekko badenii Szczerbak & Nekrasova, 1994 ia Table 1. Comparison of morphological characters of G. badenii Szczerbak & Nekrasova and G. ulikovskii Darevsky & Orlov ba- ce ee, Original description Specimens examined G. badenii G. ulikovskii G. badenii G. ulikovskii n=3 n= tt M=S n=2 Sex (M = male, F = female) (2 M, | F) (10 M, 1 F) (3 M, 2 F, (2 M, paratypes) additional material) SVL 51.5-76.5 87-108 87-106 102.1-102.7 (mean + SD) (62.3 + 0.7) (101.1 + 5.7) (93.3 + 7.6) (102.4 + 0.4) Tail length (TaL) 72.0-93.5 80-112 82.8-107.8 95.8-99.6 (mean + SD) (82.7 + 1.07) (94.1411.1,n=9) (96.9412.3,n=4) (97.7+42.7) SVL/TaL 0.82 0.94-1.24 0.81—1.28 1.03—1.06 Snout length > orbit diameter = yes yes yes Nostril in contact with rostral, yes yes yes yes supranasal, two postnasals, and first supralabial Suture on upper part of rostral present present present present Supralabials 12-15 11-14 12-14 11-14 Infralabials 13-14 9-13 11-12 12 Internasals 1-3 0-1 1 0-1 Interobitals 30-37 40-46 36-39 35-38 Gular scales 4-8 - 6-12 7-12 Number of gular scales in - 5 4-5 4-6 contact with infralabials Scales around midbody ~ 123-136 114-119 114-118 Number of dorsal tubercles 12 = 8-13 11-12 Ventral scales between axilla and groin — 51 52-56 52-54 Ventral scales between latero-ventral folds 29-35 — 31-34 32-34 Subdigital lamellae under fourth toe 18-20 15-20 16-19 18 Precloacal pores 14-18 10-15 12-14 12 Enlarged subcaudals - 56-83 60-68 59-63 Ground color of dorsum brownish grey dark grey dark grey dark grey (in preservative) Light dorsal bands 4-8 8 6-8 7-8 REFERENCES Darevsky IS, Orlov NL (1994) Eine bemerkenswerte neue, gross- wichsige Art der Gattung Gekko: Gekko ulikovskii sp. nov. aus Zentralvietnam. Salamandra 30: 71-75 International Comission on Zoological Nomenclature (ICZN) (1999) International Code of Zoological Nomenclature. Fourth Edition, Online version Nguyen VS, Ho TC, Nguyen QT (2009) Herpetofauna of Viet- nam. Edition Chimaira, Frankfurt am Main R6sler H, Ziegler T, Vu NT, Herrmann H-W, Bohme, W (2004) A new lizard of the genus Gekko Laurenti, 1768 (Squamata: Sauria: Gekkonidae) from the Phong Nha-Ke Bang National Bonn zoological Bulletin 57 (1): 15—17 Park, Quang Binh Province, Vietnam. Bonner zoologische Beitrage 53: 135-148 Szezerbak NN, Nekrasova OD (1994) A contribution to the knowledge of Gekko lizards of Southern Vietnam with descrip- tion of a new species (Reptilia, Gekkonidae). Vestnik Zoologii 1: 48-52 [in Russian] Received: 23.04.2009 Accepted: 30.04.2009 Corresponding editor: R. van den Elzen ©OZFMK 18 Book Review Finfstiick HJ, Ebert A, Weiss I (2010). Taschenlexikon der Vogel Deutschlands. 684 pp., Quelle & Meyer Verlag GmbH & Co., Wiebelsheim, ISBN 978-3-494-01471-5; 24,95 €. For many birdwatchers buying a field guide is the first step into field ornithology. After identifying their first birds, prospective birdwatchers might wish to learn more about the species they just recognized. Reaching this goal is the aim of the work at hand which tries to inform in a com- pact format suitable for carrying in a rucksack about all bird species occurring in Germany. This lexicon 1s basi- cally a summary of the ,,kompendium der Végel Mitteleu- ropas™. It contains short chapters on taxonomy, origin of German and scientific names, occurrence, status, habitat, and breeding and feeding biology of all species found in Germany. These are complemented by short descriptions of appearance, voice, body masses and peculiarities. Each species is presented with a distribution map and photo- graphs of almost always-good quality, completed by dia- grams on phenology and moult. Even species that are rarely photographed in field guides of birds from Germany like Black Lark, Siberian Thrush or Northern Parula are depicted in this book. Generally, the proximity to the “Kompendium” provides reliability and timeliness of in- formation. Nevertheless the lexicon is not error-free: Right in the beginning the explanatory distribution map is in- correct, as the light blue areas of migration periods and point occurrences are missing. The distribution area of the Green Willow Warbler has been displaced to Fennoscan- dia and Alaska, and superspecies are not always handled with care, e.g. for the Eastern black-eared Wheatear, Pied and Cyprus Pied Wheatear are missing as further mem- bers of the superspecies. Furthermore, many topics are left out that can be found yet in the “Kompendium”’, for example information on population changes, behaviour, moult and conservation measures. A general preface and introductions to the fam- ilies are nonexistent. Species are presented in alphabeti- cal order of the German names, so that possibly longer lasting thumbing through the book could be necessary if for example looking up different species of a superspecies or genus, particularly since there is no register available. Searching for scientific names is hence impossible. Read- ers should be familiar with current taxonomic findings and changes of names, as species like Cory’s Shearwater, Yel- low Wagtail and Black-eared Wheatear are provided with Bonn zoological Bulletin 57 (1): 18 Fiinfstiick/Ebert/Weifs 4 Taschenlexikon der Vogel Deutschlands Ein kompellier - Begleiter durch die ' heimische Vogelwelt a - ee. {i} “ >, RS eres ace QUELLE & MEYER | Qusieiiea eS new German names (“Sepiasturmtaucher”, ‘“Wiesen- schafstelze”’, “Mauren- and Balkansteinschmatzer’’). On the whole the lexicon is well suited for an entry into bird watching and can be recommended to everyone who is looking for a supplement to mere field guides. Wish- ing to seriously deal with field ornithology, though, re- quires purchasing additional books, such as the more cost- ly but also more detailed three volumes of “Kompendi- um der Vogel Mitteleuropas”’. Kathrin Schidelko Zoologisches Forschungsmuseum Alexander Koenig, Bonn ©ZFMK Bonn zoological Bulletin Volume 57 Issue | pp. 19-29 Bonn, April 2010 A new chameleon of the Trioceros bitaeniatus complex from Mt. Hanang, Tanzania, East Africa (Squamata: Chamaeleonidae) Patrick Krause! & Wolfgang Bohme! | Zoologisches Forschungsmuseum Alexander Koenig, Adenauerallee 160, D-53113 Bonn, Germany; E-mails: paddy.krause@web.de; w.boehme.zfmk(@uni-bonn.de Abstract. A new species of Zrioceros from the montane forests of Mount Hanang in Northwest Tanzania is described. The only known population of this species is found on Mt. Hanang in Tanzania around 2800 m a.s.l. This chameleon bears close morphological resemblance to Trioceros sternfeldi (Rand, 1963) from northern Tanzania. In order to infer phylogenetic relationships within the group, morphological and genetic data (mitochondrial 16S and 12S rRNA genes) were analysed. The new species could be compared with specimens of different populations of Trioceros sternfeldi (Mts. Meru, Kilimanjaro and Ngorongoro) and several other representatives of the family (J. hoehnelii, T. ellioti, T: rudis rud- is, T. bitaeniatus). Results of maximum parsimony, maximum likelihood and Bayesian analyses consistently placed Tri- oceros n. sp. from Mt. Hanang as a single, independent terminal clade with high genetic divergences towards all other included members of the group. Key words. Squamata, Chamaeleonidae, Trioceros, Africa, molecular phylogeny, biogeography INTRODUCTION Mt. Hanang represents one of the numerous ancient and isolated volcanic mountains which characterize the land- scape of eastern Africa. In northern Tanzania Mts. Kili- manjaro, Meru, and the Ngorongoro Crater are three of the better known extinct volcanoes which are frequently visited by tourists and lie within some of Tanzania’s most famous national parks. Several biologists identified these forested East African mountains as centers of endemism for chameleons of the “rudis group”, a subgroup of the Trioceros bitaeniatus species complex (Rand 1963; Pil- ley 2000; NeCas et al. 2003). The Trioceros bitaeniatus complex comprises various taxa of small, hornless, ovo- viviparous East African chameleons from the highland re- gions of eastern Africa, (Uganda, Kenya, Tanzania, south- ern Ethiopia and Sudan). All members of the group were placed into the separate subgenus Trioceros by Klaver & Boéhme (1986) which was recently elevated to full gener- ic rank by Tilbury & Tolley (2009). In contrast to other species of the bitaeniatus group, chameleons of the rud- is subgroup are rather stocky animals with short, broad heads and heterogeneous scalation. They belong to a se- ries isolated populations restricted to mountain forests ranging from the Ruwenzori in the north-west, south to the mountains east of Lake Malawi with outliers on Mt. Kilimanjaro, Mt. Meru and Ngorongoro in northern Tan- zania. The Tanzanian forms of this subgroup were initial- ly assigned to a rudissubspecies (Chamaeleo rudis stern- feldi) by Rand (1963), but this was long believed to be of rather specific than subspecific rank by several authors (e.g. NeCas et al. 2003). The independent taxonomic sta- tus of this form was subsequently demonstrated on the ground of morphological (Klaver & Bohme 1997; Krause 2006) and genetic data (Krause 2006). The speciation of the bitaeniatus group as a whole seems closely related to the formation of the East African rift val- leys when volcanism resulted in a separation of the an- cestral populations. This can therefore be regarded as a major factor that gave rise to present day species (Pilley 2000; Wasser & Lovett 1993). Three specimens of a curious chameleon from the Trio- ceros bitaeniatus- species complex were collected on Mt. Hanang at 2800 m altitude in March 2002, morphologi- cally similar to 7! sternfeldi (Rand, 1963) but with a very distinct colouration. This form is known only from the Mt. Hanang Catchment Forest Reserve (4° 25’ to 4° 35’ S and 35° 20’ to 35° 25’ E), situated about 4 km from the dis- trict capital Katesh and 70 km from the town Babati in the administrative region of Arusha in northern Tanzania (Figure 1). 20 Patrick Krause & Wolfgang BOhme Fig. 1. Map of Tanzania with distribution of the discussed species. 1=Mt. Hanang: 7. hanangensis n. sp.; 2=Ngorongoro: T. sternfeldi; 3=Mt. Meru: T. sternfeldi; 4= Mt. Kilimanjaro: 7. sternfeldi; 5=Usambara Mts.: 7. sternfeldi. MATERIALS AND METHODS All specimens were photographed in situ to record their natural coloration and habitus. Eversion of hemipenes in freshly dead specimens was done in the by injecting 70 % Alcohol into the organs (Glaw et al. 1999). The collect- ed voucher specimens were deposited in the Zoologisches Forschungsmuseum Alexander Koenig in Bonn (ZFMK). Additional material was examined from the Zoologische Staatssammlung Miinchen (ZSM). In order to test for morphological variation of the paratypes, chameleon body measurements were obtained with a digital caliper and evaluated with morphometric analyses methods (Krause 2006). Statistical analysis was performed using the GLM Multivariate procedure as im- plemented in SPSS 12 which provides analysis of vari- ance for multiple dependent variables by one or more fac- tor variables or covariates. In the present study the mor- phometric measurements (Krause 2006) excluding body length were chosen as dependent variables and the mor- phometric variable “snout-vent-length” was included as covariate to exclude the effect of body size on the other measured parameters. After an overall f-test has shown significance, post hoc tests (Scheffé) were applied to eval- uate differences among the specific means. Probability val- ues less than 0,05 were assumed to infer biological sig- nificance. Bonn zoological Bulletin 57 (1): 19-29 Molecular data could be obtained from tissue samples (muscle or liver, fresh or preserved in 98% ethanol) of the following taxa: Trioceros n. sp., (Mt. Hanang, Tanzania), Trioceros rudis (Uganda), Trioceros sternfeldi (Mt. Kili- manjaro, Mt. Meru, Tanzania). In order to analyse the phy- logenetic relationships within the bitaeniatus group the following taxa were also included in the analysis: 7. elli- oti, T. hoehnelii, T. bitaeniatus, T: narraoica, T. werneri, T. jacksonii, T. deremensis and T: johnstoni. Kinyongia uthmoelleri, Chamaeleo dilepis. Uromastyx acanthinura and Leiolepis belliana were used as outgroups. All spec- imens included in this study are listed in Tab. 1 and se- quences were submitted to GenBank (accession numbers AY927239-AY 927276). DNA was extracted using the Chelex method (Walsh et al. 1991). In this study a 5% Chelex solution (2,5 mg Chelating Resin [Iminodiacetic Acid; Sigma Aldrich] in 50ml ddH 0) was used (Schmitz A 2003). Two mitochondrial genes which encode for the 16S und 12S subunits of the ribosomes (16S and 12S rRNA genes) where chosen for molecular analysis. Primers 16SA (light chain 5’- CGC CTG TTT ATC AAA AAC AT -3’) and 16SB (heavy chain 5’- CCG GTC TGA ACT CAG ATC ACG T) of Palumbi et al. 1991 were used to amplify a section of the 16S rRNA gene. The primers 12SA-L (light chain; 5’ - AAA CTG GGA TTA GAT ACC CCA CTA T - 3’) and 12SB-H (heavy chain; 5’ - GAG GGT GAC GGG CGG TGT GT - 3’) of Kocher et al. (1989) were used to amplify a section of the 12S rRNA gene. PCR cycling procedure for both gene-fragments was as follows: Initial denaturation step: 90s at 94°C, 33 cy- cles: denaturation: 45s at 94°C, primer anealing for 45s at 55°C, extension for 90s at 72°C. Purification of the PCR products was conducted follow- ing standard methods using QIAquick PCR Purification Kit (Qiagen), sequencing reactions of the obtained ampli- fied DNA including cycle sequencing (standard proce- dures) were conducted by MWG-Biotech and Max Planck Institut fiir Zichtungsforschung (MPI) in Cologne using ABI Prism DNA automatic sequencer. Sequences were aligned using the computer program ClustalX (default parameters, Thompson et a. 1997). The alignment was subsequently adjusted manually using the computer program BioEdit (Hall 1999). 33 sequences (combined 16S and 12S rRNA genes) could be obtained, each comprising 861 bp, referring to the aligned sequences including gaps (16S: 479 bp; 12S: 382 bp). The alignment gaps were treated as missing charac- ter. The entire 16S+12S alignment was included in the analyses. As with two specimens (ZFMK 63223, 70752) only 12S sequences could be obtained, we conducted a separate analysis using the 12S dataset only. ©ZFMK A new chameleon of the Zrioceros bitaeniatus complex from Mt. Hanang, Tanzania 21 Fig. 2. Sulcal (left) and asulcal (right) view of the hemipenis of Trioceros hanangensis n. sp. from Mt. Hanang, Tanzania. Data were analysed using maximum likelihood and par- simony optimality criteria in PAUP 4.0b10 (Swofford 2002), further we used Bayesian inference as implement- ed in MrBayes 3.1.1 (Ronquist & Huelsenbeck 2003). For maximum likelihood, a heuristic search was conducted with 10 replicates of random stepwise addition and tree bisection-reconnection (TBR) branch swapping. Relative branch support in phylogenetic analysis was evaluated with 100 bootstrap pseudoreplicates. Parsimony analyses used heuristic searches with 100 random additions of taxa and tree bisection-recombination branch swapping. When more than a single most parsimonious tree was found, a strict consensus tree was generated. Confidence in the nodes was assessed by 2000 bootstrap pseudoreplicates (Felsenstein 1985) with random addition of taxa. Only bootstrap supports of 70 % and higher were considered as reliable, as such values were found to indicate 95 % probability of correct topology (Hillis & Bull 1993). For Bayesian analyses a GTR+I+G model was chosen accord- ing to the optimal model and parameter-values based on the AIC criteria determined in MrModeltest 2.2 (Nylan- der 2004). Two MCMC analyses for 10° generations each were conducted. The topologies of the sampled trees were used to generate a strict consensus tree. Only clades with a posterior probability of 95% or greater where consid- ered to be significantly supported. RESULTS Trioceros hanangensis n. sp. Holotype. ZFMK 82369, adult male with everted hemipe- nis. Locality: Tanzania: Mt. Hanang Forest Reserve near the village Jorodom (4° 25’ to 4° 35’ S and 35° 20’ to 35° 25’ E) at 2800 m, collected by P. Krause, March 2002. Bonn zoological Bulletin 57 (1): 19-29 Paratypes. ZFMK 82368, subadult male, same data as the holotype; ZSM 111/1981/1- ZSM 111/1981/20 collected by L. Uthmoeller (1937) from the same locality as the holotype. Diagnosis. A medium sized, stocky chameleon resembling a typical member of the Trioceros bitaeniatus complex (Rand 1963). The maximum total length is 138 mm; the tail comprises an average of 45 %, males with longer tails. Adult males and females have a similar snout-vent-length (SVL) the females being slightly larger. Body scalation is heterogeneous, with strongly enlarged denticulate scales in two distinct rows on the sides of the body. The gular crest is weak to medium, consisting of scales 1.5 times the length of their maximum width. A ventral crest is clear- ly visible in bigger specimens to almost indistinct in small- er ones. No rostral appendix is visible like in other mem- bers of the bitaeniatus complex (Trioceros narraioca Ne€as, Modry & Slapeta, 2003; Trioceros hoehnelii Stein- dachner, 1891). This chameleon bears a close morphological resemblance to Trioceros sternfeldi from northern Tanzania (Mts. Kil- imanjaro, Meru and Ngorongoro Crater) but shows a dis- tinct colour pattern, which differs remarkably from 7. sternfeldi (Sternfeld 1912a, Figures 3, 5, 6). Further, it pos- sesses scattered melanophores on the sulcus of the hemipenis which are unique as compared to 7. sternfeldi. Despite a close genetic relationship with T. e//ioti, the new form can clearly be distinguished by absence of the dis- tinct longitudinal groves on the gular pouch, a more het- erogeneous scalation and a stouter body as compared with T. ellioti, which is rather slim (Gtinther 1895; Spawls et al. 2002). The only known population of this species is found on Mt. Hanang above 2800 m in montane bushland. Description of holotype. Head short and broad, distance from tip of snout to end of parietal crest less than twice greatest width between the temporal ridges. Parietal crest not swollen posteriorly, being almost straight in profile. No rostral projection at the point of fusion of canthi ros- trales. Occipital flaps absent. Ventral crest moderately de- veloped, composed of cone-shaped, subequal scales, longest on throat, about 3/4 of eye diameter. Throat ho- mogenously covered with small scales, no indication of longitudinal grooves. Head length (from tip of snout to posterior margin of casque) 21 mm. Height of casque (from angle of mouth to top of casque) 16 mm, slightly less than length of mandible (20 mm). Caudal side of casque homogenously covered with fine granular scales. Parietal crest consists of 15 enlarged elongate scales. Two other diagonal (paraparietal) crests visible on each side of ©OZFMK 22 Patrick Krause & Wolfgang Bohme Fig. 3. Holotype of Trioceros hanangensis n. sp. (male, ZFMK 82369) from Mt. Hanang, in life. head, consisting of 4 enlarged tubercular scales. These originate in the immediate vicinity of the parietal crest and run rostrolaterally towards the orbit. Lateral crests com- posed of 4 enlarged, prominent pointed scales, extending to the supraorbital crest (11 pointed scales) and to canthi rostrales (5 scales). Body rather stout. Dorsal crest well developed, composed of scales in groups of three anteriorly and four posterior- ly. Increasing posteriorly in size within the group, giving the back a serrate appearance. Dorsal crest extends to the tail and becomes indistinct within its terminal third. The Fig. 4. Bonn zoological Bulletin 57 (1): 19-29 Trioceros hanangensis n. sp. (subadult male, ZFMK 82368) from Mt. Hanang. first scales of the series of three just behind the head are longest, reaching 1.7—2 mm length. Ventral crest moder- ate to weak, best developed on the throat; posteriorly scales are shorter but still visible up to posterior belly. Scalation heterogeneous. Sides of body, tail and upper sur- faces of limbs covered with irregularly sized, convex, small scales interspersed with larger convex scales. Lar- gest of these form a narrow row running from neck just behind temporal crests to tail. A second row of slightly smaller scales runs from shoulder to hind leg. This row is much less distinct and more irregular. Vent, inner side Trioceros sternfeldi (male) Ngorongoro Crater. ©ZFMK A new chameleon of the 7rioceros bitaeniatus complex from Mt. Hanang, Tanzania Fig. 6. Trioceros sternfeldi (male) Mt. Kilimanjaro. of legs, soles and underside of the tail almost homogene- ously scaled. No tarsal spurs. Hemipenis. Smooth margined calyces cover the asulcal side of the hemipenis, extending as far as the sulcal lips. Apical tip characterised by two pairs of finely serrate ro- tulae of roughly semicircular shape, positioned towards the sulcal side and extending around lateral side of hemipenis. Scattered melanophores could be observed in the sulcus (Figure 2). Colouration and pattern. Vivid green with lateral yel- low bar following the row of enlarged scales running from neck to the base of tail. Head light blue except gular re- gion. Eyeballs bright blue to turquoise. Variation of the paratypes. Analysis of morphometric variation between male and female specimens of Trioceros n. sp. from Mt. Hanang resulted in males having signifi- cantly broader heads than females ( x = 13,09 mm vs. x = 12,59 mm; p = 0,030) and a tendency to increased ru- gosity in adult males. There is no apparent statistical dif- ference in snout-vent-length (SVL) comparing both sex- Fig. 7. Mt. Hanang from Katesh, SSW side of the mountain. Bonn zoological Bulletin 57 (1): 19-29 N Ww es. (males: SVL = 72,30 mm, N = 9 / females: SVL = 75,52 mm, N = 14). No significant differences in relative tail length (TL) between sexes could be observed (males: TL = 66,22 mm / females: TL = 62,71 mm). Distribution. Only known from Mt. Hanang Catchment Forest Reserve (4° 25’ to 4° 35’ S and 35° 20’ to 35° 25’ E) situated about 4 km from the district capital Katesh and 70 km from the town Babati in the administrative region of Arusha. The new species is named after the type-local- ity Mount Hanang. Ecological remarks and habitat preference.: 7) hanan- gensis n. sp. was found in upland moorland at 2800 m on the moister north-eastern side of Mt. Hanang. Despite a prevaling semi-arid climate with 750-1500 mm annual rainfall, a pronounced mist effect on the eastern and north- eastern slopes is leading to a maximum of 2000 mm pre- cipitation per year at the higher regions of the mountain. The sampled specimens where observed during daytime in bushy highland vegetation (with Hypericum sp., Myr- sine africana, Protea sp.) near the source of the river Him- it, roosting on twigs at a height ranging from 100—150 cm from the ground. Phylogenetic affinities. In order to infer the phylogenet- ic relationships between the different members of the bitaeniatus group on the basis of molecular data maximum parsimony (MP), maximum likelihood (ML) and Bayesian analyses (PP) were conducted. Maximum likelihood analysis was conducted with a reduced dataset, due to computational constraints. Sequences obtained from the combined 16S+12S rRNA mitochondrial genes were analysed together with gene-fragments of Trioceros sternfeldi, T. rudis, T. ellioti, T: hoehnelii, T: bitaeniatus, T. narraoica, T. werneri, T. jacksonii, T: deremensis and T: johnstoni. The two included specimens of 7? hanangen- sis. sp. were found in a well supported clade in the MP and Bayesian trees (MP: 99; PP: 1.00), separating them Fig. 8. Type locality of Trioceros at 2800 m altitude. ©OZFMK 24 100 1.00, 96 0.99 51 69 1.00 87 al 0.99 97 0.82 Fig. 9. Patrick Krause & Wolfgang B6hme 98 1.00) 67 72 0.86 0.99 99 1.00 100 1.00 100 1.00. Kinyongia uthmoelleri Mt. Hanang, Tanzania Chamaeleo dilepis Mt. Hanang, Tanzania (V) Trioceros jacksonii Nairobi, Kenya Trioceros narraoica Mt. Kulal, Kenya Trioceros kinetensis Sudan Trioceros ellioti Kakamega Forest, Kenya (I) Trioceros ellioti Kakamega Forest, Kenya (Il) Trioceros ellioti Nandi Hills, Kenya Trioceros ellioti Kenya - Trioceros ellioti Ruwenzori Mts., Uganda (I) Trioceros ellioti Ruwenzori Mts., Uganda (Il) Trioceros hanangensis n. sp. Mt. Hanang, Tanzania (I) Trioceros rudis Ruhiza, Uganda Trioceros rudis Bwindi Forest, Uganda Trioceros sternfeldi Mt. Kilimanjaro, Tanzania - Trioceros sternfeldi Mt. Meru, Tanzania (l) Trioceros sternfeldi Mt. Meru, Tanzania (II) Trioceros hoehnelii Eldama Ravine, Kenya Trioceros hoehnelii Uganda Trioceros hoehnelii Uganda Trioceros hoehnelii Mt. Kenya (I) ~ Trioceros hoehnelii Mt. Kenya (Il) Trioceros bitaeniatus Arusha, Tanzania Trioceros bitaeniatus Mt. Hanang, Tanzania (I) Trioceros bitaeniatus Mt. Hanang, Tanzania (Il) Trioceros bitaeniatus Mt. Hanang, Tanzania (III) Trioceros bitaeniatus Mt. Hanang, Tanzania (IV) Trioceros werneri Udzungwa Mts., Tanzania Trioceros deremensis Usambara Mts., Tanzania Trioceros johnstoni Bwindi Forest, Uganda — Leiolepis belliana Uromastyx aegyptia - Trioceros hanangensis n.sp. Mt. Hanang, Tanzania (Il) Strict consensus parsimony tree derived from the combined fragment of the mitochondrial 12S and 16S rRNA genes. Parsimony bootstrap values are given above the nodes (bold), lower values are posterior probabilities produced by Bayesian infe- rence. Nodes with less than 50 % bootstrap support are not labelled. Bonn zoological Bulletin 57 (1): 19-29 ©ZFMK A new chameleon of the Zrioceros bitaeniatus complex from Mt. Hanang, Tanzania 25 71 100 71 100 78 | 74 55 97 93 51 59 Uromastyx aegyptia Chamaeleo dilepis Mt. Hanang, Tanzania (V) Trioceros jacksonii Nairobi, Kenya Trioceros narraoica Mt. Kulal, Kenya Trioceros bitaeniatus Mt. Hanang, Tanzania (IV) Trioceros rudis Bwindi Forest, Uganda Trioceros sternfeldi Mt. Kilimanjaro, Tanzania Trioceros sternfeldi Mt. Meru, Tanzania (Il) Trioceros hanangensis n.sp. Mt. Hanang, Tanzania (Il) Trioceros ellioti Kakamega Forest, Kenya (I) ~ Trioceros ellioti Nandi Hills, Kenya Trioceros ellioti Ruwenzori Mts., Uganda (Il) Trioceros hoehnelii Mt. Kenya (Il) - Trioceros hoehnelii Eldama Ravine, Kenya Trioceros werneri Udzungwa Mts., Tanzania Trioceros deremensis Usambara Mts., Tanzania - Trioceros johnstoni Bwindi Forest, Uganda Fig. 10. ML tree derived from the combined fragment of the mitochondrial 12S and 16S rRNA genes. Bootstrap values are gi- ven above the nodes, nodes with less than 50 % bootstrap support are not labelled. Bonn zoological Bulletin 57 (1): 19-29 ©ZFMK 26 Patrick Krause & Wolfgang Bohme 0.56 68 98. ee 92 97 | 0.95 64 74 0.99 93 94 0.84 | 0.99 59 59 0.69 62 0.98 1.00 0.97 | 7471 1.00 Uromastyx aegyptia Chamaeleo dilepis Mt. Hanang, Tanzania (V) Trioceros jacksonii Nairobi, Kenya Trioceros narraoica Mt. Kulal, Kenya Trioceros bitaeniatus Arusha, Tanzania Trioceros bitaeniatus Maralal, Kenya Trioceros bitaeniatus Mt. Hanang, Tanzania (I) Trioceros bitaeniatus Mt. Hanang, Tanzania (Il) Trioceros bitaeniatus Mt. Hanang, Tanzania (III) Trioceros bitaeniatus Mt. Hanang, Tanzania (IV) Trioceros rudis Ruhiza, Uganda Trioceros hoehnelii Uganda Trioceros hoehnelii Eldama Ravine, Kenya Trioceros hoehnelii Mt. Kenya (I) Trioceros hoehnelii Mt. Kenya (Il) Trioceros sternfeldi Mt. Kilimanjaro, Tanzania Trioceros sternfeldi Mt. Meru, Tanzania (I) Trioceros hanangensis n.sp. Mt. Hanang, Tanzania (i) Trioceros ellioti Kakamega Forest, Kenya (I) Trioceros ellioti Nandi Hills, Kenya Trioceros cf. bitaeniatus Sudan Trioceros rudis Ruwenzori Mts., Uganda Fig. 11. Consensus tree derived from Bayesian inference using the mitochondrial 12S gene. Posterior probabilities are given abo- ve, ML bootstrap values (bold) and MP bootstrap values (inverse) below. Nodes with less than 50 % bootstrap support are not la- belled. Bonn zoological Bulletin 57 (1): 19-29 ©OZFMK Table 1. Specimens included in this study. Species Kinyongis uthm celier’ Cham seleo dilepis Trioceres ellioti Trioc eros ellioti Troceros ellioti Trocercs ellioti Trioceros ellioti Trioceros ellioti Tricc eros hanangensis 5p. nov Triocercs hanangensis sp. nov Trioceros rucis - Thoceros rudis Trioc eros rucis Trice eros kinetensis Trioceros sternfeldi Trice eros sternfeldi Triocercs sternfeldi Trocercs hoehnelii Trioceros hoehnelit Trioceros hoehneli Trioceros hoehnelit Trioceros hoehnelii Trocercs bitaeniatus Trocercs biaenistus Trocercs bitsernatus Troceros bitaematus Tnocercs bitzeniatus Trocerce biaeniatus Trioc eros nairaoics Trioc eros jacksonii Triocercs wemeri Troceros derem ensis Trioc eros johnstoni Leiolepic bellians Uram astyx segyptia Locality Nt. Hanang, Tanzania Mt. Hanang, Tanzania (V) Kakamegs Forest, Kenya [I) Kakamega Forest, Kenya (II) Nandi Hills, Kenya Kenys Ruwenzori Mts., Uganda (1) Ruwenzori Nits., Uganda (II) It. Hanang, Tanzania (1) Nit. Hanang, Tanzania (II) Ruhiza, Uganda Bwindi Forest, Ugands Uganda S$ Sudan Mt. Kilimanjaro, Tanzania Nit. Meru, Tanzania (I) Mit. Meru, Tanzania {II} Mt. Kenya (1) Ugands It. Kenya (Il) Eldama Ravine, Kenya Ugsnda Wit. Hanang, Tanzania (1) Wt. Hanang, Tanzanis {II} Mit. Hanang, Tanzania (III) Wit. Hanang, Tanzania (IV) Arusha, Tanzania Maralal, Kenya Mt. Kulsl, Kenya Nairobi, Kenya Udzungwa IMits., Tanzania East Usambara Mits., Tanzania Bwindi Forest, Uganda Egypt Thailand Collection number A new chameleon of the Trioceros bitaeniatus complex from Mt. Hanang, Tanzania Accession number 165/125 ZFMK 82191 DOQ297231 DQz297245 ZFMK 82381 AY927263, AN 927244 ZFMK 82058 DOQ397278 DQ397258 ZFMK 20835 DO397279 DOQ297259 ZFNIK 82057 DO3S7225 DQ397229 ZFMK 70818 DO397280 DOQz297260 ZFMK 638287 DQ397281 DQ397256 ZFMK 63227 DOQ397282 DQ397257 ZFMK 82368 DQ397283 DOQ397254 ZFNIK 82369 DOQ397284 DQ397255 CAS 201781 DOQ397223 DOQ397223 CAS 201711 DOQ397285 DQ397252 ZFMKE3223 - DOQ397277 ZFMK 29714 DOQ397286 DQ397253 ZFMK 68629 DO397287 DOQ397261 TO68 (prov. ID) DO397288 DOQz97263 ZFMK 82250 DO397289 DO397262 ZFMK 70678 DO297290 DOQ397265 voucher not collected DOQ297291 DOQ297266 ZFMK 66578 DO397292 DQ397264 ZFMK 82088 DOQ397224 DQ397238 voucher net collected DQ397293 DQ397267 ZFIMIK 74949 DOQ3297222 DOQ397237 ZFMK 74952 DOQ397294 DO397265 ZFMK 82365 DOQ397295 DQ397270 ZFMK 82266 DO397296 DO397271 ZFNIK 82251 DO397297 DOs9g7272 ZFMK 70752 - DOQ397276 ZFMK 73958 DOQ397298 DO397268 CAS 199070 DO397226 DOQ397240 ZFMK 70452 DOQ397299 DOQ397274 CAS 168888 DOQ397200 DOQ297273 CAS 201597 DOQ397301 DOL97275 GenBank ABN3 1986 ABN3 1969 GenBank AB03 1994 AB03 1977 from all other included taxa. In the tree resulting from the maximum likelihood analysis, the single included speci- men of 7. hanangensis n. sp. clustered together with T. ellioti (ML: 74). Genetic distances to all other included taxa of the bitaeniatus group were relatively high (2,2—4 %) and were within the range which could be found be- tween well defined species. This confirms the independ- ent specific status of Trioceros hanangensis n. sp. Phylogenetic inference of the combined 16S+12S rRNA dataset revealed a sistergroup relationship with 7° e/lioti, which was weakly to moderately supported with all meth- ods (MP: 67; PP: 0.86; ML: 74). A genetic distance of 2,05-2,18 % could be observed between Trioceros hanan- gensis n. sp and the included specimens of 7: ellioti. Us- ing only the 12S fragment for phylogenetic inference, a much higher support for this sistergroup relationship could be observed (MP: 89; PP: 0.99; ML: 81). Nevertheless, the relationships of 7: hanangensis n. sp. towards the oth- er taxa of the bitaeniatus group, especially towards T. sternfeldi could not be finer resolved with the available limited genetic dataset, resulting in a large polytomy in all trees. Bonn zoological Bulletin 57 (1): 19-29 Conservation status. The forested area on Mt. Hanang is classified as a Catchment Forest Reserve covering ap- prox. 40 km? which is relatively small compared to oth- er reserves in Tanzania. The genuine forest of Mt. Hanang has been drawn back to the very base of the mountain and is under increasing threat by illegal logging, fires and drought. Due to the small size of the reserve, even minor disturbances are likely to affect the habitat of forest de- pendant species like 7. hanangensis n. sp. Along with the well-known mountains in northern Tanzania like Meru, Kilimanjaro, Ngorongoro and the Eastern Arc, Mt. Hanang is a further example of a remnant mountain for- est with quite unique climatic and ecological conditions. For this reason the protection and management of the re- maining forests on Mt. Hanang should receive urgent pri- ority. DISCUSSION Trioceros hanangensis n. sp. is a member of a species complex which is closely associated with the forest cov- ering many of the East African mountains. These ©OZFMK 28 Patrick Krause & Wolfgang Bohme chameleons represent a subgroup of the bitaeniatus-group (the “rudis group”’sensu Rand 1963). In contrast to other species within the bitaeniatus group, populations of the rudis group are restricted to mountain forests with wide- ly disjunct distribution, each population bearing specific morphological peculiarities distinguishing it from the oth- ers as first observed by Rand (1963). Like the bitaeniatus group as a whole, the different forms of the rudis group can most easily be understood as relics of a period equating with the last pluvial period of the Pleistocene when the rainfall in East Africa was higher and the forest was much more extensive than today. Drier pe- riods following the pluvial maxima resulted in a retreat of the forests from the lower elevations to their present positions on the higher mountains. As a consequence the chameleon populations living in these forests (Wasser & Lovett 1993) were also fragmented. Different selection pressures formed the various taxa (7. schubotzi, T. stern- feldi or T. hanangensis n. sp.) which are today restricted to relatively small areas or certain types of habitat. In con- trast to the mountains of the Eastern Arc, most volcanoes in northern Tanzania are relatively young and result from volcanic activity persisting from the Oligocene (37 myr ago) to the Quaternary. As a consequence, the origin of the different rudis-forms cannot lie very far in the past which makes it a rewarding subject for the investigation of beginning speciation processes. Despite a morphological similarity with Trioceros stern- feldi (Mts. Meru, Ngorongoro and Kilimanjaro) 7) hanan- gensis n. sp. can be regarded as valid species, chief indi- cators being a distinct colour-pattern, presence of scattered melanophores in the sulcus of the hemipenis (not present in 7: sternfeldi) and the high degree of genetic differen- tiation towards 7. sternfeldi and the other members of the group. Molecular analysis of the combined 16S+12S dataset shows a genetic differentiation between 2,2-4 %, which is clearly above the intraspecific level and confirms the independent specific status of 7) hanangensis n. sp. A weakly supported sister-group relationship towards T. ellioti is indicated by all the given trees (MP: 67; PP: 0.86; ML: 74). In contrast to T e/lioti T7 hanangensis n. sp. does not possess the distinct longitudinal groves on the gular pouch as present in 7: e//ioti, it has a more heterogeneous scalation and a stouter body than 7: ellioti (Giinther 1895; Spawls et al. 2002). Further, 7) e//ioti has a wide distri- bution ranging from western Kenya, the northern shore of Lake Victoria in Uganda, western Uganda, Rwanda and Burundi and occurs in open bushland and grassland. Limited by the available genetic dataset, phylogenetic re- lationships towards the other members of the bitaeniatus Bonn zoological Bulletin 57 (1): 19-29 group could not be sufficiently resolved . Phylogenetic affinities of 7) hanangensis n. sp. and T. sternfeldi can on- ly be assumed by means of morphological data: specimens of 7. sternfeldi from Mt. Kilimanjaro have dark eyeballs, in contrast to 7. sternfeldi from Ngorongoro and 7! hanan- gensis n. sp. which both have light blue eyeballs and are more alike in general colouration. The parietal knob ob- served by Rand (1963) associated with specimens from Mt. Meru and Mt. Kilimanjaro could not be observed in specimens from Mt. Hanang respectively Ngorongoro. These observations support the assumption that 7’ hanan- gensis n. sp. 1s more closely related to 7. sternfeldi from Ngorongoro than with the sternfeldi-populations from Meru and Kilimanjaro. The geography of the region would have allowed geneflow between the populations of Hanang and Ngorongoro in the past: particularly to the north-west of Mt. Hanang a continous forest could have existed during moister periods linking the habitat of 7 hanangensis n. sp. with the Nou- and Marang forests and further north with Oldeani (3188 m) and Ngorongoro (3000 m). Today, a dry and hot valley separates the moun- tain from the rift-wall and forms an impenetrable barrier for forest dependent species. A similar observation could be made with Kinyongia uth- moelleri, an endemic of the highlands of Hanang and Ngorongoro: the two populations show closest morpho- logical and genetic relationships (Krause 2006) which as well supports the above presumed close relationship be- tween 7. hanangensis n. sp. and T. sternfeldi from Ngorongoro. Acknowledgements. The work of the senior author (PK) was supported by a grant of the German Foreign Exchange Service (DAAD). Herpetological researches in Tanzania were authorized by Tanzania Commission for Science and Technology (COSTECH). Many thanks to Mr. Joas Kahembe, Babati, M. Nikundiwe and K. Howell (University of Dar es Salaam) who gave support with a lot of practical advice and assisted obtain- ing research permits. REFERENCES Felsenstein J (1985) Confidence limits on phylogenies: an ap- proach using the bootstrap. Evolution 39: 783-791 Glaw F, Vences M, Ziegler T, Bohme W, Kohler J (1999) Spe- cific distinctness and biogeography of the dwarf chameleon; Brookesia minima, B. peyrierasi and B. tuberculata (Reptil- ia: Chamaeleonidae): evidence from hemipenial and external morphology. Journal of Zoology 247: 225—238 Gunther A (1895) Notice of Reptiles and Batrachians collected in the eastern half of tropical Africa. Annals and Magazine of Natural History (6) 15: 523-529 ©ZFMK A new chameleon of the Zrioceros bitaeniatus complex from Mt. Hanang, Tanzania 29 Hall TA (1999) BioEdit: a user-friendly biological sequence alignment editor and analysis program for Windows 95/98/NT. 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Wasser (eds.), Cambridge University Press, Cambridge Received: 04.12.2009 Accepted: 10.02.2010 Corresponding editor: F. Herder ©OZFMK 1 00 1 y oe Bonn zoological Bulletin Volume 57 Issue | pp. 31-54 Bonn, April 2010 Reptiles from southern Benin, West Africa, with the description of a new Hemidactylus (Gekkonidae), and a country-wide checklist Klaus Ullenbruch!, Olaf Grell? & Wolfgang Bohme! 1Zoologisches Forschungsmuseum Alexander Koenig, Adenauerallee 160, D-53113 Bonn, Germany; E-mails: Klaus-Ullenbruch@t-online.de; w.boehme.zfmk(@uni-bonn.de; 2Biologenbtro GGV, Stralsunder Weg 16, D-24161 Altenholz-Stift, Germany; E-mail: olaf.grell@t-online.de Abstract. We report on a collection of reptiles made in southern Benin, mostly in the vicinity of Lama Forest, a relict rain forest surrounded by savannah habitats within the the so-called Dahomey gap. 48 species (3 chelonian, 20 saurian and 25 ophidian species were encountered) are listed with the respective voucher material and commented in respect to taxonomic or ecological information. Five lizard species (Agama sylvanus, Hemidactylus ansorgei, H. lamaensis n. sp., Holaspis guentheri, Varanus ornatus) and two snake species (Natriciteres variegata, Amblyodipsas unicolur) are new for the fauna of Benin, the second Hemidactylus species being even new to science. Some other species are reported from Benin for only the second time. This commented list of species collected from southern Benin is followed by a general country-wide checklist, which is based on literature data and also some unpublished records from northern Benin, among them the remarkable rediscov- ery of Agama gracilimembris, 9 decades after its original description from this country. Key words. Reptilia: Chelonia, Crocodylia, Squamata; Benin, West Africa, new country records; Hemidactylus lamaen- sis N. sp.; country-wide checklist INTRODUCTION Herpetologically, Benin is among the most insufficiently known countries in Africa. Bordered by Togo in the west, Burkina Faso and Niger in the north, and Nigeria in the east, Benin is best-known by its old name Dahomey, be- cause of the denomination of the huge savannah gap that separates the Upper Guinean rain forests from those of the Lower Guinean-Congolian forest block as Dahomey Gap. Among the first authors who dealt especially with Benin reptile material within the framwework of West African herpetology is Chabanaud (1916, 1917). During his cu- ratorship in the Museum of Natural History in Paris he published on acquisitions from this former part of French West Africa by the following collectors: Dr. G. Bouet, Healt Inspector of French West Africa (“Dahomey”, 1910-1913), Dr. Brot (“Haut-Dahomey”, 1908), Mr. A. Chevalier (“Dahomey”, 1909-1910), Mr. de Gironcourt (“Haut-Dahomey”, 1910), Mr. Gruvel (“Dahomey”, 1913), Mr. Primot (“Dahomey”, 1914), and Mr. Waterlot (“Dahomey”, 1914). The next important step in the explo- ration of this country was the mission to Togo and Dahomey carried out by A. Villiers in 1950 (see Villiers 1951, Loveridge 1952) which covered all important biotopes of these two countries. In 1999, Sinsin & Bergmans (1999) edited a book on rodents and snakes in Benin under mostly agricultural aspects. The table on the West African snake fauna including Benin provided for this book by Meirte is not considered here because it marks many species in this table as occurring in Benin without any reference to the source on which the information is based; on the other hand, several species are marked as questionably or not at all occurring in Benin (e.g. Scaphio- phis albopunctatus, Telescopus variegatus, Toxicodryas blandingii, Psammophis cf. sibilans, P. phillipsi: Meirte 1999) although they have been documented by museum voucher specimens in the earlier works cited above (see also the country checklist at the end of this paper). Where the savannah reaches the southern coast of West Africa the Dahomey gap stretches from roughly between Accra (Ghana) to Lagos (Nigeria), thus making Benin a seemingly pure savannah country. However, some forest 32 Klaus Ullenbruch et al. remnants such as Forét de Dan, Forét de Lama and Forét de Lokoli are still present in the southern part of Benin. The senior author of this paper (KU) had the opportuni- ty to work on a herpetological survey of the most impor- tant of these relict forest remnants, viz. Forét de Lama. This work took place within the framework of a project (“Biolama’’) of the University of Basel, Switzerland, guid- ed by Prof. Dr. Peter Nagel (see Lachat et al. 2006). The herpetological voucher material of this survey, which 1n- cluded also specimens obtained from neighbouring for- est and savannah areas (Ullenbruch 2003) is deposited in the Zoologisches Forschungsmuseum Alexander Koenig (ZFMK), Bonn, Germany. The amphibian part of this col- lection contained some important first country records of rain forest species (e.g. Ptychadena aequiplicata) and has already been included in a recent paper by Rédel et al. (2007). Amphibians collected by O. Grell in the Pendjari National Park, northern Benin (Grell 2003) have been in- corporated in the study by Nago et al. (2006). The reptil- ian records of this last-mentioned survey, mostly docu- mented by photographs, are included here only in the country-wide checklist at the end of this paper. The fau- nistic and ecological information on the reptiles of south- ern Benin (mission K. Ullenbruch) is presented here in a systematically arranged, commented species list. This list further contains some voucher material collected and do- nated to ZFMK by Dr. Werner Schréder, Gottingen, in 1984. The commented regional species list which contains also the description of one new species, 1s supplemented by a general, country-wide checklist that tries to summarize all literature data for Benin as a whole. REPTILES FROM LAMA FOREST, SOUTHERN BENIN, AND NEIGHBOURING AREAS Lama Forest or Forét de Lama is situated within a depres- sion bearing the same name, 80 km north of Benin’s cap- ital Cotonou (Figure 1). It is bordered in the north by the plateau of Abomey and in the south by the plateau of Al- lada. Lama Forest comprises a total area of 16.250 ha of “forét classée”, 9.750 ha of which belong to the Départe- ment Atlantique (Sous-préfecture Toffo) and 6.500 ha be- longing to the Départment Zou (Sous-préfecture Zog- bodomé) (Emrich et al. 1999). The center of this protect- ed area — the protectional status exists since 1947 — is formed by the Noyeau Central (4.777 ha) which is com- posed of primary forest (1.937 ha), degraded forest (1.388 ha) and of teak plantations and fallow grounds (together 1.452 ha) (Figure 2). The Noyeau Central is surrounded by ca 9.000 ha of forestry plantations. In earlier times, the primary forest covered the whole Lama depression which stretches over the south Benin in a west-east direction. Un- Bonn zoological Bulletin 57 (1): 31-54 Fig. 1. lenbruch View on Forét de Lama, South Benin. Photo: K. Ul- til 1946, large areas of natural forest were already de- stroyed, and only 11.000 ha remained. Since the 1960ies reforestations are taking place which favoured, however, mainly teak plantations. The natural vegetation has been protected only from 1988 onwards (Emrich et al. 1999). Mean annual precipitation is 1100 mm with two peaks (big and small rainy seasons) in March to June and Septem- ber to October respectively. The annual mean temperature lies between 25—29°C. The eastern part of the Lama de- pression is crossed — in north-south direction — by the Oueémé River valley. Close to Lama Forest, there are on- ly few smaller creeks. During the rainy seasons, howev- er, large parts of the area are changed into overflooded marshlands. Due to these repeated inondations and stag- nant watres, there are only six dominant tree species in Llama Forest, viz. Dialium guineense, Diospyros mespili- formis, Albizia zygia, Afzelia africana, Khaja senegalen- sis, and Anogeissus leiocarpus. The Noyeau Central con- sists mainly of a mixture of relicts of a semideciduous for- est which floristically approaches the “Forét de Samba” Fig. 2. Primary forest of Forét de Lama (Noyeau central) (left) with bordering teak plantations (right) Photo: K. Ullenbruch ©ZFMK Reptiles from southern Benin, West Africa, with the description of a new Hemidactylus 33 Fig. 3. Landscape round the village of Didja, southern Benin. Photo: K. Ullenbruch type (Mondjkannagni 1969) and of secondary forest in various developmental stages. Within this dynamic sys- tem, the South American fast-growing asteracean neophyte Chromolaema odarata is commonly found along small roads and paths and at the forest edges. In smaller plan- tations, next to the larger ones with teak, trees such as Gmelina arborea, fraké (Terminalia superba), samba wood (Triplochiton skleroxylon) and, especially in the Tof- fo sector, the fire-resistent Cassia siamea are cultivated (Specht 2002). The remaining collecting sites in southern Benin were: — Didja village, situated ca. 25 km NW of Bohicon and largely surrounded by an agriculturally formed land- scape. The area north of Didja is characterised by bush and tree savannas (Figure 3). — Dan Forest or Forét de Dan is situated ca. 15 km NE of Didja and is regularly visited by hunters from Didja. Fig. 4. Gallery forests at the Zou River, southern Benin. Pho- to: K. Ullenbruch Bonn zoological Bulletin 57 (1): 31-54 —Za-Kpota, 18 km NE of Bohicon, is likewise largely sur- rounded by an agricultural area, replaced in the north- west by large tree savannas. 7 km N of Za-Kpota, the Zou River is running, being accompanied on its banks by a small strip of dense and degraded gallery forest which belongs to the hunting grounds of the villagers (Figure 4). — Lokoli Forest or Forét de Lokoli, a 500 ha large forest on the Hounto River, ca. 100 km N of Cotonou. Numer- ous sidearms of the Hounto River run through Lokoli Forest so that it is penetrable only by boat (Figure 5). For further details, in the context of an amphibian sur- vey, see Rédel et al. (2007). Flooded forest of Forét de Lokoli. Photo K. Ullen- Fig. 5. bruch Finally, some reptiles were collected also in Cotonou and Bohicon, on house walls and in gardens, and markets were visited to inspect the reptile species offered. Most of the reptiles were collected by UK by visual encounter and fo- cal sampling. Moreover, specimens were brought by the locals, particularly by two snake hunters from Didja and Za-Kpota respectively who even received an alcohol-filled bucket in order to keep interesting snake finds for some days. The Swiss zoologists of the Biolama project provid- ed us with few, but very important voucher specimens (e.g. Hemidactylus sp.n., Holaspis guentheri) taken in trunk eclectortraps used for the entomological survey. The survey of the reptile fauna of Lama Forest, South Benin, and the neighbouring localities mentioned above revealed the existence of 48 species (2 turtle, 1 tortoise, 20 lizard, and 25 snake species) which are listed and dis- cussed in the following. OZFMK 34 Klaus Ullenbruch et al. ANNOTATED LIST OF SOUTHERN BENINIAN REPTILES Testudinidae Kinixys belliana nogueyi (Lataste, 1886) Material examined. photographic voucher only (Figure 6). Remarks. This specimen was brought to KU by a local hunter. It was taken near Bohicon and was released after photographic documentation. Fig. 6. Kinixys belliana nogueyi from near Bohicon, south- ern Benin. Photo: K. Ullenbruch Pelomedusidae Pelomedusa subrufa (Lacépede, 1788) Material examined. photographic voucher only (Figure 7). Remarks. Specimen taken by local hunter near Bohicon, photographed and afterwards released. Pelusios castaneus (Schweigger, 1812) Material examined. photographic voucher only (Figure 8). Remarks. Specimen taken by local hunter near Bohicon, photographed and afterwards released. Agamidae Agama aff. agama (Linnaeus, 1758) Material examined. ZFMK 42019-024 (3 males, 3 fe- males), Godomey near Cotonou, coll. by W. Schréder, Au- Bonn zoological Bulletin 57 (1): 31-54 Fig. 7. Pelomedusa subrufa from near Bohicon, southern Benin. Photo: K. Ullenbruch gust/September 1984; ZFMK 77061-062, Bohicon, ZFMK 77063-064, Cotonou, all coll. by K. Ullenbruch, May/June 2002. Remarks. This common and widespread species was ob- served in all habitats including human settlements where vertical structures such as tree trunks and/or house walls were preferred. It was, however, never seen in the closed forest. Fig. 8. Pelusios castaneus from near Bohicon, southern Benin. Photo: K. Ullenbruch ©ZFMK Reptiles from southern Benin, West Africa, with the description of a new Hemidactylus 35 Agama sylvanus Macdonald, 1981 Material examined. ZFMK 77065, Forét de Lama (Noyeau Central), coll. by K. Ullenbruch, 6 June 2002. Remarks. The single juvenile specimen was obtained in an arthropod trap fixed at the trunk of a dead tree (trunk eclector) in dense primary forest. A second juvenile was observed at about 5 m height and escaped upwards. Dur- ing the three months of the survey, no further specimen of this agama was seen which argues for either a secre- tive lifestyle or a general rarity, at least in this season of the year. The morphological characters of ZFMK 77065 lie with- in the range of variation of A. sylvanus which was de- scribed by Macdonald (1981) from Ghana. Joger (1990) recorded the species also from Cameroon (as A. paraga- ma sylvanus). The disjunction between these two coun- tries is partially bridged by our specimen which represents the first country record for Benin. Chamaeleonidae Chamaeleo gracilis Hallowell, 1842 Material examined. ZFMK 77056-057, environs of Did- ja, coll. by a native hunter, 10 and 30 May 2002 resp. Remarks. The two males were obtained from a native hunter on the market of Abomey who certified their ori- gin as Close to Didja. C. gracilis is widespread in the Su- dan and Guinea savannah belt south of the Sahara. The rudimentary occipital flaps distinguish it from the follow- ing, only recently described species. Chamaeleo necasi Ullenbruch, Krause & Bohme, 2007 Material examined. ZFMK 77058, male, Za-Kpota, ZFMK 77059-060, 2 females, Houégbo, all coll. by K. Ul- lenbruch, May/June 2002. Remarks. The three specimens form part of the type se- ries of this species which is otherwise known only from Togo (Ullenbruch et al. 2007). The Togolese localities (Kpalimé, Missaboué) as well as the two Benin localities are situated close to relict forest patches within the arid Dahomey gap. Za-Kpota is an agriculturally shaped site not far from the river Zou which has some degraded gallery forest vegetation, while Houégbo is not far from Lama Forest. Bonn zoological Bulletin 57 (1): 31-54 C. necasi belongs to the C. dilepis group and differs from the south Nigerian populations of the closely related C. quilensis mainly by hemipenial characters (see Ullenbruch et al. 2007). Chamaeleo senegalensis Daudin, 1802 Material examined. ZFMK 42025 (male), Bohicon, coll. by W. Schréder, August/September 1984; ZFMK 77052- 055, environs of Didja, coll. by a native hunter, 10 May 2002. Remarks. The four specimens were obtained from a na- tive hunter on the market of Abomey who certified their origin as close to Didja. C. senegalensis — a species of the C. dilepis group without any traces of occipital flaps — was found on all markets in southern Benin and proved to be much more common than C. gracilis. Eublepharidae Hemitheconyx caudicinctus (Dumeéril, 1851) Material examined. ZFMK 77066, near Didja, ZFMK 77067, Za-Kpota, both coll. by native collectors, 27 May and 18 June 2002, respectively. Remarks. A West African species distributed from Sene- gal to Cameroon which has its southernmost localities in Benin. In Benin, it is termed “snake with two heads” by the locals and thought to be very dangerous. Gekkonidae Hemidactylus ansorgii Boulenger, 1901 Material examined. ZFMK 77073-075, Lama Forest (Noyeau Central), coll. by K. Ullenbruch, 18. (77073) and 28. April 2004. Remarks. This slender, narrow-headed forest gecko with a long, cylindrical tail has only rarely been encountered and is known with certainty only from Liberia, Ivory Coast, Nigeria, and Cameroun (Henle & Bohme 2003). Bauer et al. (2006) expected H. ansorgii to occur in Benin, due to its wide West African distribution but stressed that there are no confirmed records from this country. Thus, our specimens represent the first record for Benin (Fig- ure 9). The geckos were found in the late morning (11.30h—12.30h) on tree trunks in dense primary forest at about 1.7 m height. ©OZFMK 36 Klaus Ullenbruch et al. Fig. 9. Hemidactylus ansorgei from Lama Forest (Noyeau central), first country record. Photo: K. Ullenbruch Hemidactylus angulatus Hallowell, 1852 Material examined. ZFMK 42006-009, Godomey near Cotonou, coll. by W. Schréder, August/September 1984; ZFMK 77070, Bohicon, coll. by K. Ullenbruch, 17 April 2002; ZFMK 82813-814, Cotonou, coll. by Thibault La- chet, 22/23 April 2004. Remarks. Thys van den Audenaerde (1967) distinguished between Hallowell’s forest-dwelling species Hemidacty- lus angulatus and the savannah form H. brookii guineen- sis Peters, 1868. Since the African taxa of the former con- cept of an Afro-Indian species H. brookii Gray, 1845 are not at all close relatives of the true H. brooki from India (Carranza & Arnold 2006), it is still unclear whether the name guineensis should be treated as a synonym or a sub- species of angulatus, or possibly even a species. Accord- ing to Bauer & Giinther (1991), it is a synonym of angu- latus (understood by these authors still as a subspecies of H. brookii). Examination of the lectotype (Figure 10) of H. guineen- sis Peters, 1868 [described from “Ada Foah, Guinea” (= Adafoa, Ghana; not Adafer, Mauritania, as suggested by Loveridge 1947 and Wermuth 1965!) and compared by Pe- ters (1868) only with H. verruculatus (= H. turcicus)| re- vealed that despite its provenance from coastal West Africa, it resembles the savannah form rather than the for- est form which would strengthen a partial sympatry with the forest-dwelling H. angulatus. However, the distin- guishing key character between both forms, i.e. the num- ber of granules bordering the nostril above rostral and first supralabial (Thys van den Audenaerde 1967), has been dis- proved already by B6hme (1978), and other scalation dif- ferences also seem to be insufficient for a clear separa- tion of both forms. Habitus differences could possibly be found in colour pattern, which often seems to consist of Bonn zoological Bulletin 57 (1): 31-54 Fig. 10. Hemidactylus angulatus / guineensis complex: lecto- type of Hemidactylus guineensis Peters, 1868. Photo: P. Wagn- er paired dorsal bars interrupted on the dorsal midline in the savannah populations, combined with more distinctly white-coloured single tubercles in the flank region. In the forest-dwelling populations, it consists of smaller, less symmetrically arranged darker elements, with less obvi- ous white flank tubercles in the forest populations. Steindachner (1870) described H. affinis from Gorée and Dagana, Senegal, and diagnosed it also only against H. verruculatus (= H. turcicus). He stressed the similarity of these two species, particularly concerning the size of their strongly keeled tubercles. The type series of H. affinis closely resembles the savannah-dwelling H. guineensis. In his description of H. stellatus, Boulenger (1885) com- pared it with H. brookii and H. gleadovii and stressed the presence of pure-white tubercles intermixed with a ma- jority of dark brown ones which inspired him to the name stellatus (= starred). These white tubercles are indeed more obvious in the savannah-dwelling form (Figure 11) and TT Fig. 11. Hemidactylus angulatus / guineensis complex: lecto- type of Hemidactylus stellatus Boulenger, 1885. Photo: C. Mc- Carthy ©OZFMK Reptiles from southern Benin, West Africa, with the description of a new Hemidactylus 37 Fig. 12. Hemidactylus angulatus (s.str.) from Godomey near Cotonou (left), H. mabouia from Cotonou (center) and H. aff. mabouia, also from Cotonou (right). Photo: P. Wagner brought also Chabanaud (1917 a) to the conclusion that there are two different species in this group of geckos, viz. “H. brookii” (certainly meaning angulatus) and “H. stel- latus” (= H. guineensis), at Agouagon, Benin, even in sympatry. Preliminary genetic support for two species within a H. angulatus/guineensis complex may be deduced from the tree in Carranza & Arnold (2006) where a Kenyan Rift Valley sample is closer to dry savannah samples from Mauritania and Mali (guineensis) than to a forest sample from Bioko (angulatus s.str.). Moreover, as already point- ed out by Thys van den Audenaerde (1967), the forest species H. angulatus does not occur in Central and East Africa. Bonn zoological Bulletin 57 (1): 31-54 Bauer et al. (2006) who provided locality data for many specimens of their H. angulatus (sensu lato) from all over Benin, did not distinguish between the two forms thus im- plying morphological uniformity, except their variable colour pattern. They stressed that the sympatric H. mabouia could be easily mixed up with H. angulatus what we also found to be true (but only for the forest form: an- gulatus s.str.) particularly when the diagnostic (see the key in Bauer et al. 2006) characters of tubercles on head and dorsum are concerned. Here, again the colour pattern of the dark phases of H. angulatus and H. mabouia may prove to be more reliable than the scalation (Figure 12). Concluding it becomes clear that Hemidactylus angula- tus S.]. 1s in urgent need of a taxonomic revision through- out its range. ©ZFMK 38 Klaus Ullenbruch et al. ZFMK 77070 was captured on a house wall on 19.30 h, i.e. 30 min. after sunset. Hemidactylus fasciatus Gray, 1842 Material examined. ZFMK 82808, Forét de Lama (Noyeau Central), coll. by Lamidi Konetche, April 2004. Remarks. This species was first documented for Benin by Bauer et al. (2006) who recorded specimens from the vicinity of Kétou. Our record, the second for Benin, is from dense primary rain forest. Hemidactylus mabouia Moreau de Jonnés, 1818 Material examined. ZFMK 77068-09, Cotonou, coll. by K. Ullenbruch, Datum; 82809-812, Cotonou (Maison des Stagnaires), coll. by Thibault Lachet, 22/23 April 2004. Remarks. Not found by W. Schréder in 1984 and docu- mented for the first time in Benin by Bauer et al. (2006). This species was considered to be rare in Nigeria and be- lieved to be introduced in port cities by Dunger (1968). Also, R6dder & B6hme (2008) regarded it as endemic on- ly in Central and East Africa, while it is an invasive species —next to South America, the Caribbean region and Flori- da — also in West Africa. ZFMK 77068-09 differ from typical H. mabouia by lack- ing the very characteristic dark pattern of oblique curved cross stripes (Figure 12). Similar phenotypes have been found in western Guinea (Bohme et al., unpubl. data). As no scalation differences could be detected, we regard these specimens currently as a colour morph of H. mabouia, pending new material for genetic analyses. The two specimens were collected by UK at noon time (12.00h) from house walls during a heavy thunderstorm rain. The geckos were observed to hunt for insects which were beaten to (or at least close to) the ground by the rain- water. The reduced daylight — a consequence of the dense clouds — might have helped the geckos to become active and to forage during midday. The masses of insects con- centrating under these conditions within the lower two me- ters above the ground were also used and exploited by Agama aff. agama and Mochlus guineensis. Hemidactylus lamaensis new species Holotype. ZFMK 77072, adult female, Forét de Lama (teak plantation), coll. from a pitfall by Klaus Ullenbruch, 16 April 2002. Bonn zoological Bulletin 57 (1): 31-54 Diagnosis. A small Hemidactylus with a SVL of about 5.0 cm. Characteristic is the dorsal scalation which consists of numerous irregular rows of large, convex, keeled tu- bercles which are nearly juxtaposed on the dorsal hind- part and on the tail root, separated mostly by only one row of surrounding, rather coarse granular scales. Also the tem- ple is covered by large, juxtaposed tubercles with only few or even without granules between them. Moreover, the subdigital lamellae of the first digit and toe are very low in number, the terminal unpaired lamella of digit I (manus) being the only adhesive element while digit I (toe) has two scansors: one divided proximal and one undivided termi- nal one. Description of the holotype (Figure 13). Habitus stout, head small with an elongated snout. Head & body length 47.7 mm, length of (regenerated) tail 34 mm. Snout elon- gate: distance from anterior margin of tympanum to hind- margin of eye 4 mm, from anterior margin of eye to tip of snout 5.5 mm; eye diameter 2.5 mm. Rostral broad, with a deep notch on its anterior midline which continues as a vertical suture but does not separate the rostral complete- ly. Nostril bordered by rostral, a Ist labial, three postnasal scales, and a supranasal which is separated from its fel- low by one single median scale. Scales on snout small but polygonal rather than granular, keeled, passing backwards into a tuberculate scalation of the frontal and parietal re- gion which becomes intermixed with granules from the occipital region towards the neck and dorsum. Temple covered with conical tubercles with only few intermixed granules between them. Mouthline straight and not uplift- ed in its hindpart. Supralabials (left/right) 10/10, sublabi- als (left/right 8/8). Mental triangular, followed by two pairs of postmentals, the first of them broadly meeting at the midline behind mental. Gulars small, smooth, passing pos- teriorly gradually into the semicircle-shaped, subimbricate and likewise smooth chest and ventral scales. The coni- cal and weakly keeled dorsal tubercles are arranged in 16-18 irregular longitudinal rows, circumdated by rather coarse granular scales which separate the tubercles by 1-3 rows, while on sacrum and tail root the tubercles are even nearly juxtaposed with only few scattered granules be- tween them. Limbs covered above with coarse granular scales intermixed with conical, weakly keeled tubercles. Palmar scales smooth. All fingers and toes with distinct claws, and with divided adhesive lamellae (scansors) be- neath, except the terminal one which is simple. Also the basal subdigital scales are undivided and are added to the number of subdigital lamellae when they are twice as large than the palmar scales. The values are: 1-5-6-7-6 (left manus), |-5-6-7-5 (right manus), 2-6-7-8-7 (left pes), and 2-6-6-7-6 (right pes). The regenerated tail is dorsally cov- ered with very regular subimbricate smooth scales which gradually pass into several longitudinal rows of oblique- ly enlarged, imbricate subcaudal plates. ©ZFMK Reptiles from southern Benin, West Africa, with the description of a new Hemidactylus 39 Fig. 13. Hemidactylus lamaensis n. sp., holotype from Lama Forest (Noyeau central) in dorsal (a), ventral (b) and lateral (c) view. Photo: P. Wagner Bonn zoological Bulletin 57 (1): 31-54 OZFMK 40 Klaus Ullenbruch et al. Colour in preservative. Upper side rather uniformly light brownish, with a darker brown head, and one indistinct darker crossband on the neck and on the shoulder respec- tively. Underside uniform whitish-yellowish. Etymology. The species name refers to Lama Forest, so far the only known locality of the new Hemidactylus. It is intended to stress the importance of this relict forest within the Dahomey gap also in conservational aspect as it may harbour still more unknown rain forest species. Distribution and natural history. Only known from the type locality. The single female was taken from a pitfall on the forest floor in a young teak plantation bordering the primary forest of the Noyeau Central. This does not argue against an arboricolous lifestyle, because also tree- dwellers like H. fasciatus were observed to climb down from their shelter in big fig trees and to forage on the ground some hours after sunset (BGhme 1975). Comparisons. This unique single specimen cannot be as- signed to any West African species (see Bauer et al. 2006). Its snout is elongated but not to the extent observed in H. ansorgei. Its most obvious character is the possession of densely arranged, large, keeled tubercles which are less numerous as compared to those of the UH. angulatus/guineensis complex. On the hindpart of the dor- sum and the tail root, they are even nearly juxtaposed. Al- so in contrast to the situation found in the former com- plex is the dense coverage with tubercular scales of the temporal region. Moreover, it has the lowest number of subdigital lamellae of all Beninian and West African Hemi- dactylus. Assuming that the original tail in this species has also se- ries of tubercles (not present in the regenerated tail of the unique holotype), the key given by Bauer et al. (2006) for the geckos of Benin can be completed as follows: 13a. Dorsal tubercles prominent, keeled ............... 14 13b. Dorsal tubercles not, or only weakly keeled, sepa- rated by one another by a distance greater than by tuber- cle width; dorsum of head atuberculate or with few small tubercles; color variable, the typical pattern of a series of dark curved crossbands distinct in the dark phase only BHO tpn Neath etna wen ce aac re: Hemidactylus mabouia 14a. Dorsal tubercles separated by one another by approx- imately the width of one tubercle; tubercles usually promi- nent on dorsum of head; temple granular By eisai Hemidactylus angulatus/guineensis complex 14b. Dorsal tubercles separated by one another by less than a tubercle width, on sacrum and tail root even abutting; temple covered with conical tubercles a Snaislatee Danis Set arers AGES Hemidactylus lamaensis sp. nov. Bonn zoological Bulletin 57 (1): 31-54 Scincidae Panaspis togoensis (Werner, 1902) Material examined. ZFMK 77079-080, Forét de Lama, coll. by K. Ullenbruch, 24 and 28 April 2002. Remarks. Both voucher specimens were collected in a transition zone of primary forest and a Chromolaema clearing, after heavy rainfalls around noon time. Consid- ered to be ombrophilous by Perret (1973), activity after rains was also observed by Rédel et al. (1997). Accord- ing to Fuhn (1972) and Schmitz et al. (2005), Lygosoma (Riopa) dahomeyense Chabanaud 1917 is a synonym of P. togoensis. Mochlus guineensis (Peters, 1879) Material examined. ZFMK 77076, Forét de Lama; ZFMK 77077-078, Cotonou, all coll. by K. Ullenbruch, 16 and 13 April 2002 respectively. Remarks. The specimen from Lama Forest was taken in a pitfall trap placed in an old teak plantation close to the primary forest. The two specimens from Cotonou were caught in the garden of a house during heavy rainfall when they were foraging for prey negatively influenced by the rains (see above, under Hemidactylus mabouia). Remarkably, the specimen from Lama Forest differs in some characters from the two Cotonou specimens. It is more delicately built, having thinner limbs and a longer and slimmer tail. Moreover, the eyes seem to be smaller, the overall colouration is distinctly darker, and the sub- caudal scales are spotted with black, while the Cotonou specimens have the underside of their tails purely white. More material is necessary to evaluate these differences (Figure 14). Trachylepis affinis (Gray, 1838) Material examined. ZFMK 42004-005, Godomey near Cotonou, coll. by W. Schréder, August/September 1984; ZFMK 77081-090, Forét de Lama, coll. by K. Ullenbruch, 16 April to 1 May 2002. Remarks. The two specimens from Godomey come from a farmland/garden environment, the Lama Forest individ- uals, however, were all collected within dense primary for- est (Noyeau Central), on clearings grown with Chromo- laema, and in the neighbouring teak plantations. The lizards were found either in leaf litter and on the grassy floor, or on tree trunks. When disturbed they tried almost ©OZFMK Reptiles from southern Benin, West Africa, with the description of a new Hemidactylus 4] Fig. 14. Mochlus guineensis, from Cotonou (above) and Lama Forest (Noyeau central) (below). Photo: P. Wagner invariably to escape upwards. They were mainly active be- tween 11.00h and 15.00h, at air temperatures of 31—32°C. Half of the Lama Forest sample has a distinctly striped colour pattern with a conspicuous, white sublabial stripe running along the lower flank towards the tail, while the other half lacks this stripe. Trachylepis perroteti (Duméril & Bibron, 1839) Material examined. ZFMK 41998-999 (2 males), Godomey near Cotonou, coll. by W. Schréder, August/Septeber 1984. Remarks. According to Mausfeld-Lafdhyia et al. (2004) and Chirio & LeBreton (2007), 7: perroteti consists of at least two distinct species which require further study (see also Stoll 2008). But because the true 7. perroteti (s.str.) is the more southernly distributed from of the two cryp- tic siblings in Cameroon, the two males from near Coto- nou are most likely belonging to this latter species. Whether this is also true for the North Beninian popula- tions (see the country checklist below) remains to be stud- ied. Lacertidae Holaspis guentheri Gray, 1863 Material examined. ZFMK 77071, Forét de Lama (teak plantation), coll. by K. Ullenbruch, 26 June 2002. Remarks. The only voucher specimen was obtained from an arthropod trap (trunk eclector) fixed on a dead teak tree (Tectona grandis) within an old teak plantation bordering the primary forest of Forét de Lama. This little lacertid Bonn zoological Bulletin 57 (1): 31-54 with the ability of gliding flight has not been recorded so far from Benin, thus representing the first country record. Varanidae Varanus exanthematicus (Bosc, 1792) Material examined. ZFMK 77004-005, near Didja, coll. by local hunter, 1 May 2002. Remarks. Both specimens were obtained from a local hunter at Abomey who certified their origin as being close to Didja. Varanus niloticus (Linnaeus, 1766) Material examined. ZFMK 42200, Godomey near Coto- nou, coll. by W. Schréder, August/September 1984; ZFMK 77001, juv., Ouémé River near Togbota, coll. by local hunter, 15 April 2002. Remarks. ZFMK 77001 was found under a heap of dry wood on the bank of a small tributary of the Ouémé Riv- er, close to the settlement of Togbota. Varanus ornatus (Daudin, 1803) Material examined. ZFMK 77002-003, near Didja, coll. by native hunter, 10 and 30 May 2002 respectively. Remarks. Both specimens were obtained from local hunters at Abomey who had collected them in the vicin- ity of Didja. One of them originated from Lakpo, 8 km NE of Didja, which is in immediate vicinity of a further relict forest patch, viz. Foret de Dan. A big, adult speci- men was seen in hectic flight in Lama Forest and could not be caught. On markets, V. ornatus was much less commonly seen than V. niloticus which reflects also the scarcity of relict forest patches in the Dahomey gap which serve as refu- gia for this forest species. For Benin, our specimens rep- resent the first country record (B6hme & Ziegler 1997). Pythonidae Python regius (Shaw, 1802) Material examined. ZFMK 41993-995, Godomey near Cotonou, coll. by W. Schréder, August/September 1984; Photographs from Za-Kpota and Forét de Lama. ©OZFMK 42 Klaus Ullenbruch et al. Remarks. Of ZFMK 41995, only head and neck are pre- served. This species was seen in great numbers on all mar- kets visited. These great numbers certainly pose a serious conservation problem which could be buffered by farm- ing royal pythons for the international pet trade. Howev- er, such farms will be very difficult to be controlled as far as wild-caught specimens are concerned. Python sebae (Gmelin, 1788) Material examined. None. Remarks. Rock pythons could be seen only twice on mar- kets by UK during his survey. He was told by the people that they fear the large snakes and normally kill them whenever possible. This species is subject to superstitious beliefs by the local people. For instance, a scientific as- sistant of the BIOLAMA project even told that rock pythons are capable to cast out a net out of their mouth to catch humans and to eat them subsequently. Colubridae (sensu lato) Afronatrix anoscopus (Cope, 1861) Material examined. ZFMK 77006 (juvenile), Forét de Lokoli, coll. by K. Ullemnbruch, 5 May 2002. Remarks. The existence of this aquatic species in Benin was considered as likely by Chippaux (2001). Our spec- imen (Figure 15) was the first documented record from Benin until very recently Chirio & Ineich (2009) published a find from Atakora Province which makes our specimen the 2nd published record. It is, however, still interesting to note that both new records of this species are very dis- tant from each other, ranging from a relict, swampy rain forest to dry Sudan savannah. Crotaphopeltis hotamboeia (Laurenti, 1768) Material examined. ZFMK 42001-002 (juveniles), Godomey near Cotonou, coll. by W. Schréder, August/September 1984. Remarks. None. Dasypeltis gansi Trape & Mané, 2006 Material examined. ZFMK 77014, Za-Kpota, coll. by na- tive hunter, may 2002. Bonn zoological Bulletin 57 (1): 31-54 Fig. 15. Afronatrix anoscopus, juvenile, from Lokoli Forest, first country record. Photo: K. Ullenbruch Remarks. Trape & Mané (2006 a) partitioned the West African egg-eater snakes formerly referred to as D. scabra into 3 different species, one of them with a likewise new subspecies: D. confusa, D. gansi (with D. g. gansi and D. g. latericia), and D. sahelensis. According to the original description and to the key and colour plates of Trape & Mané (2006 b) the specimen from Za-Kpota clearly rep- resents the nominotypic subspecies of D. gansi. It is the second voucher from Benin, the first (Trape & Mané 2006 a) Benin record being from Lanta (where it is, according to the same authors, sympatric with D. confusa). The record of Chabanaud (1916) of “D. scabra palmarum” refers very likely also to D. g. gansi which is the only of the newly described taxa which tends to be patternless. Dispholidus typus (Smith, 1829) Material examined. ZFMK 77029, between Bohicon and Didja, coll. by K. Ullenbruch, May 2002. Remarks. Found dead on the road between the two above- mentioned places. Lamprophis fuliginosus (Bote, 1827) Material examined. ZFMK 77011, Didja; 77015-017, Za- Kpota; 77041, Forét de Lama; 77051, Forét de Dan, all coll. by K. Ullenbruch, 14-18 June 2002. Remarks. ZFMK 77041 was found dead on a path with- in Lama Forest, just between primary forest and the neigh- bouring teak plantation. Although still rather fresh, its head was already badly damaged by ants. ©ZFMK Reptiles from southern Benin, West Africa, with the description of a new Hemidactylus 43 Lycophidion semicinctum Duméril, Bibron & Duméril, 1854 Material examined. 77007, near Didja, coll. by native hunter, May 2002; ZFMK 77050, near Bohicon, coll. by K. Ullenbruch, May 2002. Remarks. The Bohicon specimen was found dead on a road. Meizodon regularis Fischer, 1856 Material examined. ZFMK 42003, Godomey near Coto- nou, coll. By W. Schréder, August/September 1984. Remarks. Known from, among else, Ghana (where the type locality is situated), and Togo west of Benin, as well as from Cameroon further east (Roux Estéve 1969), this species was to be expected in Benin but was obviously only indicated for Benin by Chippaux (2001). ZFMK 42003 represents the second country record and first pub- lished voucher specimen for Benin. Natriciteres olivacea (Peters, 1854) Material examined. ZFMK 77039, Forét de Lama, coll. by K. Ullenbruch, May 2002. Remarks. Found on a cloudy day at 11.20h on the forest floor. This underlines that this species is not confined to savannah habitats as claimed by Broadley (1983) but that at least in West Africa also forests are inhabited (see Vil- liers 1975, Chippaux 2001: distribution map). Natriciteres variegata (Peters, 1861) Material examined. ZFMK 77042-046, Foret de Lama, coll. by K. Ullenbruch, May 2002. Remarks. All five individuals were found on the forest floor. The species proved to be rather common but was never seen outside the forest. Also Dunger (1972a), Broadley (1983) and Chippaux (2001) regard N. variega- ta as a forest species. The only earlier record of it was Cha- banaud’s report from “Dahomey”, thus our small series represents the first documented locality record for Benin. Philothamnus irregularis (Leach, 1819) Material examined. ZFMK 77025-026, Za-Kpota, coll. by native snake hunter, May 2002; ZFMK 77040, Foret de Lama: Koto village, coll. K. Ullenbruch, May 2002. Bonn zoological Bulletin 57 (1): 31-54 Remarks. The last mentioned specimen was killed by a local farmer in the presence of KU on a sun-exposed grassy area. Philothamnus semivariegatus (Smith, 1847) Material examined. ZFMK 77034-035, Bohicon, coll. by K. Ullenbruch, May 2002. Remarks. Both snakes were encountered on the compound of a mill. One of them was found on the floor of a house belonging to that compound. Encounters in human habi- tations and even within buildings were also reported from Nigeria by Dunger (1973a). Psammophis elegans (Shaw, 1802) Material examined. One specimen, uncatalogued, Coto- nou (stored in the private collection of M. Serge Attignon, Cotonou), coll. by a S. Attignon, 11 May 2002. Remarks. The only available specimen of this snake was found in front of a house in the outskirts of Cotonou. Psammophis phillipsi (Hallowell, 1844) Material examined. ZFMK 41996, Godomey near Coto- nou, coll. by W. Schréder, August/September 1984. Remarks. The large adult, dark-brown unicoloured female is allocated to Hallowell’s species because the mid-region of its ventral plates has a broad median zone shaded with bluish-grey colour. We think that this is a species of forest- ed and moist savannahs which is unicoloured above al- ready as a juvenile has and lacks the typical striped juve- nile dress of the following form (Bohme et al. 1996). Psammophis cf. phillipsi (Hallowell, 1844) Material examined. ZFMK 77024 & 77028, Za-Kpota, coll. by a native snake hunter, May 2002; ZFMK 77047, Abomey, coll. by K. Ullenbruch, May 2002. Remarks. The three specimens belong to the big-growing form of the Psammophis sibilans species complex which is dorsally striped as a juvenile and unicoloured above and below as a large, old adult. It inhabits the dry Sudanian and Sahelian savannahs and is mostly referred to as P. cf. phillipsi (e.g. Bbhme 1978, Bohme et al. 1996) or, recent- ly, again as true P. sibilans (Trape & Mané 2006 b). Al- ©OZFMK 44 Klaus Ullenbruch et al. Fig. 16. Psammophis sudanensis from Bohicon, southern Benin. Photo: K. Ullenbruch so earlier records of P. sibilans from West Africa (e.g. Cha- banaud 1916) refer to this form which, in our opinion, is not conspecific with the true P. sibilans from Egypt. Many not yet fully grown but nonetheless adult (mature) spec- imens may retain more or less distinct reddish dorsolat- eral stripes along the body. The belly is light whitish to yellowish and patternless (no dark hairlines along the ven- tral plates), but supralabials, sublabials and gulars show some dark pigmented spots (see BOhme et al. 1996). The anal shield is entire. Its taxonomic status and nomenclat- ural status needs still to be assessed. Psammophis sudanensis Werner, 1919 Material examined. ZFMK 77012, Didja, coll. native hunter; ZFMK 77018-019; ZFMK 77036-038, Bohicon, and ZFMK 77047, Abomey, all coll. by K. Ulenbruch, May 2002. Remarks. The specimens represent various age classes. All have distinct hairlines along the outer parts of the ventral shields, however, these hairlines are brownish in younger and darker in the older individuals. The zone between these hairlines is differently coloured than the two outer, ventrilateral zones, the latter being white while the inner is yellowish-whitish. This ventral pattern is therefore of the “subtaeniatus type” and agrees with the type speci- men of sudanensis Werner, 1919 (NMW 1986 from Kadugli, Kordofan) which actually was described as P. subtaeniatus sudanensis (Werner 1919, Bohme 1987). In all specimens, the anal shield is divided. ZFMK voucher specimens from e.g. northern Senegal (Dakar, Fété Olé), Algeria (Tamanrasset), North Cameroon (Mokolo) (see B6hme 1986) and Chad (ZFMK 74520) are Bonn zoological Bulletin 57 (1): 31-54 Fig. 17. The same specimen from below. Photo: K. Ullenbruch very close to typical P. sudanensis but differ in having e1- ther faint, light brown hairlines (instead of blackish ones) or no hairlines at all; their undersides are in both cases of a uniform, bright porcellanic white. They correspond to the type specimen of P. leucogaster Spawls, 1983 (BMNH 1980: 261, from Wa, northern Ghana, see Spawls 1983), and further sampling must show whether both forms have zones of intergradation. It seems that if there is a distinction between typical sudanensis and leucogaster in West Africa, this is a question of a north-south rather than a west-east disjunction. ZFMK 77036-038 were found on the compound of a mill at Bohicon. One of them was lying in the afternoon on the terrace of a house, the other two were killed during grass mowing (Figs 16 and 17). Rhamphiophis oxyrhynchus (Reinhardt, 1843) Material examined. ZFMK 41997, Godomey near Coto- nou, coll. by W. Schréder, August/September 1984; ZFMK 77020-023, Za-Kpota, coll. by a native hunter, May 2002. Remarks. None. Scaphiophis albopunctatus Peters, 1870 Material examined. ZFMK 77009-010 (juvenile and adult), Didja, collected by a local snake hunter, May 2002. Remarks. The juvenile specimen contained 3 newborn mice which could not be identified to the familial or gener- ic level. ©OZFMK Reptiles from southern Benin, West Africa, with the description of a new Hemidactylus 45 Atractaspididae Amblvodipsas unicolor (Reinhardt, 1843) Material examined. ZFMK 77027, Za-Kpota, coll. by na- tive hunter, May 2002. Remarks. When fixing this snake, some white nematodes appeared between the scales and tried to leave the speci- men. This was not observed in any other snake during preparation. Elapidae Dendroaspis viridis (Hallowell, 1844) Material examined. ZFMK 77049, Bohicon (market), coll. by K. Ullenbruch, May 2002. Remarks. Since this snake was found on a market, no fur- ther information on its precise provenance is available. Elapsoidea semiannulata Bocage, 1882 ZFMK 77013, near Didja, coll. by a native snake hunter, May 2002. Remarks. Represents the West African subspecies E. s. moebiusi Werner,1897 (Trape & Mané 2006). Naja nigricollis Reinhardt, 1843 Material examined. ZFMK 77032, Foret de Lama; ZFMK 77033, Bohicon, both coll. by K. Ullenbruch, May 2002. Remarks. One of the two juvenile specimens was found dead on a road at the margin of Lama Forest, the other was killed during grass mowing in the mill compound of Bohicon. Viperidae Bitis arietans (Merrem, 1820) Material examined. ZFMK 77048, environs of Abomey, coll. by native hunter, May 2002. Remarks. B. arietans is very commonly offered on mar- kets. Usually, the head is cut off, dried and used for Voodoo ceremonies while the rest of the animal is used as food. Bonn zoological Bulletin 57 (1): 31-54 Causus maculatus (Hallowell, 1842) Material examined. ZFMK 77030, between Bohicon and Didja, ZFMK 77031, 9 km east of Abomey, both coll. by K. Ullenbruch, May 2002. Remarks. The first specimen of this ubiquitous species was found dead on the asphalt road between Bohicon and Did- ja, the second one was killed by locals next to the street to Abomey in a crop field. Echis ocellatus Stemmler, 1970 Material examined. ZFMK 77008, near Didja, coll. by lo- cal snake hunter, May 2002. Remarks. None. CONCLUSIONS Of the 48 species recorded above from several localities in southern Benin, including the relict rain forests Forét de Lama and Forét de Lokolhi, only eight can be regard- ed as true forest species. These are: Agama sylvanus, Hemidactylus ansorgei, H. fasciatus, H. mabouia, H. lamaensis sp. n., Holaspis guentheri, Varanus ornatus, and —as the only snake — Psammophis phillipsi. Two of them, Hemidactylus fasciatus and H. mabouia, are recorded from Benin for the second time, H. mabouia, however, being a recent invasive colonizer. The remaining five lizard species, all from Lama Forest, are first country records for Benin, one of them, H. Jamaensis sp. n., being new to sci- ence. Three snake species, viz. Afronatrix anoscopus, Amblyo- dipsas unicolor, and Natriciteres variegata, are found in both forest and savannah habitats. Of these, Afronatrix anoscopus 1s recorded here for the second time from Benin, but for the first time from southern Benin (the very recent first record being from the Atakora chain in the northwestern part of the country). Amblyodipsas unicol- or and Natriciteres variegata are both representing the re- spective first country records documented by voucher specimens. The remaining 40 species are widely distributed in the West African savannas and have in most cases been recorded from Benin before. Nonetheless, particularly some snakes have been listed and/or plotted in maps of review papers and books by e.g. David & Ineich (1999), Chippaux (1999, 2001, 2006), or Trape & Mané (2006 b) without locality and voucher documentation of the respec- tive country records, so that our specimens are in some ©ZFMK 46 Klaus Ullenbruch et al. cases nonetheless the first documented records, despite earlier citations of the respective species from Benin. In the following country-wide checklist, we included al- so some new data derived from the mission to Pendjari National Park by one of authors (OG). Among these, the by far most remarkable find is the rediscovery of Agama gracilimembris in Benin, from where it was first described about ninety years ago. COUNTRY-WIDE CHECKLIST OF THE REP- TILES OF BENIN The following list contains several records of species col- lected by the Lama Forest survey of the senior author which have not been documented before from the terri- tory of Benin. However, as nearly all these species were already known from the neighbouring countries in the west (Togo) and in the east (Nigeria), their presence in Benin is not surprising and was already regarded as probable in several cases by Villiers (1951), Chippaux (2001) or Bauer et al. (2006). More general books on West Africa or even Africa as a whole (e.g. Villiers 1976, Welch 1982, Chip- paux 1999, 2001, 2006, Trape & Mané 2006 b) often pose the problem that a species is assumed to occur in a spe- cific country (e.g. formulations as “from Senegal to Ugan- da’’) rather than being documented from a specific local- ity within the respective country; or that it is plotted on a map without documentation of the origin of the respec- tive data. For completeness’ sake, we included in this list some species recorded by Chabanaud (1916) which are definite- ly lacking in West Africa and thus are probably due to misidentifications. They are marked with an asterisk (*). A re-examination of the respective voucher material as far as still extant was, however, beyond the scope of this pa- per. It is also noted here that the most important place of origin of Chabanaud’s material collected by Dr. Bouet is constantly spelled Agouagon in his papers, as it is also by Villiers (1951). However, more recent authors (e.g. Hoog- moed 1974, Salvador 1982) spell it “Agouagou” under which name it is also found on modern road maps. Here, we follow the original spelling. It was reflecting the still poor state of knowledge of the reptilian fauna of Benin when Loveridge (1952) stated that among the turtles and lizards from the A. Villiers mission (1950) no less than 8 species were new for the country, among them even Agama agama! More than five decades later Bauer et al. (2006) recorded — next to more first records of geckos and a new species of Hemidactylus — Bonn zoological Bulletin 57 (1): 31-54 also the common H. mabouia as new for the country, and in this paper, we can list — again next to another new Hemi- dactylus — further 8 species as new for Benin’s reptile fau- na. Our reference to Pendjari NP and to Bemberéké are based on data and photographic documentations collect- ed by one of us (OG) and of a small donation of snakes to ZFMK by T. & M. Imthurn (2000). For the latter, the ZFMK catalogue numbers are given. Geoemydidae *Mauremys leprosa — Chabanaud (1917): Porto Novo (as Clemmys leprosa). Not in Fritz & Havas (2007). Accord- ing to Fritz (pers. comm.) this record is certainly due to an error and should be deleted from the country’s species list. Testudinidae Kinixys belliana nogueyi — Loveridge (1951): Bassila; this paper: Bohicon-Abomey area; Pendjari NP. Pelomedusidae Pelomedusa subrufa — Loveridge (1951): Bassila; Zag- nanado; this paper: Lama Forest surroundings; Pendjari NP. Pelusios castaneus — Sinsin et al. (1999): Benin, unspec- ified (as P. subniger); Branch (2008): on map, but with- out locality data; this paper: Lama Forest surroundings; Pendjari NP (ZFMK 83613). Trionychidae Cyclanorbis elegans — Gramentz (2008): “Benin”, with- out specific locality. Cyclanorbis senegalensis — Gramentz (2008): Pendjari NP (based on Grell 2003); this paper: Pendjari NP (Figure 18). Crocodylidae Crocodylus suchus — This paper: Pendjari NP (Figure 19). Osteolaemus tetraspis — Sinsin et al. (1999): “Sud-Benin” including the Departments of Zou, Mono, Ouémé, and At- lantique. ©ZFMK Reptiles from southern Benin, West Africa, with the description of a new Hemidactylus 47 Fig. 18. Cyclanorbis senegalensis, juvenile from Pendjari NP, northern Benin. Photo: O. Grell Eublepharidae Hemitheconyx caudicinctus — Chabanaud (1917 a): “Haute Dahomey”; Agouagon (as Psilodactylus caudicinc- tus); Bauer et al. (2006): Kétou; Diho (=Idiho); this pa- per: Didja; Za-Kpota. Gekkonidae Hemidactylus ansorgei — This paper: Lama Forest (1st country record). Hemidactylus angulatus/guineensis complex — Cha- banaud (1917 a): Agouagon (as H. brookii and H. stella- tus); Grandison (1956, in part): “Dahomey”; Loveridge a ry Fig. 19. Crocodylus suchus, Pendjari NP. Photo: O. Grell Bonn zoological Bulletin 57 (1): 31-54 (1947): Ajuda; Godomey; Grand Popo; Porto Novo; Zomai; Loveridge (1952): Abomey; Bassila; Kous- soukoingou; Bauer et al. (2006): Attogon; Niaouli; Abomey; Collines du Dassa-Zoume; Manigri; Pendjari NP; this paper: Godomey; Bohicon. Hemidactylus beninensis — Bauer et al. (2006): Collines du Dassa-Zoume (new species description). Hemidactylus fasciatus — Bauer et al. (2006): near Kétou; this paper: Lama Forest (2nd country record). Hemidactylus mabouia — Bauer et al. (2006): Godomey, Abomay, Niaouli; this paper: Cotonou (2nd country record). Hemidactylus muriceus — Bauer et al. (2006): near Kétou. Hemidactylus lamaensis — This paper: Lama Forest (new species description). Lygodactylus conraui — Bauer et al. (2006): Lokoli Fo- rest. Phyllodactylidae Ptyodactylus ragazzii — Bauer et al. (2006): Chutes de Koudou, PN du W. Tarentola ephippiata — Bauer et al. (2006): Pendjari NP. ©ZFMK 48 Klaus Ullenbruch et al. Fig. 20. Agama gracilimembris, Pendjari NP, first rediscovery in Benin after its first description from this country nine decades ago. Photo: O. Grell Agamidae Agama agama complex — Loveridge (1952): Bassila; Kou- tiacou; Zagnanado; this paper: Godomey nr. Cotonou; Co- tonou; Bohicon; Pendjari NP. Agama sylvanus — This paper: Lama Forest (1st country record). Agama gracilimembris — Chabanaud (1918): “Dahomey” (original description); this paper: Pendjari NP. Note: This little agama with unresolved intrageneric re- lationships was subsequently found in Nigeria and in the Central African Republik (Grandison 1968, 1969, Joger 1990), the gap between these two records being bridged by a find in northern Cameroon (Béhme 1975). The pho- tographic record from Pendjari NP (Figure 20) is the first find of this rare agama in Benin since its original descrip- tion in 1918, a rediscovery after 9 decades! Chamaeleonidae Chamaeleo africanus — This paper: Atakora chain (1st country record, based on photograph). Note. A photograph of a “chameleon” published in a pop- ular book on Benin by Englebert (1973, cited after Con- rad 1999) without identifying the species shows clearly a specimen of C. africanus. Its inclusion by Conrad (1999) into her unpublished thesis means that it is still the first published country record for Benin. Chamaeleo gracilis — This paper: Abomey-Didja; Pend- jar1 NP. Bonn zoological Bulletin 57 (1): 31-54 Chamaeleo necasi — Ullenbruch et al. (2007), this paper: Bohicon-Za-Kpota, Houegbo. Chamaeleo senegalensis — Loveridge (1952): Zagnanado; this paper: Bohicon; Abomey-Didja; Pendjari NP. Scincidae Chalcides thierryi — Loveridge (1952): Koussokoingou (as Chalcides ocellatus thierryi). Mochlus guineensis — Chabanaud (1917): Agouagon; this paper: Lama Forest; Cotonou. Panaspis togoensis — Chabanaud (1917): Agouagon (as Lygosoma (Riopa) dahomeyense, syn. fide Schmitz et al. 2005); Loveridge (1952): Bassila (as Lygosoma (Panaspis) breviceps togoense) Grandison (1956): “Dahomey” (as P. breviceps togoense); Fuhn (1972): “Dahomey: Agouagou” (as Panaspis kitsoni); this paper: Lama Forest; Pendjari NP. Trachylepis affinis — Chabanaud (1917): Satadougou (as Mabuia raddonii); Loveridge (1952): Abomey; Bassila; Koussokoingou; Tanogou (as Mabuya blandingii); this pa- per: Godomey nr. Cotonou; Lama Forest; Pendjari NP. Trachylepis perrotetii— Chabanaud (1917): Agouagon (in part as Mabuia albilabris, fide Hoogmoed 1974); Loveridge (1952): Abomey; Bassila; Koussokoingou (as Mabuya perroteti); this paper: Godomey nr. Cotonou; Pendjari NP. Trachylepis maculilabris — Chabanaud (1917): Agouagon; Loverdige (1952): Bassila. Trachylepis quinquetaeniata sharica Sternfeld, 1917 — Chabanaud (1917): Agouagon (in part as Mabuia albi- labris, fide Hoogmoed 1974); Loveridge (1952): Bassila (as Mabuya quinquetaeniata sharica); this paper: Pend- jari NP. Gerrhosauridae Gerrhosaurus major zechi — Loveridge (1952): Kous- sokoingou. Lacertidae Acanthodactylus boueti — Chabanaud (1917): Agouagon (as A. (Latastia) boueti); Loveridge (1952), Salvador (1982): Bassila. ©OZFMK Reptiles from southern Benin, West Africa, with the description of a new Hemidactylus 49 Heliobolus nitidus — Loveridge (1952): Bassila (as Eremias n. nitida); this paper: Pendjari NP. Holaspis guentheri — This paper: Lama Forest (1st coun- try record). Amphisbaenidae Cynisca leucura — Chabanaud (1917a): Agouagon; “Da- homey” (as Amphisbaena leucura); Loveridge (1952): Natitingou. Varanidae Varanus exanthematicus — This paper: Abomey-Didja. Varanus niloticus — Chabanaud (1917a): Agouagon; “Haut Dahomey”; this paper: Godomey nr. Cotonou; Ouémé River near Togbota; Pendjari NP (photographic voucher of an unusually pale olive-coloured juvenile, Fig- ure 21). Varanus ornatus — This paper: Abomey-Didja; Lama For- est (1st country record). Leptotyphlopidae Leptotyphlops bicolor — Chabanaud (1916): Porto Novo (as Glauconia. b. gruveli). Leptotyphlops brevicauda — Bocage (1887): “Dahomey” (as Stenostoma brevicauda); Hahn (1980): “Dahomey” (as Leptotyphlops brevicaudus). *Leptotyphlops nigricans — Chabanaud (1916): “Da- homey” (Identification probably erroneus, taxon not in west Afria). Typhlopidae Ramphotyphlops braminus — Trape & Mané (2006 b): Cotonou. Typhlops punctatus — Chabanaud (1916): “Dahomey”; Chabanaud (1917a,b): Ouidah, Agouagon. Boidae Gongylophis muelleri — Trape & Mané (2006 b): on grid map but without specific locality. Bonn zoological Bulletin 57 (1): 31-54 Fig. 21. Varanus niloticus, subadult, unusally light-coloured specimen from Pendjari NP. Photo: O. Grell Python regius — Villiers (1951): “trés commun”, voucher specimen from Abomey; this paper: Godomey nr. Coto- nou; Za-Kpota; Lama Forest. Python sebae — Villiers (1951): “Trés commun partout”, specimens from Abomey; this paper: South Benin. Colubridae Afronatrix anoscopus — Chirio & Ineich (2009): Niangou NE of Tanguieta; Atakora Prov.; this paper: Lokoli For- est (2nd published country record, regarded already as probably present by Villiers 1951 and Chippaux 2001). Bamanophis dorri — Chippaux 1999, 2006; N Benin: un- specified (as Coluber dorri); Trape & Mané (2007): on grid map but without specific locality (as Hemorhhois dor- ri; Schatti & Trape (2008): unspecified. Crotaphopeltis hippocrepis — Rasmussen (2000): Soubrou- kou. Crotaphopeltis hotamboeia — Chabanaud (1916): “Daho- mey” (as Leptodira hotamboeia); Chabanaud (1917 b): Agouagon (as Leptodira hotamboeia); this paper: Godo- mey nr. Cotonou. Dasypeltis gansi — Chabanaud (1916): “Dahomey” (as D. scabra palmarum); Trape & Mané (2006 a): Lanta; this paper: Za-Kpota. Dispholidus typus — This paper: Bohicon-Didja (1st coun- try record; regarded as probably present by Villiers 1950). Grayia smythii — Chabanaud (1916): near Sakété, Porto- Novo. ©ZFMK 50 Klaus Ullenbruch et al. Hormonotus modestus — Chabanaud (1916): “Dahomey”; Chabanaud (1917 b): near Lac Azzi. Lamprophis fuliginosus — Chabanaud (1916): “Dahomey” (as Boaodon fuliginosus); Chabanaud (1917b): Agouagon (as Boodon fuliginosus); Villiers (1951): Koussokoin- gou/Atakora (as Boaedon fuliginosus); this paper: Dan Forest; Lama Forest; Didja, Za-Kpota. Lamprophis lineatus — Chabanaud (1916): “Dahomey” (as Boaodon linatus); Chabanaud (1917b): Agouagon (as Boodon lineatus); Villiers (1951): Abomey (as Boaedon lineatus). *Lycophidion capense — Chabanaud (1916): “Dahomey” (listing of capense and capense multimaculatum proba- bly due to identification error: both taxa not in West Africa). Lycophidion semicinctum — Villiers (1951): Abomey; this paper: Bohicon; Didja. Mehelya crossi — Chabanaud (1916): “Haut-Dahomey” (as Simocephalus capensis); Loveridge (1940): Bassila; Vil- liers (1951): Abomey. Meizodon coronatus — Trape & Mané (2006 b): on grid map but without specific locality. Fig. 22. Psammophis elegans from Pendjari NP, dorsal pattern. Photo: O. Grell Bonn zoological Bulletin 57 (1): 31-54 Meizodon regularis — Chippaux (2001): Benin (on map, without documented locality); this paper: Godomey nr. Cotonou (2nd country record). Natriciteres olivacea — Chabanaud (1916): “Dahomey” (as Tropidonotus olivaceus); this paper: Lama Forest (1st doc- umented locality record for Benin). Natriciteres variegata — This paper: Lama Forest (1st country record). Philothamnus irregularis — Trape & Mané: on grid map but without specific locality; this paper: Lama Forest. Philothamnus semivariegatus — Chabanaud (1916): “Da- homey”; Chabanaud (1917a): Cotonou, Agouagon; Villiers (1951): Abomey; this paper: Lama Forest. Polemon neuwiedii — Chabanaud (1916): “Dahomey” (as Miodon neuwiedi); Agouagon (as Miodon neuwiedi). Prosymna meleagris — Chabanaud (1916): “Dahomey”; Chabanaud (1917a): Agouagon. Psammophis elegans — Loveridge (1941): Ouidah; this pa- per: Cotonou; Bembéréké ZFMK 72885); Pendjari NP (Figure 22). Psammophis lineatus — Chabanaud (1916): “Dahomey”; Chabanaud (1917 b): Agouagon. Psammophis cf. phillipsi— Chabanaud (1916) “Dahomey” (as P. sibilans); this paper: Abomey; Za-Kpota; Pendjari NP (Figure 23). Psammophis phillipsi (s.str.) — Chabanaud (1916): “Da- homey” (as P. regularis); this paper: Godomey near Coto- nou. Psammophis praeornatus — Chabanaud (1916): “Da- homey” (according to Villiers 1951 not documented). Psammophis schokari — Chabanaud (1916): “Dahomey” (as P. shokari: considered to be of doubtful provenance by this author). Psammophis sudanensis — This paper: Didja; Za-Kpota, Abomey, Bohicon (1st country record). Rhamphiophis oxyrhynchus — Chirio & Ineich (1991): Benin, unspecified; Trape & Mané (2006): on grid map, but without specific locality; this paper: Godomey nr. Cotonou; Za-Kpota. ©OZFMK Reptiles from southern Benin, West Africa, with the description of a new Hemidactylus 51 Fig. 23. Psammophis cf. phillipsi from Pendjari NP, showing the characteristic reddish dorsilateral stripes. Photo: O. Grell Scaphiophis albopunctatus — Bocage (1895, 1896): Aju- da; Chabanaud (1916): “Dahomey”; Chabanaud (1917a): Agouagon; Broadley (1994): Parakou, and Segbana; this paper: Lama Forest (Didja). Telescopus variegatus — Chabanaud (1916): “Dahomey” (as Zarbophis variegatus), Chabanaud (1917a): Agouagon (as Tarbophis variegatus); Villiers (1951): Abomey (as Tarbophis variegatus). *T. semiannulatus — Chabanaud (1916): “Dahomey” (as Tarbophis semiannulatus. Repeated by Villiers (1951), but probably due to misidentification, as taxon not in present in West Africa). Thelotornis kirtlandii — Villiers (1951): Bassila. Toxicodryas blandingii — Chabanaud (1916): “Dahomey” (as Dipsadomorphus blandingii); Villiers (1951): Zag- nanado, Abomey (as Boiga blandingii). Toxicodryas pulverulentus — Chabanaud (1917b): Porto Novo (as Dipsadomorphus boueti, syn. fide Villiers 1951). Atractaspididae Amblyodipsas unicolor — Considered to be probably in Benin by Villiers (1950) and Meirte (1999); Chippaux (2001): On map, but without documented locality; this pa- per: Za-Kpota (first documented country record). Aparallactus modestus — Boettger (1898): Grand Popo; Witte & Laurent (1947): ’Gross Popo”. Atractaspis aterrima — Villiers (1951): Abomey; Ras- mussen (2005): “Benin, without locality”. Bonn zoological Bulletin 57 (1): 31-54 Atractaspis dahomeyensis — Barboza du Bocage (1887): Zomai; Villiers (1951): Abomey; Trape & Mané (2006b): Parakou. Atractaspis watsoni — Trape & Mané (2006 b): on grid map but without specific locality. Elapidae Dendroaspis viridis — Chabanaud (1916): “Dahomey” (as Dendraspis viridis); this paper: Bohicon. Elapsoidea semiannulata moebiusi — Chabanaud (1916): “Dahomey” (as Elapechis guentheri); Chabanaud (1917 b): Agouagon (as Elapechis guentheri); this paper: Lama (Didja). Naja melanoleuca — Chabanaud (1916): “Dahomey”; Vil- liers (1951): Abomey. Naja nigricollis — Chabanaud (1916): “Dahomey”; Cha- banaud (1917): Agouagon; Villiers (1951): Abomey; this paper: Lama Forest; Pendjari NP. Naja senegalensis — Trape et al. (2009): Niénié. Viperidae Bitis arietans — Chabanaud (1916): “Dahomey”; Cha- banaud (1917 b), Villiers (1951): Koussokoingou/Atako- ra, Zagnanado (as B. lachesis); this paper: Abomey; Be- mbéréké (ZFMK 72886). Bitis nasicornis — Chabanaud (1916): “Dahomey”. Causus maculatus — Chabanaud (1916): “Dahomey” (as C. rhombeatus); Chabanaud (1917b): Agouagon (as C. rhombeatus); Villiers (1951): Koussokoingou/Atakora (as C. rhombeatus); this paper: Bohicon/Didja, Abomey. Echis. ocellatus — Chabanaud (1916): “Dahomey” (as E. carinatus); Chabanaud (1917 b): Agouagon (as E. cari- natus); Villiers (1951): Koussokoingou/Atakora (as E. car- inatus); this paper: Didja; Bembéréké (ZFMK 72867, 73091); Pendjari NP. Acknowledgements. We thank Prof. Dr. Peter Nagel and Dr. Ralf Peveling, University of Basel, that they provided the op- portunity to KU to work within the BIOLAMA project. Thanks are also due to Lamidi Konetche and Thibault Lachet for some additional gecko specimens. WB thankfully acknowledges Dr. Werner Schréder, Gottingen, and Mr. Thomas & Mrs. Marianne Imthurn, Bembéréké/Benin, for some reptile from Godomey and Bembéreékeé, respectively. ©OZFMK Nn i) We further thank PD Dr. Mark-Oliver Rédel and Mr. Frank Tillack, Museum ftir Naturkunde Berlin, for the loan of relevant gecko type material, and Dr. Colin McCarthy, Natural History Museum London, and Dr. Heinz Grillitsch, Naturhistorisches Museum Vienna, for photos of further important type material of geckos and a snake. Prof. Dr. Uwe Fritz, Senckenberg naturhistorische Sammlungen Dresden, kindly commented on a chelonogical issue while Dr. Andreas Schmitz, Muséum d’His- toire Naturelle, Geneva, helpfully commented on skink taxon- omy and on the manuscript as a whole. Mr. Philipp Wagner (MSc) and Mrs. Ursula Bott (both from ZFMK Bonn) provid- ed again support with their photographic and technical skills. REFERENCES Angel F (1933, “1932”) Les serpents de l’Afrique occidentale frangaise. Bulletin du comité d’études historiques et scienti- fiques de l’ AOF15 (4): 613-858 Bauer AM, Giinther R (1991) An annotated type catalogue of the geckos (Reptilia: Gekkonidae) in the Zoological Museum, Berlin. Mitteilungen aus dem Zoologischen Museum in Ber- lin 2: 279-310 Bauer AM, Tchibozo S, Pauwels OSG, Lenglet G (2006) A re- view of the gekkotan lizards of Bénin, with the description of anew species of Hemidactylus (Squamata: Gekkonidae) Bocage JV Barboza du (1887) Mélanges erpetologiques. II. 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Tropical Zoology, Firenze 20: 1-17 Villiers A (1951) Mission A. Villiers au Togo et a Dahomey. Etu- des dahomeyennes (IFAN), 5: 1-25 ©OZFMK 54 Klaus Ullenbruch et al. Wermuth H (1965) Liste der rezenten Amphibien und Reptilien. Gekkonidae, Pygopodidae, Xantusiidae. Das Tierreich, Ber- lin 80: 1-xxii + 1-246 Werner F (1919) Wissenschaftliche Ergebnisse der mit Unter- stiitzung der Kaiserlichen Akademie der Wissenschaften in Wien aus der Erbschaft Treitl von F. Werner unternommenen zoologischen Expedition nach dem Anglo-Agyptischen Sudan (Kordofan) 1914. IV. Bearbeitung der Fische, Amphibien und Reptilien. Denkschriften der Kaiserlichen Akademie der Wis- Bonn zoological Bulletin 57 (1): 31—54 senschaften, Mathematisch-Naturwissenschaftliche Klasse 96: 437-509 Witte GF de, Laurent R (1947) Révision d’un groupe de Colu- bridae africains. Mémoires du Muséum Royal d’ Histoire Na- turelle de Belge, Bruxelles (2) 29: 1-134 Received: 19.03.2010 Accepted: 09.04.2010 Corresponding editor: F. Herder ©ZFMK Bonn zoological Bulletin Volume 57 Issue | pp. 55-64 Bonn, April 2010 Fitness components in the relationship between Rhopalapion longirostre (Olivier, 1807) (Insecta: Coleoptera: Apionidae) and Alcea rosea (Linnaeus, 1758) (Malvaceae). Analysis of infestation balance of a herbivorous weevil and its host plant Gertha Wilhelm', Hans L. Nemeschkal', John Plant? & Hannes F. Paulus? ' Faculty of Life Sciences, Department of Theoretical Biology, University of Vienna, Althanstr. 14, A-1090 Vienna, Austria; E-mails: a9404775@unet.univie.ac.at; Hans.Leo.Nemeschkal@univie.ac.at; * Faculty of Life Sciences, Department of Evolutionary Biology, University of Vienna, Althanstr. 14, A-1090 Vienna, Austria; E-mails: john.plant@aon.at; hannes.paulus@univie.ac.at Abstract. Multivariate statistics (principal components analysis, path analysis) were used to investigate fitness compo- nents of the interactions between the weevil Rhopalapion longirostre (Olivier, 1807), Apionidae, Coleoptera and its host plant Alcea rosea (Linnaeus, 1758), Malvaceae. We focused on the activities of the larvae such as the choice of seeds for consumption, the preparation of seed chambers as a site for pupation, as well as the construction of escape holes through which the adults later emerge. The analyses revealed that the optimal conditions for successful development of the weevils depended on the availabil- ity of seed capsules characterized by a high number of well developed seeds, few undeveloped and few spoiled seeds. The high number of larvae, pupae and not emerged adults found in the seed capsules corresponds with the successful emergence of adults. Egg deposition by the females in appropriate flower buds of the host plant, together with larval con- tribution to overall reproductive success are important fitness components in the life cycle of the weevil. The reproduc- tive success of the plant is partly guaranteed by a high number of seed chambers, many well developed seeds, many not infested seeds, and few undeveloped seeds. Furthermore, the extended blooming period of the host plant serves as an es- cape strategy by developing buds early or late in the season that cannot become entangled in the life cycle of R. lon- girostre. Key words. Alcea rosea, Malvaceae, Rhopalapion longirostre, Apionidae; Host plant-parasite interaction, infestation bal- ance, larval contribution, path analysis, PCA, reproductive success INTRODUCTION Life history theory provides a framework for analyzing the contribution of major features of the life cycle to re- productive fitness and seeks to place these features in an evolutionary context (Roff 1992; Stearns 1992; Stearns & Hoekstra 2000). The life history events of most of the 62,000 described species of weevils or snout beetles (Cur- culionoidea, Coleoptera) (Alonso-Zarazaga & Lyal 1999), and their immature stages are incompletely documented and more or less episodic. As holometabolous insects, weevils undergo a complete metamorphosis which pro- duces larval and adult stages each with their own life habits and physiological and behavioral traits thus enhanc- ing the possibility for adaptation, specialization and evo- lution. The phenomenal diversity of weevils was discussed recently in evolutionary terms by Oberprieler et al. (2007). Considered as a group, weevils utilize nearly all parts of host plants from almost all plant taxa (Anderson 1995). The larvae of most Apionidae, with about 1800 described species (Kuschel 1995), feed internally in stems, leaves, buds, galls, fruits or seeds of their specific host plants. The evolution of larval habits and host plant associations are taken for analysis of larval role in weevil’s diversification (Marvaldi et al. 2002). In this paper we examine several fitness components of the weevil Rhopalapion longirostre (Olivier, 1807) which is monophagous on its host plant Alcea rosea (Linnaeus, 1758), with special focus on larval contribution to repro- ductive success (Reavey & Lawton 1991). Seeds of A. 56 Gertha Wilhelm et al. rosea, presumably derived from Anatolia (Hammer 1994), were brought to France in the 18 century. The beauty and popularity of this garden plant, known as the hollyhock, passe-rose, rose trémiére, or Stockrose, was lovingly recorded in verse by King Ludwig XVI of France in the 18‘ century: L- Amour de moi s’y est éclosé / dedans un petit jardinet, / oi croit la rose et le muguet / et aussi bien la passe-rose. Today, the plant occurs in many parts of Europe, North and South America, mainly due to its cultivation in gar- dens. The development of the larvae of R. longirostre takes place in the buds and seeds of A. rosea and the expansion of the weevil’s distribution range has proceeded in inti- mate association with that of the host plant (Perrin 1995; Gonget 1997). Females of R. longirostre have a body size from 2.4 to 3.4 mm excluding the rostrum (Freude et al. 1983). They em- ploy their extremely elongated rostrum (average size 2.2 mm, standard deviation +/- 0.2 mm) (Wilhelm 2004), to bore holes into developing buds of A. rosea. The plant at- tains a height of 1 to 3 meters and can persist for more than two years. The buds appear in a long inflorescence composed of 30 to 50 buds. Every bud is surrounded by thick inner and outer sepals which have stiff grey-green hairs (Adler et al. 1994). After, or even during copulation, with the male sitting on her back, the female bores a chan- nel into a selected bud using her rostrum. She turns around and inserts her ovipositor into the bore channel penetrat- ing the outer and inner sepals of the plant to lay three to four eggs into it. Female weevils prefer to bore into buds located in the middle of the inflorescence. These buds will usually not bloom in the following 2 weeks thus allow- ing enough time for the development of the larvae. If a female is among the first to bore a hole into the bud, she chooses the side of the plant which receives the morning sunlight. If several females have oviposited in the same bud, the boring channels are widely spaced from each oth- er, presumably to avoid competition among the larvae for food. The first instar larvae are legless, as most of wee- vils larvae have lost the development of legs (Crowson 1955; Stehr 1991; Marvaldi 1997; Farrell 1998), about 0.4 mm long, and they hatch after 3 to 4 days. If the bore chan- nel is in the upper part of the bud then the larvae of the first stage consume the surrounding pollen grains. How- ever, if the bore channel is located at the bottom side of the bud, then the larvae feed only on the soft tissue of the bud. All larvae bore feeding channels toward the direc- tion of the ovules in the developing seed capsule, using their strong mandibles to bite and chew. By the time the bud has blossomed and before it falls off, all larvae must have migrated below the ovary. Within a few days the up- per part of the blossom wilts and falls to the ground. When the seeds of A. rosea are fully developed, each larva tries Bonn zoological Bulletin 57 (1): 55-64 to enter a seed chamber from the bottom of the seed cap- sule and consume the entire contents of the seed from within. Prior to pupation, the last instar bores a hole on the broadest side of the seed chamber to later escape when it reaches the adult stage. The escape hole is filled in with a secretion produced by the larva to protect the pupa and the developing weevil against outside influences, e.g. wet- ness and sunlight. Pupation lasts between 2 to 3 weeks de- pending on weather conditions. The darkening of the adult weevil’s exoskeleton is visible through the transparent membrane of the pupa beginning at the apex of the ros- trum and the elytra. The adult beetle slowly emerges from the dry and stiff seed chamber over a period of several hours using its rostrum to get out through the escape hole. A duration of 8 to 10 weeks is required from egg deposi- tion to the emergence of adult weevils. Additional details on the biology of R. /ongirostre are found in Dieckmann (1977), Behne (1998), Pupier (1997), Sprick et al. (2002) and Wilhelm (2004). The interaction between the host plant and the weevil is reflected in a successful joint expansion in Europe over the last twenty years. To demonstrate this a null hypoth- esis was postulated which denied significant interactions between the life cycles of host plant and weevil. To test null hypothesis in detail, we posed the following ques- tions: (1) What parameters lead to optimal prerequisites for suc- cessful development of weevils in seed capsules? In par- ticular, what are the conditions of capsules which contain larvae, pupae and immature adults? Do these conditions correspond with successful emergence of adults? (2) Which counter strategies are taken by the host plant to survive or to ensure its successful growth and repro- duction? (3) How effective is larval contribution to the overall re- productive success of the weevil? MATERIALS AND METHODS To address these questions we examined dry seed capsules collected from 20 Alcea rosea plants growing on the ter- race at the University of Vienna, Althanstrasse 14, in the first week of Nov. 2006. For measurements we examined only dry capsules (n=205) having a complete set of seed chambers. Seed capsules, which were incomplete because some seed chambers had already fallen to the ground, were not investigated. The plants, 1-20, contained the follow- ing number of seed capsules: 7, 10, 11, 4, 5, 6, 21, 4, 16, oh, (UNS 3}, 24 IS, 13), 7, OO, O ©ZFMK Fitness components in the relationship between Rhopalapion longirostre and Alcea rosea 57 infested bud host plant ie seed capsule (whole) seed capsule (schema) a eS yt adult weevil Fig. 1. Brief outline of life history stages of the weevil Rhopalapion longirostre on its host plant, Alcea rosea. Schema showing various conditions of a seed capsule: seed chambers with 1) fully developed seeds, 2) seeds infested with larvae, 3) pupae and clo- sed escape holes, 4) not emerged adults and closed escape holes, 5) escape holes, 6) spoiled seeds, 7) undeveloped seeds. Abbre- viations of infested bud: b bore channel, 0 ovules, p pollen, s sepal; sc seed chamber. The diameter of each capsule was measured 5 times to cancel out error. The number of dry seeds per capsule and the condition of every capsule were documented. The seed chambers were opened under a dissecting microscope to determine the inner condition. The infested condition was indicated by the presence of larvae, pupae or not emerged adults, or empty seed chambers with open escape holes. The uninfested condition was indicated by the presence of seed chambers which are fully developed, undeveloped or spoiled. In the following, the data were analyzed thor- oughly by multivariate statistics to estimate the importance of animal and plant related factors and their interactions on the reproductive fitness of weevil and plant. For the following case study nine character variables of the seed capsules were measured for the statistical analy- ses. Diameter of capsules in mm: diam (plant related: PR) Number of seed chambers per capsule: nrse (PR) Bonn zoological Bulletin 57 (1): 55-64 Number of open beetle escape holes per capsule: nreh (an- imal related: AR), i.e. estimate of the number of beetles which have successfully escaped. Number of dead larvae per capsule: nrla (AR) Number of dead pupae per capsule: nrpu (AR) Number of dead, not emerged adults per capsule: nrne (AR) Number of fully developed seeds per capsule: nrds (PR) Number of undeveloped seeds per capsule: nrus (PR) Number of spoiled seeds per capsule: nrss (PR) In preparation of multivariate analyses, the original data were transformed as follows: The first variate was loga- rithmically transformed (natural log). In order to fulfill the ©ZFMK 58 Gertha Wilhelm et al. Table 1. Correlation matrix of transformed variates and significant levels: *** = highest significant P-value < 0.001, ** = highly significant P-value < 0.01, * = significant P-value < 0.05. Bonferroni corrected. Variates: diam diameter of seed capsule, nrse number of seed capsules, nreh number of escape holes, nrla number of larvae, nrpu number of pupae, nrne number of not emer- ged adults, nrds number of fully developed seeds, nrus number of undeveloped seeds, nrss number of spoiled seeds. The signi- ficance of correlation coefficients was determined by a random permutational test (Nemeschkal 1999). diam nrsc nreh nrla nrpu nrne nrds nrus nrss diam 1.0000 nrse 0.075632 1.0000 nreh 0.057043 0.167632* 1.0000 nrla 0.032532 0.293509*** - 0.135968 1.0000 nrpu 0.013449 0.2803 19*** - 0.044457 0.538041 *** 1.0000 nme = - 0.07478 0.129488 0.128997 0.161018 0.212016*** 1.0000 nrds 0.036070 - 0.046134 - 0.247253%** — - 0.304999%** = - 0.31991*** - 2.242189** 1.0000 nrus 0.000937 0.163294 - 0.257581*** — - 0.028526 - 0.088356 - 0.046297 — - 0.306250*** 1.0000 nrss - 0.001917 0.254107*** - 0.26796 1*** 0.096727 0.208869** 0.088184 - 0.402289*** 0.165270* 1.0000 Table 2. Factor matrix, axis | to 7, and significant axes of loadings. *** = highest significant, P-value < 0.001, ** = highly si- gnificant P-value < 0.01, * = significant P-value < 0.05, ns= no significance P-value > 0.05, eigenvalues, and percentages of the total variance. The significant levels were determined by a random permutational test (NEMESCHKAL 1999, 10000 iterations). Potential win for animal and plant. + A, + P; potential loss: - A, - P. Abbreviations of axes: SUPLA for ‘successful plant (seed) production’; SUAN for ‘successful animal production’; SUMIX for ‘integration of the two trends, successful animal and plant production’; LIFA for ‘late infestation axis’ and DASA for ‘damaged seed and animal axis’. Factor axes Fl F2 F3 F4 F5 F6 F7 Abbreviations SUPLA SUAN SUMIX LIFA DASA nn. nn. var 3 nreh - 0.083229ns_ - 0.72391 6*** 0.568846*** - 0.237805** 0.174577ns_ - 0.024872ns 0.2403 88*«« var 4 nrla 0.680778 - 0.168066« - 0.46383 1** - 0.297076*** -0.174923ns 0.394877*** 0.135561ns var 5 nrpu 0.726492*** - 0.245330 — - 0.380804 - 0.114225ns 0.074140ns_ - 0.498353*** 0.010510ns var 6 nrne 0.430808*** — - 0.379749 +«« O.211681** 0.631854 - 0.475213*** 0.012534ns 0.012746ns var 7 nrds - 0.743359%* — 0.012772ns - 0.532765*** 0.217806** -0.116773ns -0.103023ns 0.302864++« var 8 nrus 0.217084+« 0.687623 **« 0.397469» - 0.274868** - 0.443069 -0.147826ns 0.168382ns var9 nrss 0.559095*** 0.45827 1 ««« 0.118490ns 0.384199 0.525592** 0.073805ns 0.184544 +« Eigenvalues 2.096067 1.439697 1.184405 0.827693 0.778592 0.442973 0.230572 percentage of 29.94 20.57 16.92 11.82 11.12 6.33 3.29 total variance animal win/loss -A +A +A +A -A Plant win/loss +P -P +P +P -P linearity requirement, variates 2 to 9 were substituted by their square roots. Finally, a product moment correlation matrix was calculated between all variates over the 205 capsules (Table 1). The correlation structure of variates 3 to 9 was then ana- lyzed by a principal components analysis (PCA, for de- tails see Morrison 2004). An orthogonalization was reached via PCA by calculating a full analysis and extract- ing all possible, i.e. 7, axes (Stat. Graphics — Version 7.3 for MS DOS). The eigenvectors were scaled according to Bonn zoological Bulletin 57 (1): 55—64 eigenvalues leading to a factor matrix (Table 2). The sig- nificance of loadings on principle axes was determined by a random permutational test, 10000 random permutations each, (Nemeschkal 1999). Note that the original variates are represented on factor axes by their loadings. A load- ing is the product moment correlation of an original vari- ate with the factor axis. For details and further statistical explanation, see Morrison (2004). Path analysis, a special type of regression analysis, (Stat. Graphics — Version 7.3 for MS DOS) was applied to rep- ©ZFMK Fitness components in the relationship between Rhopalapion longirostre and Alcea rosea 59 resent the complicated animal-plant relations in diagram- matic form (Path diagrams, Figs. 2 and 3). In addition, path analysis has the advantageous option to focus dependen- cies either on plant or for animal variables (for theoreti- cal details on path analysis, see Sokal & Rohlf (1995). For reasons of simplicity, our analyses are mainly based on procedures of the general linear model. In order to test whether a loss of information in the data structure has oc- curred due to data transformations, we calculated, in par- allel log-linear analyses with the non-transformed data (see Appendix). RESULTS Of the 205 examined seed capsules, each contained an av- erage of 35 individual seed chambers (ll 34.7758, ul 35.6798; ll: lower limit, ul: upper limit of 95% confidence of average). Of the 35 seeds chambers, an average of 13 (Il 11.6393, ul 13.4734) were fully developed, inferring the reproductive success of the host plant, 7 (11 6.0002, ul 7.6396) had open escape holes, inferring the potential re- productive success of the weevils. The remaining 15 seed chambers (on average per capsule) were either infested (documented by dead larvae, pupae or not emerged adults), undeveloped or spoiled by mold or mites. The results from the PCA and the path analyses reveal de- tailed information on the complex system of interactions between weevil and host plant. For the sake of simplifi- cation, the diameter and number of seed chambers were omitted from the PCA and path analyses since the simple correlation between the diameter (diam) and the remain- ing variates are not significant (Table 1) and since the number of seed chambers (nrsc) is a summation of vari- ates 3 to 9. A total of 7 axes are significant (Table 2). Factor axes | to 5 are the most important, accounting for over 90% of the total variance. Results summarized for the five most important factor ax- es: F1 represents nearly 30% of total variance. Positive and negative signs of the loadings indicate the reverse propor- tionality in the variational trends: nrla, nrpu, nrne, nrss ( positive and very highly significant) and nrus (highly sig- nificant) versus nrds (negative and very highly significant). F2 represents 21% of total variance. Cumulatively, the ax- es F1 and F2 explain half of the total variance. The load- ings are contrasted to each other by their signs (+/-); i.e. nrus and nrss (very highly significant) and with positive Bonn zoological Bulletin 57 (1): 55-64 signs, versus nreh, nrpu, nrne (very highly significant) and nrla (significant), both with negative signs. F3 explains 17% of the total variance, and by accumula- tion of axis Fl to F3 more than two thirds of total vari- ance are explained. The signs of loadings, nreh, nrne and nrus (very highly significant) contrast with those of nrla and nrpu (very highly significant). F4 represents about 12% of total variance. Axes F1 to F4 cumulatively explain more than three quarters of total vari- ance. The loadings are grouped by positive signs for nrne, nrss (very highly significant) and nrds (highly significant) and negative signs for nrla, nrus (very highly significant) and nreh (highly significant). F5 explains 11% of total variance. The positive sign of the loading nrss (very highly significant) contrasts with those of nrne and nrus (very highly significant). Factor axes 6 and 7 together provide 9% of total infor- mation and will not be further considered in this study. Path Analyses Two path analyses were conducted to focus on the main influences affecting the animal’s or plant’s life cycle. Vari- ables 3 to 9 served as the basis for both path analyses. The first analysis determines the influence of variables 4 to 9 on variable 3 (nreh: number of escape holes per capsule, AR = animal related) (Fig. 2). In the second analysis vari- able 7 (nrds: number of fully developed seeds per capsule, PR = plant related) the variables 3 to 6, 8 and 9 take in- fluence on variable 7 (Fig. 3). Path Analysis I (Fig. 2): Since variable 3, nreh, number of escape holes per capsule, may be an indicator of fit- ness for the weevil, it was subjected to a multivariate path analysis. Simple arrows indicate the directions of influ- ences of the predictors on criterion (= direct paths), dou- ble arrows indicate the correlation between predictor vari- ables. The strength of influence is given by the value of the path coefficient. The path coefficient is a standardized multiple regression coefficient (path coefficient = regres- sion coefficient times standard deviation of predictor di- vided by the standard deviation of criterion). The path co- efficient quantifies the amount of change in criterion when progressing in independent variables. There are 4 direct paths from the predictors to the criterion. The first direct path proceeds from var 4, nrla, to the criterion and is neg- ative and highly significant. It infers that the more dead larvae per capsule, the fewer escape holes. The second di- rect path from var 7, nrds, to the criterion is negative and very highly significant: the more well developed seeds, the fewer open escape holes. The third direct path proceeds OZFMK 60 Gertha Wilhelm et al. ky 0.5380 I ee OFS 19310 ** 0.208869 * 0.165270 *** _0 306250 Variables nreh number of escape holes nrla number of larvae nrpu number of pupae nrne number of not emerged adults nrds number of fully developed seeds nrus number of undeveloped seeds nrss number of spoiled seeds Fig. 2. Path diagram | showing 6 predictor variables and a residual effecting one criterion (var 3, nreh, number of escape holes, an animal related variate), 2= ().414525. The path diagram explains 41.45% of the variance of the criterion. Only significant paths are shown. The significance of paths was determined by random permutational test; 10000 iterations each; Bonferroni Correction. Simple arrows indicate the direction of influences of the predictors to the criterion (= direct paths), double arrows indicate the cor- relation between predictor variables. The strength of influence is given by the value of the path coefficient. from var 8, nrus, to the criterion and is negative and very highly significant: the more undeveloped seeds, the few- er escape holes. The fourth direct path proceeds from var 9, nrss, to the criterion and is negative and very highly sig- nificant: the more spoiled seeds per capsule, the fewer open escape holes. In addition, several indirect effects are apparent. Both direct and indirect paths in the diagram must be combined to indicate an overall influence on the criterion. The most important indirect path proceeds from nrla to the criterion via nrds, and is very highly signifi- cant: the more larvae, the fewer well developed seeds and the more escape holes (indirect effect: -0.304999 « -0.624623 = 0.190509). Direct and indirect paths are mu- tually counteracting, therefore it is the summation of all effects that indicates the overall influence on the criteri- on, which in this case is negative: direct effect: nrla > nreh -0.277150 plus indirect effect: nrla > nrds > nreh 0.190509 results in an overall effect of -0.086641. A sec- ond indirect effect proceeds from nrla to the criterion via Bonn zoological Bulletin 57 (1): 55-64 nrpu, nrne and nrds: the more larvae, the more pupae and the more not emerged adults means that there are fewer well developed seeds and more escape holes in the seed capsule (0.538041 * 0.212016 * -0.242189 * -0.624623 = 0.017257). Another indirect effect between nrpu and nrds to the criterion is very highly significant: the more pupae, the fewer well developed seeds and the more es- cape holes (-0.31931* -0.624623 = 0.19945). The indirect effect which proceeds from nrds and nrus to the criterion is very highly significant: the more developed seeds, the fewer undeveloped seeds and the more escape holes (- 0.30625) * -0.389241 = 0.11921. The indirect effect which proceeds from nrds and nrss to the criterion is very high- ly significant: the more developed seeds, the fewer spoiled seeds and the more escape holes (-0.402289 * -0.421838 = (0.1697). Overall, the indirect effects indicate a decrease of the negative influence from the direct paths to the cri- terion. ©ZFMK Fitness components in the relationship between Rhopalapion longirostre and Alcea rosea 61 *** -0.267961 Variables nreh number of escape holes nrla number of larvae nrpu number of pupae nrne number of not emerged adults nrds number of fully developed seeds. nrus number of undeveloped seeds nrss number of spoiled seeds Fig. 3. Path diagram 2 showing 6 predictor variables and a residual effecting on criterion (var 7, nrds, plant related variate, num- ber of fully developed seeds per seed capsule), R? = 0.545329. The path diagram explains 54.53% of the variance of the criterion. The significance of paths was determined by random permutational test, 10000 iterations each, Bonferroni Correction. Path Analysis II (Fig. 3): The second path analysis, using the same variables as in the first analysis, was calculated to determine the influence of variables 3—6, 8 and 9 on variable 7, nrds, number of fully developed seeds per cap- sule, which is the most important plant related factor. Since variable 7 may be an indicator of fitness for the plant, it was subjected to a multivariate path analysis. There are 4 direct paths from the predictors to the crite- rion. The first direct path proceeds from var 3, nreh, to the criterion and is negative and very highly significant: the more escape holes, the less fully developed seeds per capsule. The second direct path from var 4, nrla, to the criterion is also negative and very highly significant: the more larvae, the fewer well developed seeds. The third di- rect path, from var 8, nrus, to the criterion is negative and very highly significant: the more undeveloped seeds, the fewer developed seeds per capsule. The fourth direct path, from var 9, nrss, to the criterion is also negative and very highly significant: the more spoiled seeds, the fewer well developed seeds. Bonn zoological Bulletin 57 (1): 55—64 There are several indirect effects between the predictors and the criterion. The indirect effect proceeds from var 3 and var 8 to the criterion and is very highly significant: the more escape holes, the less undeveloped seeds and the more fully developed seeds (-0.257581 * -0.387095 = 0.09971). The indirect effect between nreh and the crite- rion, via nrss, is very highly significant: the more escape holes, the fewer spoiled seeds and the more fully devel- oped seeds (-0.267961 * -0.407705 = 0.10925). Both paths show a decrease of the negative influence of escape holes on the criterion. Another indirect effect from nrne, via nr- pu, via nrla to the criterion is very highly significant: the more not emerged adults, the more pupae and the more larvae, the fewer well developed seeds will be found (0.212016 * 0.538041 * -0.256493 = -029259). This indi- rect effect shows a negative influence on the criterion. In summary, both the PCA and the path analyses explain animal - plant interactions; path diagram | focuses on the animal, while path diagram 2 centers on the plant. OZFMK 62 Gertha Wilhelm et al. DISCUSSION The relationships between insects and plants are of utmost importance for terrestrial global ecology. Many of these relationships involve specialized herbivorous insects and their host plants on which the females oviposit and on which the larvae feed (Schoonhoven et al. 1998). The choice of an oviposition site is instrumental for the fitness of many weevils (Messina 2004). In a case study approach we investigated fitness components of the interactions be- tween the weevil, R. longirostre and its host plant, A. rosea. The life cycles of the weevil and its host plant must be considered in connection with each other. With the help of multivariate statistics it was possible to isolate sever- al main factors which contribute toward the survival of the host plant, as well as the successful development of the weevil. The lengthy inflorescence of A. rosea may con- tain of more than 30 buds in different development stages ranging from fully developed buds at the bottom, which will blossom soon, to immature tiny buds at the top. Host plant preferences of the female weevils for oviposition sites are buds located in the middle of the inflorescence. Exactly these offsetting development stages of the buds are reflected in the PCA (Table 2). In summary, the PCA provides an overview of the interaction system between the plant and the weevil. Factor axis 1, which is the most important one, is inter- preted as the axis which reflects successful seed develop- ment: The variate of final production outcome in the plant, i.e. nrds, number of developed seeds, is contrasted to the counteracting variables of numbers of undeveloped seeds, nrus, and spoiled seeds, nrss, and indicators of weevil de- velopment, nrla, nrpu and nrne. The axis is to be read as follows: the more developed seeds, the less dead larvae, pupae and not emerged adults in the capsules but also the less undeveloped and spoiled seeds. In contrast to this in- terpretation, axis | could be read as: the more dead lar- vae, pupae and not emerged adults inside the capsules and the more undeveloped and spoiled seeds, the less devel- oped seeds. Note that the variable of plant final success, nrds, is exclusively represented on axis | but not the vari- able of the animal final success, nreh. Consequently, we assigned to axis | the abbreviation SUPLA for “success- ful plant” (i.e. seed) production. The next most important axis, factor axis 2, is to be in- terpreted as mainly reflecting weevil development and pro- duction. The variate of final production outcome in the an- imal, i.e. nreh, number of escape holes, is combined in variation with the variates of animal development, i.e. nr- la, nrpu, and nrne. Whereas the first variate, nreh, signi- fies production success, the remaining variates nrla, nrpu Bonn zoological Bulletin 57 (1): 55—64 and nrne stand for the unsuccessful weevil development in the host plant. However, all the animal related variates are contrasted to plant variates which stand for unsuccess- ful seed development, i.e. nrus and nrss — undeveloped and spoiled seeds. The axis loading read as: The less undevel- oped and spoiled seeds, the more escape holes but also the more dead larvae, pupae and not emerged adults. Note that the variate of final production outcome in the plant, nrds, is not significantly represented on this axis. Consequent- ly, we designate this axis SUAN, “successful animal” pro- duction, as a main infestation axis. Highly infested cap- sules were located in the middle of the plant’s inflores- cence. Early infestation of seed capsules guarantees high reproductive success for the weevil as shown by the high proportion of open escape holes. In cases where a high number of seeds were infested, the number of undevel- oped and spoiled seeds was very low. Factor axis 3 reflects the mutual interaction of weevils and plant production. In one group of variates the final pro- duction outcome of the animal, nreh, and the pre-devel- opmental stage to that final outcome, i.e. nrne, are com- bined with the number of undeveloped seeds. In contrast, in the second group of variates, the final production out- come of the plant, nrds, is combined with the developmen- tal variates of the weevil, nrla and nrpu. The axis load- ings can be read in two different ways, both immediate- ly indicating a competitive race between animal and plant to maximize their fitness (as shown for a seed predatory weevil and its host plant, Tojyu, Sota 2006). On the one hand, the more escape holes, as well as not emerged adults and undeveloped seeds in capsules, infer fewer developed seeds, as well as larvae and pupae. On the other hand, the more developed seeds, as well as larvae and pupae, infer fewer escape holes, not emerged adults and undeveloped seeds. There are two counter acting trends: One, early in- festation by the weevil which is characterized by a high number of open escape holes and not emerged adults to- gether with a high number of undeveloped seeds. Since larvae only infest well developed seeds, few seeds remain for plant development. The second trend concerns late in- festation and incomplete weevil development. It is char- acterized by a high number of larvae and pupae together with a high number of developed seeds. The abbreviation SUMIX (successful animal and plant production) repre- sents the integration of the two trends. Factor axis 4 can be interpreted as late infestation. Both variates, nrds and nreh, are highly significant and contrast each other. In the one group of variates, nrds is combined with nrss and nrne. In another group, nreh is combined with nrla and nrus. The variate loadings are interpreted as the following: many developed seeds and many spoiled seeds infer few undeveloped seeds in capsules, while the presence of fewer escape holes and fewer larvae infer ©ZFMK Fitness components in the relationship between Rhopalapion longirostre and Alcea rosea 63 fon) more not emerged adults. A high number of spoiled seeds indicates that adverse weather conditions, mostly in late summer, could have influenced negatively both seed and weevil development, and in particular, the escape of adult weevils. We abbreviate this axis LIFA, “late infestation axis”. Factor axis 5 is an axis representing damaged seeds and late developmental stages of the weevils. The variate, nrss, contrasts with nrus and nrne. The loadings are interpret- ed as follows: The presence of many spoiled seeds infers fewer undeveloped seeds and fewer not emerged adults, and vice versa. This axis is termed DASA, “damaged seed and animal axis”. The main important axes of the PCA provide a compre- hensive explanation about the complex system of inter- action between animal and plant. The influences and trends of both path diagrams must be seen in the light of direct paths and indirect effects (see Figs. 2 and 3). Our null-hypothesis was falsified by the significant correspondence between the life cycle of host plant and weevil. In path diagram 1, all direct paths (with- out exceptions) from the different variates to the criteri- on have strong negative influences. The sequence of in- direct effects between the number of larvae > number of well developed seeds > criterion, as well as the indirect influence nrla > nrpu > nrne > nrds > criterion, nreh, are very important. As larval feeding habits are highly con- servative some of these habits are apparently irreversible, e.g. feeding on seeds (Marvaldi et al. 2002). Larvae of the last developmental stage choose well developed seeds to enter, to consume and to prepare the seed chamber for pu- pation and future escape. The sequence of indirect effects, nrds > nrus > criterion, or nrds > nrss > criterion, are also very important. For the performance of a larva, it is vital to find a well developed seed instead of an undevel- oped or spoiled seed. In path diagram 2, the 4 direct paths (without exceptions) have also strong negative influences on the criterion nrds. The indirect effects are advantageous for both, plant and animal production. The sequence, nrne > nrpu > nrla > criterion, indicates that the high number of infested seeds are based on a high number of formerly well developed seeds. The sequences, nreh > nrus > criterion, nrds, and nreh > nrss ~ nrds, infer that high numbers of escape holes and few undeveloped or spoiled seeds increase the number of well developed seeds. A plant in a healthy con- dition guarantees the performance of the weevils. The time of infestation is crucial for the weevils. Our analyses re- veal that an infestation occurring late in the season in- volves a high likelihood of poor plant condition, as shown in the indirect effect nrpu > nrss > nreh. Bonn zoological Bulletin 57 (1): 55—64 In conclusion, we address the initial questions posed in the introduction. 1) What are the optimal conditions for the reproduction of weevils? The most important optimal condition for wee- vil reproduction is the timely presence of many well de- veloped seeds. 2) What are the optimal conditions for the successful de- velopment of plants? The optimal conditions for the suc- cessful development of plants are few undeveloped and few spoiled seeds. 3) Is the contribution of larvae to the overall reproductive success of the weevils important? Yes, in both path diagrams, the animal related variate, 1.e. number of larvae, has a highly significant direct path to the first criterion, number of escape holes, as well as to the second criterion, well developed seeds. Obviously lar- vae benefit from capsules with a very high number of well developed seeds. Since the number of larvae has an ex- clusively direct impact on both criteria, this influence is interpreted as a high larval contribution to the reproduc- tive success of R. longirostre. Our results strongly support the view of an intricate rela- tionship between the life cycles of A. rosea and R. lon- girostre. The plants’ period of bud development lasts from April to October. The weevils utilize a window within this range to fulfill their development, beginning in May af- ter hibernation in the soil and ending in September with the completion of their life cycle and the emergence of the next adult generation from the seed chambers. The time periods before and after weevil activity ensure enough op- portunity for successful production of seeds. 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(1991) Immature Insects, Kendall/Hunt, Dubugue, Iowa Toju H, Sota T (2006) Adaptive divergence of scaling relation- ships mediates the arms race between a weevil and its host plant. Biology Letters 2: 539-542 Wilhelm G (2004) Die Lebensgeschichte von Rhopalapien lon- girostre (Olivier). Diploma dissertation, Department of Evo- lutionary Biology, University of Vienna, 105 pp Volume 2. APPENDIX Poisson regression analysis Dependent variate: nreh Regression coefficients (predictors): beta- 0 (constant): 4.632211 < 0.0001 beta- | (nrla): -0.170142 0.0002 *** beta- 2 (nrpu): -0.040228 0.4612 ns. beta- 3 (nrne): 0.040783 0.5624 ns. beta- 4 (nrds): -0.405291 < 0.0001 *** beta- 5 (nrus): -0.267374 < 0.0001 *** beta- 6 (nrss): -0.258430 < 0.0001 *** global model, deviance: 530.341739, P-value = < 0.0001, *** The significance was tested by random permutational tests (10000 iterations each). Dependent variate: nrds Regression coefficients (predictors): beta- 0 (constant): 3.797212 < 0.0001 beta- 1 (nreh): -0.166088 < 0.0001 *** beta- 2 (nrla): -0.098854 0.0072 ** beta- 3 (nrpu): -0.061230 0.1904 ns. beta- 4 (nrne): -0.079595 0.1760 ns. beta- 5 (nrus): -0.158951 <0.0001 *** beta- 6 (nrss): -0.170892 <0.0001 *** global model, deviance: 340.101076, P-value = < 0.0001 *** The significance was tested by random permutational tests (10000 iterations each). Received: 10.10.2008 Accepted: 25.02.2009 Corresponding editor: M. Schmitt ©ZFMK Bonn zoological Bulletin | Volume 57 | Issue | | pp. 65-73 Bonn, April 2010 Life-history of Gnopharmia kasrunensis Wehrli, 1939 and G. colchidaria Lederer, 1870 (Geometridae, Ennominae) and their distribution in Iran, with first host-plant records for the genus Hossein Rajaei Shoorcheh Zoologisches Forschungsmuseum Alexander Koenig, Adenauerallee 160, 53113 Bonn, Germany; E-mail: rajaei@daad-alumni.de Abstract. For the first time, observations on the life-history of the little known species Gnopharmia kasrunensis Wehrli, 1939 and G. colchidaria Lederer, 1870 are reported; eggs, larvae, pupae, adults and habitats are illustrated. Suitable dif- ferential features for a correct identification of both Iranian Gnopharmia species as well as a distribution map are pro- vided. Zusammenfassung. Erstmals wird tiber Beobachtungen zur Biologie der wenig bekannten Arten Gnopharmia kasrunen- sis Wehrli, 1939 und G. colchidaria Lederer, 1870 berichtet. Eier, Larven, Puppen, Imagines und ihre Lebensraéume wer- den beschrieben. Fiir die korrekte Bestimmung der beiden iranischen Gnopharmia-Arten werden geeignete differential- diagnostische Merkmale sowie eine Verbreitungskarte vorgelegt. Key words. Gnopharmia kasrunensis, colchidaria, biology, distribution, Iran INTRODUCTION The genus Gnopharmia Staudinger, 1892 belongs to the tribe Macariini (Ennominae) (Wehrli, 1953: 565; Scoble & Kriiger 2002) and consists of several species which are difficult to determine by external means only. Parsons et al. (1999) list thirteen species in this genus. At present, Gnopharmia is in the process of taxonomic revision by the author. By discovery of one new species and five syn- onymies, the number of valid species in the genus has been reduced to nine. All species occur in the Middle East and Asia Minor, their geographic range extending from Turkey to Pakistan. Five of the species occur in Iran. Our present knowledge on the biology and distribution of Gnopharmia species is very poor, partly due to difficul- ties of their correct identification. In the original descrip- tions of species in this genus (e.g. Ebert 1965, Erschoff 1874, Lederer 1870, Staudinger 1892, Wehrli 1938, Wehrli 1939, Wehrli 1941, Wiltshire 1967, Wiltshire 1970) no host plant records or notes about their biology are given. Prout (1915) listed three Gnopharmia species and six sub- species, and Wehrli (1953) mentioned seven species (and 11 subspecies). Both described the external features and Wehrli also characters of their genitalia, but they did not mention host plants or larval stages. The host-plant index published by The Natural History Museum, London (HOSTS, 2007), also does not record any host plant for species of the genus Gnopharmia. After a number of unsuccessful breeding experiments with Prunus spp. and other Rosaceae species by various lepi- dopterists, the first correct suggestion about the host plant of Gnopharmia was made by Bernd Miller (Berlin; pers. comm.), who assumed the wild Almond which occurs abundant in the habitats where Gnopharmia species oc- curs (Prunus, subgenus Amygdalus; Rosaceae). The first breeding of a Gnopharmia was made by Robert Trusch (Karlsruhe; pers. comm.) in 2008 with material from East- Iran (Prov. Ostan-e Khorasan, Birjand, Kuh-e Mirza Arab, 2.040 m a.s.1); larvae were fed with leafs of wild Almond shrubs from the field in Iran, but the experiment had to be finished because of the lack plant material in Germany; the species of the reared larvae remained undetermined During May and July 2009, the author collected females of G. kasrunensis (Fig. 13) and G. colchidaria (Fig. 14) at several localities in Iran. The present paper illustrates the larval stages of both species and gives an overview of their distribution. MATERIAL AND METHOD Females of Gnopharmia species have been collected at six Localities (Table 1) by using a light trap. For laying eggs, 66 Hossein Rajaei Shoorcheh they were caged inside small plastic boxes with tissue pa- per, eventually adding suitable plant material. In Tole-Heidari valley (province Fars, south of Jahrom, N 28°28’56” E 053°23’09”, altitude 1112 m a.s.].) May 15, 2009, one female of G. kasrunensis has been collect- ed (culture A, table 1). This habitat is very dry and there are scattered bushes of different plant species. Prunus ly- cioides (Rosaceae; det. H. Akhani and M. Yazbek) as a possible host plant is common. Three females of G. kas- runensis were found in Ras-Kuh (province Kerman, road Baft-Sirjan, 2 km after Baft, N 29°17°27” E 056°35°37”, altitude 2540 m a.s.l.) May 20 to May 21, 2009 (cultures B, C and D, table 1). This locality near Ras-Kuh village has a southeast-facing aspect (Fig. 1). The dominant plant species in the area is Prunus scoparia (det. H. Akhani and M. Yazbek) (Fig. 3 and 4). Material of this plant species has been added to the plastic boxes containing females. Two females of G. colchidaria have been collected at the mountains near the village Ab-Asemani, (province Fars, road from Estahban to Sarvestan, 20 km to Sarvestan, N 29°05’51” E 053°26712”, 1890 m a.s.l., May 22, 2009) (cultures E and F, table 1). This biotope has a northern ex- posure. Prunus scoparia was common here too, but there also were wild Pistacia trees (Anacardiaceae), many bush- es of Astragalus (Fabaceae) and Echinops (Asteraceae), and several Umbelliferae spp. (Umbelliferae) (Fig. 2). One female of G. colchidaria has been collected at Osh- torankuh (province Lorestan, between Dorud and Gahar lake, near Cheshmeh-Khorram, N 33°22’41” E 049°11713 © 2360 ma.s.l., Jun 22, 2009 (culture G, table 1). Five females of G. colchidaria were found at Gardaneh-Gar- rin (province Hamadan, 25 km to Nahavand from Noorabad, N 34°02’48” E 048°20731”, 2135 ma.s.1., Jun 25, 2009 (cultures H—L table 1) and at Khani-Sefid in Gar- daneh-Khan near the border of Iran-Iraq (province Ko- rdestan, 10 km from Saghez to Baneh, N 36°04713” E 045°59’31”, altitude 1976 ma.s.l., June 26, 2009) one fe- male of G. colchidaria has been collected (culture M, table 1). No eggs or larvae have been observed in nature. Pu- pation of G. kasrunensis larvae (culture C) took place in the field (Iran), that of G. colchidaria (culture F) in the laboratory in Karlsruhe-Germany. In the field, the study of egg-morphology was done using a hand lens. For this study the species have been identified by using male genitalia of freshly emerged adults. Identifying fe- males by their genitalia 1s still problematic, as differences between species are minute and not fully understood at present. In some cases (e.g. G. colchidaria) only a female emerged and for identification genitalia dissection of si- multaneously collected males were used. There are clear characters in the male genitalia, especially in the aedea- gus, that can be used to separate the species. The aedea- gus of G. kasrunensis 1s thicker and shorter (length 1.2—1.5 mm) than that of G. colchidaria (length 1.4—1.8 mm) and bears a group of 2—6 long, rod-like, subapical spines (length up to 0.3 mm) with rounded tip (Figs 24, 28). In contrast, the aedeagus of G. colchidaria has a subapical group of short, more or less cone-shaped spines (max1- mum 0.15 mm in length), 1—3 of them situated more dis- tally and 2-3 (rarely up to 5) more basally (Figs 25, 29). The tips of the ‘octavals’ (a pair of distal processes at ster- nite 8) are more strongly curved in G. kasrunensis (Figs 22, 26), than in G. colchidaria (Figs 23, 27). The geni- talia capsules of both species are similar in size and shape, Table 1. Species, date and number of collected females and their eggs, larvae and pupa, and dates of hatching and pupation, ac- cording to cultures. Code of Species Date of Date of N. eggs Date of N. larvae Dateof N. pupa culture collecting laying egg hatching pupation A (loc. 1) G. kasrunensis 15.5.09 17.5.09 27 21.-22.5.09 21 — 0 B (loc. 2) G. kasrunensis 20.5.09 23.—24.5.09 37 27.-28.5.09 28 = 0 C (loc. 2) G. kasrunensis 20.5.09 23.—24.5.09 14 27.-28.5.09 12 25.6.09 1 D (loc. 2) G. kasrunensis 21.5.09 24.-25.5.09 28 28.-29.5.09 22 = 0 E (loc. 3) G. colchidaria 22.5.09 28.—29.5.09 24 30.—31.6.09 17 — 0 F (loc. 3) G. colchidaria 22.5.09 28.-29.5.09 33 30.-31.6.09 21 2.7.09 1 G (loc. 4) G. colchidaria 22.6.09 25.6.09 18 29.-30.6.09 16 — 0 H (loc. 5) G. colchidaria 25.6.09 27.-28.09 16 1.-2.7.09 16 _ 0 I (loc. 5) G. colchidaria 25.6.09 27.28.09 45 1.—2.7.09 36 — 0 J (loc. 5) G. colchidaria 25.6.09 27.28.09 42 ].—2.7.09 38 - 0 K (loc. 5) G. colchidaria 25.6.09 27.28.09 U7 1.—2.7.09 14 — 0 L (loc. 5) G. colchidaria 25.6.09 27.28.09 40 1.—2.7.09 36 - 0 M(loc. 6) G. colchidaria 26.6.09 28.-29.6.09 9 2.-3.7.09 7 = 0 Bonn zoological Bulletin 57 (1): 65—73 O©OZFMK Life-history of Gnopharmia kasrunensis and G. colchidaria and their distribution in Iran 67 Plate 1. Figs 1-2: Biotopes of Gnopharmia species: 1, G. kasrunensis (Ras-Kuh, cultures B, C and D); 2, G. colchidaria (Ab- Asemani village, cultures E and f); Figs 3-4: host-plant: 3, tree of Prunus scoparia; 4, leaves and fruits of Prunus scoparia; Fig. 5: eggs of G. kasrunensis (culture C); Figs 6-8: larvae of G. kasrunensis: 6, L2; 7, L4; 8, L3; Figs 9-11: larvae of G. colchidaria: ONES OS WD tle: Bonn zoological Bulletin 57 (1): 65-73 ©ZFMK 68 Hossein Rajaei Shoorcheh Plate 2. Figs 12-14: Adults of Gnopharmia: 12, G. colchidaria (female, freshly emerged; culture F); 13, G. kasrunensis (holo- type, male); 14, G. colchidaria (male) (scale 1 cm); Figs 15-16: Pupa of G. colchidaria: 15, ventral view, 16, lateral view (scale 2mm); Fig. 17: Pupa of G. kasrunensis, ventral view (one day after pupation); Fig. 18: Pupa of G. kasrunensis, posterior part (ma- le, ventral view; Cr: cremaster); Fig. 19: id., pupa of G. colchidaria, female (scale 1mm); Fig. 20: floricomous setae (distal part) on female ovipositor (scale on figure); Fig. 21: Setal map of G. colchidaria and G. kasrunensis larvae. Bonn zoological Bulletin 57 (1): 65—73 OZFMK Life-history of Gnopharmia kasrunensis and G. colchidaria and their distribution in Iran 69 but the basal process of the sacculus is distinctly longer in G. colchidaria. In G. kasrunensis both of these projec- tions seems to be equal in length (Figs 30, 31). For SEM photography male genitalia have been dehydrat- ed in 70%—96% and 100% ethanol. For drying with HMDS (1, 1, 1, 3, 3, 3-Hexamethyldisilazan) the HMDS- protocol (Oshel 1997) was used. According to this proto- col the genitalia have been transferred from 100% ethanol to a 1:1 ethanol-HMDS mixture and then to 100% HMDS. The specimens were dried by evaporation of HMDS. SEM photos were made using a Hitachi S-2460N SEM at Zoologisches Forschungsmuseum Alexander Koenig. RESULTS Rearing report and field observations The female of culture-A (G. kasrunensis) laid eggs in a plastic box containing leaves of Prunus lycioides. Most larvae hatched from these eggs, but died because of lack of fresh food plants. Leaves of P. lycioides carried with us had dried out and no P. /ycioides or another possible host plant was available. Females of cultures B—F (G. kas- runensis) laid eggs (Prunus scoparia was added to the containers) just two to three days after collecting and two days later, L1-larvae started to feed successfully. Despite attempting to provide relatively stable conditions to the larvae, many of them died in different stages. Of 79 Ll-larvae of G. kasrunensis in cultures B, C and D, just 42 survived to L2, and only 18 of them moulted again (after 7-8 days). L3-larvae were feeding almost continu- ously, but they were more active in the morning and in the afternoon. Larvae continued to perish during this stage as well. 6—8 days later just 7 larvae moulted to L4. They fed again for 5—6 days, but during this time all but one died. Fully grown (25.6.2009) the last remaining larva of this species (culture C) stopped feeding and rested under pieces of tissue paper in its container, then pupated (see Fig. 17), without spinning or producing a cocoon. L1 stage of G. colchidaria took 7—9 days. Many of the larvae again died later during several instars. Of 38 L1- larvae (cultures E and F) just 24 moulted to L2. 5 larvae died in this instar. L3-stage took 7-8 days and of 19 lar- vae just three reached L4. The feeding period of the last instar is longer than that of the last species (9 days). Ful- ly grown, just one larva pupated (Figs 15, 16). At a temperature of 25—28° C, a male of G. kasrunensis emerged after 18 days (July 12, 2009; dissected). At a Bonn zoological Bulletin 57 (1): 65-73 same temperature, one female of G. colchidaria emerged from pupa after 16 days (July 18, 2009; Fig. 12). All seven females of the cultures G—M (collected from Lorestan, Hamadan and Kordestan provinces) laid eggs (a total of 187 eggs) which were brought to Germany, in order to find and to test suitable food-plants there. 163 L1- larvae hatched but none of the plants offered was accept- ed and all larvae died. Host-plants Prunus scoparia has been offered to the L1-larvae of G. kasrunensis and G. colchidaria (cultures B—F) and was readily accepted by all of them. This plant may be con- sidered as the main food resource for both, G. kasrunen- sis and G. colchidaria, also under natural conditions. To the L4-larva of G. colchidaria (cultures E and F), feed- ing on P. scoparia before, I offered two other Prunus species (Prunus haussknechtii C. K. Schneid. and Prunus lycioides (det. H. Akhani and M. Yazbek), but no feeding marks were found in either of them. To Ll-larvae of G. kasrunensis from Tole-Heidari (cul- ture A) the common almond (Prunus dulcis (Mill.) D.A.Webb) and later Prunus haussknechtii C. K. Schneid. have been offered, but none of them was accepted. Prunus scoparia was not available at that time. All larvae of the cultures G—M have been brought to Ger- many. It took 2 days to find a plant of Prunus dulcis (Miller) D.A. Webb. During this period I offered Cratae- gus monogyna Jacq., but it was not accepted by the lar- vae and more than half of them died. The others died in the next days because they also did not accept Prunus dul- cis. Morphology of preimaginal stages Ovum Ovum: Elliptical, surface rough, green in colour (just af- ter deposition), turning red-ochre after one or two days (Fig. 5). Most of them were deposited separately in a ver- tical, upright position, but some were laid in clusters as shown in Fig. 5. Under warm conditions, L1-larvae developed very quick- ly and emerged just after 4—5 days (table 1). There is no significant difference in the shape or the struc- ture between the eggs of these two species. OZFMK 70 Hossein Rajaei Shoorcheh Plate 3. Figs 22-23: SEM photos of sternite A8, with ‘octavals’: 22, G. kasrunensis; 23, G. colchidaria; Figs 24-25: SEM pho- tos of apical part of aedeagus: 24, G. kasrunensis; 25, G. colchidaria (scale on figures); Figs 26-27: Sternite A8: 26, G. kasrunen- sis (holotype, male); 27, G. colchidaria (male); Figs 28-31: male genitalia: 28, G. kasrunensis (aedeagus); 29, G. colchidaria (ae- deagus); 30, G. colchidaria (genital capsule); 31, G. kasrunensis (genital capsule). Scale 1 mm. Bonn zoological Bulletin 57 (1): 65—73 ©ZFMK Life-history of Gnopharmia kasrunensis and G. colchidaria and their distribution in Iran 71 Larval stage G. kasrunensis Wehrli, 1939 L1: 1.42.1 mm (5 days after haching) in length, with two longitudinal pale-brown bands on green background, start- ing from the posterior end of the medio-dorsal area and continuing to the head. Head and end of the body are brown. A longitudinal brown band, similar to the dorsal band, runs ventrally from posterior end to the head. The thoracic legs and abdominal prolegs are dark brown. L2: (Fig. 6): length 6.2—-7.00 mm (11 days after hatching), brown medio-dorsal bands change (in compare to L1) from brown to more green, similar to the ground colour, but at the end of L2 the rest of brown bands are distinct- ly visible on two-thirds posterior of body. The brown ven- tral band vanishes toward head. In this stage, there are two dorsal processes at the base of the D2 seta dorsally on seg- ment A2 (see Fig. 21). The tips of these processes are dark brown or black. L3 (Fig. 8): 9.6—10.5 mm (20 days after hatching), exhibits nearly the same colour patterns as found in the fully grown larvae. The dorsal area is completely green with medio- dorsal brown bands (from A10 to A6). Ventrally, the green colour is extended, but on A6—10 there are still soft brown colour band. During L3, the colour of the dorsal process- es at the base of the D2 seta (of the A2 segment) changes from dark brown or black to brown. L4 (Fig. 7): 19.2—25.8 mm (28 days after hatching), com- pletely green in colour, with a longitudinal medio-dorsal brown bands running from the A10 to the A6 segment. The dorsal processes of the A2 segment are brown, but in two cases, changed to green. G. colchidaria Lederer, 1870 L1: 1.2—2.2 mm (7 days after hatching), coloration very similar to G. kasrunensis, but here longitudinal dorso-me- dial bands start from the posterior end and continue along two-thirds of the body dorsally (from A10 to A4—5). L2: (Fig. 9): 6.5—7.1 mm (12 days after hatching), brown dorso-medial bands continue to the head, the ventral band extends slightly to the lateral. The two dorsal processes of A2 are clearly dark brown to black in colour (like in G. kasrunensis). L3: (Fig. 10) 9.7-10.7 mm (20 days after hatching), com- pletely different in comparison with G. kasrunensis: dor- sal and ventral longitudinal brown bands extend lateral- ly and the green colour is reduced to the lateral area. Black dorsal processes on the A2 segment are clearly visible. Bonn zoological Bulletin 57 (1): 65—73 L4: (Fig. 11): 19.2-26.1 mm (26 days after hatching), pre- dominantly brown, with some yellow-green colour later- ally and dorsally. The A2 dorsal processes are brown, like the surrounding area. The position of the primary setae of the L4- larvae of both species does not exhibit any differences. (see setal map, Fig. 21). For nomination of seta used McGuffin (1977) and Stehr (2005). Pupa Pupae of both species are obviously very similar in size and colour. (Figs 15-19). They are slender, about 12.8—13 mm in length, the head section rather acute, almost trian- gular; the elongated cremaster (length 0.6 mm) bears two terminal spines of different length (right spine broken in fig. 19). Until the second day after pupation the colour re- mains green, and then changes to brown. The pupal exu- via is mid-brown, too. More material is needed in order to compare the pupal characters of both species thorough- ly. Distribution (Fig. 32). According to our present knowl- edge, G. kasrunensis and G. colchidaria occur sympatri- cally in most parts of the Zagros Mountains. G. kasrunen- sis has been described from Kazeroun (Fars prov., Iran) and is distributed in the central and eastern Zagros with some small populations in western Zagros and central Al- borz. Only two specimens of this species were collected at Sayhakil in Musandam (Oman). G. colchidaria occurs in nearly all parts of Zagros, furthermore there are popu- lations in south-eastern Iran, and in the Alborz (vic. Tehran, Qazvin, Zanjan, and Gilan). The species has been described from Helenendorf (East Azerbaijan, Transcau- casia; present name: Xanlar), and there are additional re- ports from Armenia (ssp. melanotaenia Wehrli, 1938) and other regions of S. Caucasus. The map is based on mate- rial present in the collections studied. Probably these two species have a wider distribution, but the author so far had no access to further material proving this. Bionomic data. According to the analyzed label data, de- rived from several collections, G. kasrunensis has been collected between the second week of February and the second week of September. The flight period of G. colchi- daria dates from early March to late October. G. kasrunen- sis flies at altitudes from 200 up to 2200 m, G. colchidaria from sea level to 2800 m. Both species probably are (at least) bivoltine in most places, but may be univoltine in the North of Iran and at higher elevations. Overwintering stage obviously is the pupa. ©ZFMK 72 Hossein Rajaei Shoorcheh 48° S26 Caspian Sea 56° 60° 36° 32° Kerman Ei: A $ ifn iraz a A U-~e e@ ie Br : e ew 28° : e. @n Bandar-e Abbas ry Persian Gulf 1000-2000 m > 2000 m ® G.colchidaria A G.kasrunensis Fig 32. Distribution map of G. kasrunensis and G. colchidaria in Iran. DISCUSSION Despite intensive search on many Prunus shrubs in a num- ber of Iranian localities for immature stages of Gnopharmia, no eggs, larvae or pupae were found in the nature. Of course, it is often difficult to find eggs of moths in the field, but in case of Gnopharmia these difficulties may also be related to the existence of a so-called “flori- comus” on the ovipositor of Gnopharmia females. Numer- ous specialized (“scoop-shaped”) setae (Fig. 20) on the papillae anales probably enable the females to scratch par- ticles from the surface of the bark to cover and camou- flage the eggs. This behaviour has been observed in the W. European Ennomine species Theria primaria Haworth, 1809 (Stokoe 1948: 186, pl. 39:1), females of which have a very similar ovipositor with almost identical, specialized Bonn zoological Bulletin 57 (1): 65-73 setae. This hypothesis is supported by the fact that the papillae anales of almost all females studied are spoiled with dust or bark-particles retained near the setal bases (Scoble & Kriiger, 2002, D. Stiining, Bonn, pers. comm., unpublished). Prunus scoparia as the most probable natural host plant of G. kasrunensis and G. colchidaria is well distributed in many parts of the Zagros Mountains. During the last years this plant has also been intensively grown by local people and became more widely distributed, as “bitter al- monds’ became more frequently used by the pharmaco- logical industry in Iran. G. kasrunensis and G. colchidaria fly sympatrically at many localities in Iran (Fig. 32). Lar- ©ZFMK Life-history of Gnopharmia kasrunensis and G. colchidaria and their distribution in Iran 73 vae of both species are very similar morphologically and the young larvae (L1, L2) are also without clear differ- ences in colour and pattern. However, grown larvae (L3, L4) can be separated. The reason for the high mortality of the larvae during all stages is not clear to us. Alteration of climate in the ve- hicle, low quality of the food plant, obstruction by sun- light or artificially damp environment (wet paper towels were provided to keep the food fresh) could have been the reason for bacterial or viral infection which killed the lar- vae. Further studies on the biology of this interesting moth genus will be carried out during the 2010 collecting sea- son. Acknowledgments. I deeply thank Axel Hofmann (Breisach) and Dr. Jorg-Uwe Meineke (Kippenheim) for their helpful ad- vice during our common field work, and express my sincere thanks to Prof. Dr. Hossein Akhani (Tehran) and Mariana Yazbek (Lebanon) for the determination of host plants. Furthermore, I do not forget valuable comments by Dr. Robert Trusch and Giin- ter Ebert (Karlsruhe), Dr. Dieter Stiining (Bonn) and Dr. Axel Hausmann (Munich). I thank Lily Wescott and Tom Swinehart (Bonn) for proofreading of the first draft, Stefan Scharf (Karl- sruhe) for preparing the digital map of Middle-East, Ulla Kreutz (Meckenheim) for processing of the digital images, Bernd Miiller (Berlin), Michael Falkenberg (Karlsruhe) and Maryam Bo- roomandi (Bonn) for their friendly help during different steps of this project. Special thanks go to Dr. Montazami, Alireza Naderi, Mohammad-Ebrahim Sehhati and other colleagues of Iran Department of Environment for their collaboration during our 2009 excursion and also colleagues of Hayk Mirzayans In- sect Museum in the Iranian Research Institute of Plant Protec- tion (HMIM, Tehran) for their friendly collaboration in the proj- ect Gnopharmia. | am grateful to Hamid Valipoor (Ghoochan) and Mahdi Nasiri (Dorud) for their aid in the field trip of 2009 in Iran, to Professor Dr. J. W. Wagele (ZFMK, Bonn) for his pro- motion and DAAD (Deutscher Akademischer Austausch Dienst) for the financial support of this project. REFERENCES Ebert G (1965) Afghanische Geometriden (Lep.) I. Stuttgarter Beitrage zur Naturkunde 142: 1-32, Pl. 14 Erschoff NG (1874) Cheshuekrylyja (Lepidoptera). — In: Fedchenko AP: Puteshestvie vy Turkestan [Journey to Turkestan], vol. 2 Zoogeograficheskaja izsledovanija part 5, p. E-VI, 1-127, Pl. 1-6 HOSTS (a Database of the World’s Lepidopteran Hostplants, hosted by Natural History Museum, London 2007): Bonn zoological Bulletin 57 (1): 65—73 http://www.nhm.ac.uk/jdsml/research-curation/research/proj- ects/hostplants/index.dsml Lederer J (1870) Contributions a La Faune des Lépidoptéres de la Transcaucasie. Annales de la Société Entomologique de Belgique. 13: 39, 48, pl. 2, Fig. 1 McGuffin WC (1977) Guide to the Geometridae of Canada (Lepidoptera), II subfamily Ennominae 2. Memoirs of the Entomological Society of Canada. No. 101, Fig. 152 Oshel Ph (1997) HMDS and Specimen Drying for SEM. Microscopy today 5 (4):16 Parsons MS, Scoble MJ, Honey HR, Pitkin LM, Pitkin RB (1999) The catalogue. In: Scoble MJ, ed. Geometrid Moths of the world: a catalogue (Lepidoptera, Geometridae). Collingwood: CSIRO Publishing. 1—1046 Prout LB (1912-1916) Die spannerartigen Nachtfalter. In: Seitz A (ed.): Die Gross-Schmetterlinge der Erde 4: i-v, 1-479, pls. 1—25. Stuttgart, Alfred Kernen Verlag Scoble MJ, Kriiger M (2002) A review of the genera of Macariini with a revised classification of the tribe (Geometridae: Ennominae). Zoological Journal of the Linnean Society 134: 257-315 Staudinger O (1892) Neue Arten u. Varietaten von palaarktischen Geometriden. Deutsche entomologische Zeitschrift Iris. 5:181-188, pl. 2, Fig. 12-13 Stehr FW (2005) Immature Insects. Kendall/Hunt Publishing Co., U.S. 1-768, p. 290-303 Stokoe GHT (1948) The Caterpillars of British Moths, 2nd se- ries. London: F. Warne Wehrli E (1938) Nachtrag zu meiner Arbeit in Nr. 31 d. J. Sei- te 354 [Neue Gattungen, Untergattungen, Arten und Rassen von Geometriden (Lep.).]. Entomologische Rundschau 55: 433-434 Wehrli E (1939) Einige neue Arten und Rassen aus dem sudwestlichen Iran und aus dem Irak, sowie ein neues Subgenus aus dem letzteren. (Lepid. Geometr.). Mitteilungen der Miinchner Entomologischen Gesellschaft 29: 70-71 Wehrli in Brandt W (1941) Beitrag zur Lepidopteren-Fauna von Iran (4). Einige neue Geometriden. Mitteilungen der Miinchner Entomologischen Gesellschaft 31: 864-886, PI. 28-30 Wehrli E (1939-1954) 6. Subfamilie: Geometrinae. In: Seitz A (ed.): Die Gross-Schmetterlinge der Erde 4, Die Spanner des palaearktischen Faunengebietes (Suppl.): 254-722, Pls 19-53. Stuttgart, Alfred Kernen Verlag Wiltshire EP (1967) Middle East Lepidoptera XX, A third contribution to the Fauna of Afghanistan. Beitrage zur naturkundlichen Forschung in Siidwestdeutschland 26: 137-169 Wiltshire EP (1970) Osterreichische entomologische Expe- ditionen nach Persien und Afghanistan. Annalen des Naturhistorischen Museums in Wien 74: 387-398 Received: 15.11.2009 Accepted: 20.02.2010 Corresponding editor: D.Stiining ©ZFMK Bonn zoological Bulletin | Volume 57 | Issue | | pp. 75-83 Bonn, April 2010 Two new species of the Plutodes costatus-group from the Philippines and Indonesia (Lepidoptera, Geometridae, Ennominae) Dieter Stiining Zoologisches Forschungsmuseum Alexander Koenig, Adenauerallee 160, 53113 Bonn, Germany; E-mail: d.stuening.zfmk@uni-bonn.de Abstract. Two new species, Plutodes magdelinae sp. n. from Mindanao Island, the Philippines, and P. thorbeni sp. n. from Sulawesi, Indonesia, are described. Pattern and coloration clearly indicate that they are members of the P. costa- tus-group. Besides the well-known, wide-spread and often abundant P. costatus Butler, 1886, other members of this species- group are P. warreni Prout, 1923, and P. moultoni Prout, 1923. The latter species, described from Peninsular Malaysia, is redescribed. It is very similar externally to P magdelinae sp. n., but clearly different in the genitalia morphology. Its known range of distribution is fairly extended, based on new records. P. hermanowskii Stining, 2005 (Luzon, Philip- pines) is the most recently described species in this group. DNA barcoding of all involved taxa revealed that they are well differentiated also on the genetic level. Key words. Plutodes, new species, P. magdelinae sp. n., P. thorbeni sp. n., Philippines, Indonesia, DNA barcoding. INTRODUCTION Species of the genus Plutodes Guenée [1858] are charac- terized by broadly unipectinate antennae in both, males and females, and their striking wing-pattern of yellow ground colour with bands and patches in reddish-brown or grey, the latter often highlighted by margins of irides- cent scales. Parsons et al. (1999) listed 27 species as a whole, distributed from the NW Himalayas (Sabathu, type-locality of PR warreni) and Nepal (Yazaki, 1992, 1993, 1994: eight species) to tropical Australia (P. sig- nifera Warren, 1896; cf. McQuillan & Edwards, 1996). Holloway [1994] redescribed the genus and recorded sev- en species from Borneo. He also provided a definition for the tribe Plutodini. Species of the P. costatus-group are distinguished from the typical species of Plutodes (like the type-species P. cyclar- ia Guenée and its relatives) externally by larger size and by the hind wings coloured brown with only some rem- nants of the yellow ground colour. Also in the forewings the range of brown colour is enlarged, most definitely so in P. costatus and P. warreni. P. moultoni has the yellow areas of the forewings more extended, but has the apical patch, which is very conspicuous in the two species men- tioned before, reduced to minor orange-coloured remnants. Instead, the apical region is marked by a pair of black, out- wardly pointed spots. PR. hermanowskii is transitional to the core-group in having oval apical patches to the forewings, similar to the majority of species related to P cyclaria. The two new species described herein clearly belong to the P. costatus-group, the one from Mindanao being strik- ingly similar externally to P. moultoni, but clearly differ- ent in male genitalia. P. thorbeni from Sulawesi is also dis- tinctive in its habitual features, but closer to P. costatus externally. DNA barcodes were obtained at the Canadian Centre for DNA Barcoding, University of Guelph, Canada, in the course of a project called ‘All Leps’ (cf. http:// www.lep- barcoding.org). MATERIAL AND METHODS This study is based mainly on material of the Lepidoptera collection of the Zoologisches Forschungsmuseum Alexander Koenig, Bonn, Germany (ZFMK)). In addition, the collections of the Zoologische Staatssammlung, Mu- nich (ZSM), including the collection Claude Herbulot, and two important private collections (coll. M. D. Sommerer, Munich, and coll. M. Schaarschmidt, Leipzig) were stud- ied. The relevant material housed at The Natural History Museum, London (NHM), has been checked by J. D. Hol- loway, and images of the holotype of P. moultoni Prout and its labels were provided by the collection manager at the NHM Lepidoptera department. 76 Dieter Sttining Figs 1-15. Adults of Plutodes spp. 1-4. Plutodes moultoni Prout, 1: &’, N. Thailand, 2: underside; 3: &@, Sumatra, 4: underside of 3; S—6. Plutodes magdelinae sp. nov., holotype @, Mindanao; 5: upper side, 6: underside; 7-12. Plutodes thorbeni sp. nov., Su- lawesi, 7-8: holotype &, 9-12: @ paratypes (8, 10, 12: underside); 13-15. Plutodes costatus Butler; 13: @, Sumatra; 14: 9, Suma- tra; 15: underside of 9. Bonn zoological Bulletin 57 (1): 75-83 ©ZFMK Two new species of the Plutodes costatus-group from the Philippines and Indonesia Ta External features and genitalia of all species were exam- ined. The illustrations of whole moths (upper side and un- derside) and moth genitalia were prepared from digital im- ages, and were subsequently edited in Adobe Photoshop. The terminology of the genitalia largely follows that of Klots (1956). A total of 38 specimens of the ““Plutodes costatus species- group” has been processed at the Canadian Centre for DNA Barcoding (CCDB) in order to obtain DNA barcode sequences (COI, 5’, barcode fragment, 658 bp.), most of them successfully, though partly 15 years old (for details of the procedure see Ivanova et al., 2006). Abbreviations. NHM: The Natural History Museum, London; ZFMK: Zoologisches Forschungsmuseum Alexander Koenig, Bonn, Germany; ZSM: Zoologische Staatssammlung, Munich, Germany. TAXONOMY Redescription of Plutodes moultoni Prout (Figs 1-4, 16-18, 19, 20) Plutodes moultoni Prout, 1923, Annals and Magazine of Natural History (9) 11: 321. Type locality. [Peninsular Malaysia] Selangor, Bukit Ku- tu, 3457 ft. Holotype o’. NHM, London. Male (Figs 1-4). Wingspan 31—38mm, length of forewing 17—20mm. Apex rounded, termen evenly curved, tornus rounded. Fovea lacking. Vein R, of forewing arising from cell close to the common stem of R,_;, anastomosing with Sc at varying length. R, branching off from the middle of the common stem of R;,,4. Forewings yellow, with a grey- ish-brown basal area and a distal, indistinct, orange patch, both parts broadly connected. The connecting band is al- ways present, the orange patch may be more or less dis- tinctly developed. Distal to and rarely connected with the orange patch are two blackish-brown, subapical, irregu- larly triangular spots, their tips pointing towards the ter- men. Basal area with an ill-defined anterior margin, reach- ing well into the discal cell, an acute protuberance at the position of the discal vein and a short concavity prior to the connection with the apical patch. Posterior half with indistinct, transverse, greyish striation. Iridescent scales are found along basal two-thirds of costa, anterior mar- gin of basal greyish-brown area, around orange patch and densely covering the triangular spots. Fringe of termen yellow, that of hind-margin brown. Bonn zoological Bulletin 57 (1): 75-83 Hind wings with margin evenly rounded, fringe yellow. A narrow yellow marginal band from apex to tornus, slightly broader at apical half. Remaining part of hind wing dark blackish or greyish brown, with extensive trans- verse striation. Distal margin of dark area lined with iri- descent scales. Discal spot or streak not visible. In some specimens a postmedial, double, indistinct zigzag-line present. Underside of both wings lighter yellow, forewings with a transverse, slightly curved, blackish bar from tornus to- wards middle of costa, but not reaching the latter, less strongly developed or absent in Malaysian (including the type) and Sumatran specimens (see Figs 2, 4). A band proximal to this bar rather greyish-brown than yellow. Hind wings on underside with a large, blackish, oval patch, touching or almost reaching the anterior margin near apex. Basal area of wings light yellow or suffused with darker scales (the latter character found again in specimens from Malaysia and Sumatra). Distally a broad, light yellow mar- ginal band of even width. Fringe yellow. Head with frons distinctly narrower than eye-diameter, smoothly scaled, greyish-brown, but with groups of up- right, narrow scales at dorsal margin. Vertex with larger, yellow scales and white scales between the antennal bases, basal segments and dorsal side of lower part of antennae also white. Palps slightly up-curved, of a light greyish- brown colour, the third segment small, tapering. Proboscis well developed. Antennae unipectinate, with longest branches about 7,5 times as long as the diameter of the flagellum shaft. Terminal one third of antennae simple. Patagia and anterior parts of tegulae yellow, posterior part of tegulae, thorax and abdomen greyish brown. Tibiae of hindlegs not dilated, thus without hair-pencil. Setal comb on third abdominal sternite as well as sterno-tympanal spine lacking. Tympanal organ without lacinia. Male genitalia (Figs 19, 20). Valves elongate, weakly sclerotized, apex rounded, lamina internally densely se- tose. Valve costa, like the whole genitalia capsule, weak- ly sclerotized, the sacculus being the only strongly scle- rotized part. It reaches a little more than half the length of the ventral valve margin, bearing an acute distal tooth- like process with a second tip at some distance pointing basally. A strong basal process with a more rounded tip is directed dorsally. There is also a roundly protruding area opposite to the sacculus on the external surface of the valve. Uncus broad, dorso-ventrally flattened, dorsally se- tose, with a dense group of stiff, narrow, upright scales near base, slightly curved ventrad, with pointed tip. Socii present, small, but with long setae. Gnathos with narrow lateral arms, central part a broad plate at right angles to- wards the lateral arms, without medial process. Transtil- ©OZFMK 78 Dieter Sttining ie 7 RE Figs 16-18. Holotype & of Plutodes moultoni Prout, 1923. 16: upper side; 17: underside, 18: labels. Images reproduced by kind permission of the Trustees of the Natural History Museum, Lon- don. la a triangular, curved, cap-like plate, its tip pointing dis- tally, with a pair of short, narrow processes arising from the proximal corners and directed caudally. Juxta a round plate, with a v-shaped incision with irregular margins dor- sally. Saccus narrow, strongly elongate, rounded distally. The aedeagus straight, club-shaped, with a stout proximal and a narrow distal half, the vesica with an elongate, se- mi-cylindrical sclerotization and a small group of cornu- ti, a long, curved, distal one and 3—4 distinctly smaller ones, arising from a common base. Bulbus ejaculatorius moderately large, about half the length of the aedeagus. Male genitalia of specimens from N Thailand a little small- er and slightly differing to those from Sumatra in the length-proportions of the saccular processes (Fig. 20). Bonn zoological Bulletin 57 (1): 75-83 Female similar to male, wingspan 35—37 mm, length of forewing 19-21 mm, with forewings a little broader and termen more strongly curved. Pectinations of antennae about one fifth shorter than in males and a little narrow- ei Material studied. Type material. Holotype, o’, [Peninsu- lar Malaysia], Bukit Kutu, Selangor, April 1915, 3457’; coll. NHM (see Figs 16-18). J. D. Holloway (London, pers. comm.) mentioned two more specimens from Peninsular Malaysia in coll. NHM, but Prout (1923) ob- viously only had one specimen at hand when he described this species (i.e. the holotype). Further material. 19, [Peninsular] Malaysia, Taiping [~ 4°51’N 100°46’E], May [19]79; 40’, N. Sumatra, vic. Tiga-Dolock, 18., 25.v.1972, leg. Ro- esler & Ktippers; coll. ZFMK; 1c’, N. Sumatra, Dolok Me- rangir, 21.1x.—5.x.1967, leg. Dr. E. Diehl, ex colli Dr Re Bender, coll. ZSM. (more specimens from N. Sumatra in coll. M. D. Sommerer, Munich). 2c", Borneo, Sabah, route Keningau-Kimanis PK 28.5, 1300m, 8., 11.viti.1991, leg. Haxaire. Herbulot collection in coll. ZSM; 1c’, S. Vietnam, Prov. Quang Tri, Partung, Da Krong, 1.—3.vi.2005; coll. ZFMK; 1o', SE Thailand, Prov. Trat, Kho Chang, 0m, 25.— 30.x1.2004, leg. T. Ihle; coll. ZFMK; 8", N. Thailand, Chiang Mai Prov., Doi Suthep, 1050m, 25.x1.2000, leg. D. Stiining; 4c", same locality, 1100m, 15.-19.1.2008, leg. T. Ihle; 10’, N. Thailand, Chiang Mai Prov., Doi Angkhang, W Fang, 1600m, 21.v.2004; 19, N. Thailand, Chiang Mai Prov., Doi Inthanon, 1500m, 28.—30.iv.2006, leg. T. Ihle; 19, N. Thailand, Mae Hong Son Prov., ca. 30km E Khun Yuam, 950m, 1.v.2006, leg. T. Ihle; coll. ZFMK; 16 specimens from Doi Suthep, Chiang Mai Prov., are kept in the collection of Bro. Amnuay Pinratana, St. Gabriel’s College, Bangkok. 1c, S. China, S. Yunnan Prov., Mangxi Ba Mts., Simao distr., 18km S Simao city, 1280m, 16.i11—10.1v.2000, ex coll. Dr. R. Brechlin. Coll. ZFMK. Distribution and bionomics. Peninsular Malaysia (type- locality), Borneo, Sumatra; Vietnam, SE & N. Thailand, S. China (first record for all localities except Peninsular Malaysia). Specimens have been collected in January, March-April, May, June, Sept.—October, November; at el- ©ZFMK Two new species of the P/utodes costatus-group from the Philippines and Indonesia 79 evations between 0 and 1600m. The immature stages are unknown. Plutodes magdelinae sp. nov. (Figs 5, 6, 21) Male (Figs 5—6). Wingspan 37—38mm, length of forewing 21—22mm. Apex rounded, termen evenly curved, tornus rounded. Fovea lacking. Vein R, of forewing arising from anterior vein of cell close to the common stem of R_; — both at rather large distance from upper corner of cell — , anastomosing with Sc at varying length. R, branching off from the middle of the common stem of R3,4. Forewings yellow, with a greyish-brown basal area and a distal, in- distinct, orange patch, both parts narrowly connected. The connecting band is always present, the orange patch may be more or less distinctly developed. Distal to the orange patch are two blackish-brown, subapical, elongate trian- gular spots, their tips pointing towards the termen. Basal area with an ill-defined anterior margin, reaching just a little beyond the posterior margin of the discal cell. Pos- terior half with indistinct, transverse, greyish striation. [ri- descent scales are not very obvious, weakly covering basal two-thirds of costa, anterior margin of basal greyish-brown area, orange patch and the submarginal area near the tri- angular spots. Fringe of termen yellow, that of hind-mar- gin brown. Hind wings with margin evenly rounded, fringe yellow. A narrow yellow marginal band from apex to tornus, dis- tinctly broader at apical half. Remaining part of hind wing dark blackish or greyish brown, with extensive transverse striation. Distal margin of dark area lined with iridescent scales. Discal spot visible as a short, curved, white streak. Postmedial ziczac-line absent. Underside of both wings (Fig. 6) lighter yellow, forewings with a broad, posterior, greyish-brown band between hind margin and posterior vein of cell, but not reaching termen. Hind wings on underside with a large, blackish-brown, oval patch, almost reaching the anterior margin near apex. Basal area of wings strongly suffused with darker scales. Distally a rather broad, light yellow marginal band of even width. Fringe yellow. Head with frons distinctly narrower than eye-diameter, smoothly scaled, greyish-brown, with thin white lateral margins, but with groups of upright, narrow scales at dor- sal margin. Vertex with larger, yellow scales and white scales between the antennal bases, basal segments and dor- sal side of lower part of antennae also white. Palps slight- ly up-curved, basal segment predominantly white, second and third of a light greyish-brown colour, the third seg- ment small, tapering, with a white tip. Proboscis well de- veloped. Antennae unipectinate, with longest branches Bonn zoological Bulletin 57 (1): 75-83 about 8,5 times as long as the diameter of the flagellum shaft. Terminal one third of antennae simple. Patagia and anterior parts of tegulae yellow, posterior part of tegulae, thorax and abdomen greyish brown. Tibiae of hindlegs not dilated, thus without hair-pencil. Setal comb on third ab- dominal sternite as well as sterno-tympanal spine lacking. Tympanal organ without lacinia. Male genitalia (Fig. 21). Valves elongate, weakly scle- rotized, apex rounded, lamina internally densely setose. Valve costa, like the whole genitalia capsule, weakly scle- rotized, but sacculus with a prominent and strongly scle- rotized, plate-like process, inserting in an upright position close to the ventral margin of the saccular region. It is get- ting broader distally, with an acute tip, narrower basally, with a rounded, very short, dorsal bulb. Outer margin of the plate densely covered with short, appressed setae. Length a little more than half the length of the ventral valve margin. Uncus broad, dorso-ventrally flattened, dor- sally setose, with a dense group of stiff, narrow, upright scales near base, slightly curved ventrad, with pointed tip. Soci very small, marked by a few long setae only. Gnathos with narrow lateral arms, central part a broad plate at right angles towards the lateral arms, without me- dial process. Transtilla a triangular, curved, cap-like plate, its tip pointing distally, with a pair of short, narrow processes arising from the proximal corners and directed caudally. Juxta a narrow plate, with a deep, v-shaped in- cision dorsally. Saccus narrow, strongly elongate, round- ed distally. The aedeagus almost straight, club-shaped, with a stout proximal and a narrow distal half, the vesica with an elon- gate, semi-cylindrical sclerotization and a small group of 5—6 cornuti, not much differing in size. Bulbus ejacula- torius moderately large, about half the length of the aedea- gus. Diagnosis. P. magdelinae is distinguished from the exter- nally very similar P. moultoni by slightly larger size, the wing-shape (forewings more elongate with more strong- ly oblique termen), a narrower greyish-brown basal area (anterior margin just reaching the posterior vein of discal cell, distal margin more strongly concave). The subapi- cal, black, triangular spots are larger and more distinctly acute towards the termen. In the hind wings, the yellow marginal band is much narrower at tornal half. On under- side, the oblique bar on forewing from tornus to costa is absent, the basal area of hind wing is much darker. In the male genitalia, the elongate, simple plate on sac- culus without a separate basal process is distinctive. Female unknown. ©OZFMK 80 Dieter Sttining Figs 19-22. c'-genitalia of Plutodes spp. 19: P. moultoni Prout, Sumatra (vesica partly everted); 20: P. moultoni Prout, N. Thai- land; 21: P magdelinae sp. nov., paratype, coll. no. LEP 2010/ 469-2; 22: P. thorbeni sp. nov., paratype, coll. no. LEP 2010/ 472- 3. (bottom left: aedeagus, bottom right: enlarged apical part of aedeagus). Type-material. Holotype, o', “Philippinen, Mindanao (Prov. Davao del Sur), Mt. Apo, SE-route via Kapatagan, 1570m, 10.-12.July 1996, prim. forest, leg./ ex coll. Dr. Ronald Brechlin. Coll. ZFMK, coll. no. LEP 2010/468 (Biodat). Paratypes. 2c’, same data as holotype, coll. nos. LEP 2010/469-1, 469-2; 1c", Philippinen, Mindanao, Mt. Ki- tanglad, Intavas, 600m, 8° 10’N 124°56’E, 23.24. ii. 2009, leg. J. H. Lourens & K. Knoblich, coll. ZFMK, coll. no. Lep 2010/470; Bonn zoological Bulletin 57 (1): 75-83 Io, Philippinen, Mindanao, Mt. Dalongdong, 9-17. vil.1999. Coll. M. Schaarschmidt, Leipzig. [“Dalong- dong” correctly is named Dolongdong and 1s a village on a high plateau at the western slopes of Mt. Kalatungan, 45km SW Talakag, 40km W Maramag, 07°53’N 124°40’E; J.H. Lourens, Luzon, in litt.] Distribution and bionomics. Probably endemic to Min- danao Island, the Philippines, only known from Mt. Apo, Mt. Kitanglad and the Dolongdong high plateau near Mt. Kalatungan. Specimens have been collected in February, July and August at 600m and 1570m a.s.l. The immature stages are unknown. ©OZFMK Two new species of the Plutodes costatus-group from the Philippines and Indonesia 81 Etymology. The present new species is dedicated to Magdeline Camille Vlasimsky (formerly Frankfurt a. M., Germany, now Dallas/ Texas), in recognition of her and her parents’ Lezlie and Stan valuable contribution to the taxonomic research of the author and to conservation of biodiversity. Plutodes thorbeni sp. n. (Figs 7-12, 22) Male (Figs 7-12). Wingspan 34-39mm, length of forewing 19—23mm. Apex slightly acute, termen evenly curved, tornus rounded. Fovea lacking. Vein R, of forewing arising from anterior vein of cell close to the common stem of R,_;— both at rather large distance from upper corner of cell- , anastomosing with Sc at varying length. R, branching off from the middle of the common stem of R3,,4. Forewings yellow, with a greyish-brown basal area and a distal, more or less distinct, irregularly quadrangular, dark brown to orange-brown, rarely light or- ange patch, both parts broadly or narrowly, sometimes not connected. Internal surface of the patch with a dark, waved line. Apical corner of the patch forming an elongate, black spot, its tip connected with the apex of the forewings. The spot is most distinctive in specimens with a reduced, or- ange, apical patch. Basal area with an ill-defined anteri- or margin, reaching well beyond the posterior margin of the discal cell, with an acutely triangular protuberance at the position of the discal vein and a broad concavity pri- or to the connection with the apical patch. Posterior half with indistinct, transverse, greyish striation. A margin of iridescent, leaden-grey scales along the anterior margin and around the apical patch, including the black apical streak, but not along distal margin. Fringe of termen yel- low, that of hind-margin brown. Hind wings with margin evenly rounded, fringe yellow, a narrow yellow marginal band from apex to middle of termen, with rather straight internal border. Larger part of hind wing dark blackish or greyish brown, with extensive transverse striation. Distal margin of dark area lined with iridescent scales. Discal spot weak, visible as a short, curved, white streak. Indistinct postmedial double waved or ziczac-line present in most specimens. Underside of both wings (Figs 8, 10, 12) yellow, fore- wings with a tiny, black, apical spot and a short, transverse, almost straight, blackish bar from tornus towards the cen- tre of the wing. There is a greyish-brown band between hind margin and posterior vein of cell, more or less agree- ing with the dark basal area of the upper side and also a dark patch representing the apical patch of upper side. Hind wings on underside with a large, blackish-brown, oval patch, broadly touching the anterior margin near apex. Basal area of wings yellow, often suffused with darker Bonn zoological Bulletin 57 (1): 75—83 scales. Distally a narrow, light yellow marginal band from apex to the middle of termen, basal half represented by the yellow fringe scales only. Head with frons distinctly narrower than eye-diameter, smoothly scaled, greyish-brown, with thin white ventral margin, but with groups of upright, narrow scales at dor- sal margin. Vertex with larger, yellow scales and white scales between the antennal bases, basal segments and dor- sal side of lower part of antennae also white. Palps slight- ly up-curved, basal segment predominantly white, second and third of a light greyish-brown colour, the second edged white distally, the third segment small, tapering, with a white tip. Proboscis well developed. Antennae unipecti- nate, with longest branches about 8,5 times as long as the diameter of the flagellum shaft. Terminal one third of an- tennae simple. Patagia and anterior parts of tegulae yel- low, posterior part of tegulae, thorax and abdomen grey- ish brown. Tibiae of hindlegs not dilated, thus without hair-pencil. Setal comb on third abdominal sternite as well as sterno-tympanal spine lacking. Tympanal organ with- out lacinia. Male genitalia (Fig. 22). Valves elongate, weakly scle- rotized, apex rounded, lamina internally densely setose. Valve costa, like the whole genitalia capsule, weakly scle- rotized, but sacculus with a prominent and strongly scle- rotized, triangular plate with an acute tip distally, insert- ing in an upright position close to the ventral margin of the saccular region. Basally, it is getting narrow and widens again to a rounded, dorsal bulb. Outer margin of plate and bulb scobinate and slightly setose. Length of sac- culus and its process a little more than half the length of the ventral valve margin. Uncus broad, dorsally rounded, ventrally flattened, dorsally setose, slightly curved ven- trad, with pointed tip, with a dense group of stiff setae near base. Socii very small, marked by a few long setae only. Gnathos with narrow lateral arms, central part a broad plate at right angles towards the lateral arms, without me- dial process. Transtilla a triangular, curved, cap-like plate, its tip pointing distally, with a pair of short, narrow processes arising from the proximal corners and directed caudally. Juxta a narrow plate, with a deep, v-shaped 1n- cision dorsally. Saccus narrow, strongly elongate, round- ed distally. The aedeagus club-shaped, with a stout proximal and a narrow, slightly curved distal half, the vesica with an elon- gate, semi-cylindrical sclerotization and a small group of 5—6 cornuti, not much differing in size. Bulbus ejacula- torius moderately large, about half the length of the aedea- gus. Diagnosis. P. thorbeni is in no way similar to P. moultoni or P. magdelinae, but exterrnally quite close to P. costa- ©ZFMK 82 Dieter Stiining tus, the type of this species-group (Figs 13—15). Both are of the same size and the arrangement of yellow and brown pattern elements is very similar. In detail, P thorbeni is distinguished from P. costatus in the apical forewing patch being smaller, darker und more strongly irregular. P. costa- tus has the internal surface of this patch rather grey in- stead of brown. P. thorbeni has a black, acute apical spot or streak, best seen in specimens with a reduced apical patch (Figs 9, 11). It is strongly highlighted by iridescent scales and absent in P. costatus. There is a curved, black- ish antemedial line in P. costatus, absent in P. thorbeni. Fringe of termen of forewings is completely yellow in P. thorbeni (partly grey in P. costatus) and there is more of the yellow ground colour distal to the brown pattern ele- ments. In the hind wing, the tornal half is narrowly yel- low in P. thorbeni, but brown — including fringe — in P costatus. The underside is very different in both species, i.e. the large black patch of the hind wing seen in P. thor- beni is absent in P. costatus, the surface being light grey instead in the latter (Fig 15). The underside of P. thorbeni is much more similar to P. magdelinae and P. moultoni. Also the male genitalia are strongly different, especially in the shape of the saccular process, which is also more similar to P. magdelinae (Fig 21), and cannot be mistak- en. Female unknown. Type material. Holotype, o’, “Indonesia, C. Sulawesi, Tambusisi mts., 1800m, 1° 45’S 121° 27°E, 11. 1996, na- tive collector”. Coll. ZFMK, coll. no. LEP 2010/471. Paratypes. 60’, same data as holotype; 20’, South Sula- wesi, Quarles Mountains, ca. 30km N Rantepao, Polo Polo, 2200m, 26.—28.1x.1995, leg. H. Schnitzler; 3c", Cen- tral Sulawesi, Sempuraga (=Sampuraga), 1500—1700m, vi. 1996, G. Lecourt & J. Martin leg.; 2c", C. Sulawesi, vic. Mamasa, 2° 57’S 119° 24’E, 17.-18.x.1995, leg. local col- lector (Gala); 1c, S. Sulawesi, Puncak Palopo, 900—-1300m, x.1997, leg. local collector, ex coll. Dr. R. Brechlin. Coll. ZFMK, coll. nos LEP 2010/472-476. 2°, S. Sulawesi, Puncak Palopo, 900—1300m, 11.1998, leg. local collector, coll. M. Schaarschmidt, Leipzig, Germa- ny. 2c", C. Sulawesi, Mt. Tambusisi, x. 1195, native collec- tor; 70", same locality, 11. 1996; 2c’, same locality, i1. 1996. Coll. K. Yazaki, Tokyo. Further material studied. J. D. Holloway sent images of three male specimens, collected by him during the ““Pro- ject Wallace” of the Royal Entomological Society. Col- Bonn zoological Bulletin 57 (1): 75-83 lecting site: “Sulawesi Utara, Dumoga Bone N.P., Site 15, 1140m, 6.8. vili.1985”. According to his recommenda- tion, these three undissected males which also show slight external differences compared to the paratypes mentioned above and which come from a locality in Northern Sula- wesi rather far away from the type locality are not recor- ded as paratypes. Distribution and bionomics. Probably endemic to Su- lawesi, but widely distributed on the island. Specimens have been collected in February, June, September and Oc- tober at elevations between 900m and 2200m a.s.1. The immature stages are unknown. Etymology. The present new species is dedicated to Thor- ben Briese (Lippstadt, Germany), in recognition of his and his parents’ Dr. Anette Meyer-Briese and Dr. Harald Briese valuable contribution to the taxonomic research of the au- thor and to conservation of biodiversity. DNA BARCODING RESULTS Results obtained from the Canadian Centre for DNA Bar- coding (CCDB) revealed clear specific distances between all species involved: pairwise distances are between 3.13% and 7.25% in the P. costatus-group. Of the species treat- ed in the present paper, P. moultoni and P. thorbeni ex- hibit the closest relationship (medial pairwise distance 3.54%). In addition, an unusually deep divergence was found among the analysed specimens of P. warreni Prout, indi- cating a cryptic new taxon within material from Laos, Viet- nam and Thailand. Morphological studies on this group are carried out at present and will be published in near fu- ture, together with a more comprehensive account on the barcoding results of the whole group. Preliminary results can be viewed under www.boldsystems.org/ Plutodes. Acknowledgements. Thanks are due to A. Schintlmeister, Dres- den, for presenting Geometridae material from the Philippines collected by Karel Cerny, Victor Sinjaev and himself to the Lep- idoptera & Trichoptera Section of ZFMK. I also thank Jan Lourens, Lucena City, Luzon, the Philippines, for presenting valuable specimens of Plutodes, including one paratype of a species described herein. I am thankful to J. D. Holloway, NHM London, and M. Sommerer, Munich, for various data and infor- mation about species of P/utodes in the collections under their care. The latter also checked, besides his private collection, the Herbulot collection now housed at the ZSM. K. Yazaki, Tokyo, and M. Schaarschmidt, Leipzig, provided data and sent images of specimens in their private collection. I am also very thank- ful to Mrs. Ulla Kreutz, Meckenheim, for preparing the figures and plates presented in this article. The careful processing of the material by the CCDB team (University of Guelph) is grateful- ly acknowledged as well. ©ZFMK Two new species of the Plutodes costatus-group from the Philippines and Indonesia 83 REFERENCES Holloway JD (1993) [1994]. The Moths of Borneo, part 11: Geometridae, Ennominae. Malayan Nature Journal 47: 1-309, 19 pls Ivanova NV, de Waard JR, Hebert PDN (2006) An inexpensive, automation-friendly protocol for recovering high-quality DNA. Molecular Ecology Notes 6: 998-1002 Klots AB (1956) Lepidoptera. In: Tuxen SL, ed. Taxonomist’s glossary of genitalia in insects. Copenhagen: Ejnar Munks- gaard, 97-111 McQuillan PB, Edwards ED (1996) Geometridae. In: Nielsen ES, Edwards ED, Rangsi TV (eds): Checklist of the Lepi- doptera of Australia. Monographs on Australian Lepidoptera, CSIRO Publishing, Collingwood Parsons M, Scoble MJ, Honey MR, Pitkin LM, Pitkin BR (1999) In: Scoble MJ (ed.): Geometrid Moths of the World: a Cata- logue (Lepidoptera, Geometridae). CSIRO Publishing/ Apol- lo Books, Collingwood/Stenstrup Bonn zoological Bulletin 57 (1): 75-83 Stiining D (2005) Plutodes hermanowskii sp. n., a new species of the P. costatus — group from the Philippines. Tinea 21: 9-22 Yazaki K (1992) Geometridae. In: Haruta T (ed.): Moths of Nepal, part 1. Tinea 13 (Suppl. 2): 5-46 Yazaki K (1993) Geometridae. In: Haruta T (ed.): Moths of Nepal, part 2. Tinea 13 (Suppl. 3): 103-121 Yazaki K (1994) Geometridae (part). In Haruta T (ed.): Moths of Nepal, part 3. Tinea 14 (Suppl. 1): 5-40 Yazaki K (2004) Notes on geometrid moths (Lepidoptera, Geo- metridae) from Nanling Mts, S. China (III). Tinea 18: 117— 126 Received: 10.12.2009 Accepted: 30.01.2010 Corresponding editor: F. Herder ©ZFMK Bonn zoological Bulletin ] Volume 57 Issue | pp. 85-89 Bonn, April 2010 Proclinopyga ulrichi sp. nov.: the first fossil aquatic dance fly of the subfamily Clinocerinae (Diptera: Empididae) Bradley J. Sinclair Canadian National Collection of Insects, Ottawa Plant Laboratory — Entomology, CFIA, Ottawa, Canada; E-mail: bradley.sinclair@inspection.ge.ca Abstract. Proclinopyga ulrichi sp. nov. is described from Baltic amber, and represents the first definitive clinocerine fos- sil species. Palaeoparamesia proosti Meunier, 1902, considered a possible species of Clinocerinae, is transferred to the Trichopezinae. Key words. Taxonomy, Baltic amber fossil, new species, morphology INTRODUCTION The Empidoidea comprise five families and numerous subfamilies of mostly predaceous flies (Sinclair & Cum- ming 2006). Some 150 fossil species of Empidoidea (ex- clusive of Dolichopodidae s.str.) are described (Evenhuis 1994; Solorzano Kraemer et al. 2005). Fossils are de- scribed from most major empidoid lineages, extending the origin of these lineages to at least the Cretaceous period (Grimaldi & Cumming 1999). The subfamily Clinocerinae consists of primarily aquat- ic associated taxa, with adults mostly encountered on emergent rocks in streams and rivers (Sinclair 1995). Two fossil species have been classified as clinocerines. Fushunempites furvus Hong, 2002, from Eocene Chinese amber is possibly a clinocerine on the basis of the long and narrow cell r,. Unfortunately the photo of the speci- men (Hong 2002a, b) does not permit a definitive deci- sion concerning subfamily assignment. Palaeoparamesia proosti Meunier, 1902 has also been viewed as a possi- ble clinocerine. A specimen of this Baltic amber species, identified by Meunier (1908), was examined in this study and found not to represent a clinocerine (see Discussion). Consequently, the description herein of a new Baltic am- ber species of the recent genus, Proclinopyga Melander, represents the first unequivocal clinocerine fossil. The age of Baltic amber is assumed to be 40 million years (Eocene). MATERIALS AND METHODS To prepare the pieces for identification, the amber was cut and polished using an IsoMet® Low Speed Saw (Buehler, Lake Bluff, Illinois, USA) cutting machine and a Phoenix® Beta Grinder-Polisher (Buehler) polishing ma- chine with SiC grinding paper for metallography, grit 800, 1200 and 2500, Microcut® Abrasive Paper (Buehler) plain backing P 4000, and paperboard. To protect the holotype of the new species the piece was embedded in synthetic resin Araldite® 2020 (XW396/XW397) (Huntsman Ad- vanced Materials, Everberg, Belgium) (M. Solorzano Kraemer, Bonn, pers. comm. 2008). Terms used for adult structures primarily follow those of McAlpine (1981), except for the antenna where terms of Stuckenberg (1999) were used. Homologies of the male terminalia follow those of Sinclair & Cumming (2006). The Baltic amber specimen of Palaeoparamesia examined in this study was on loan from the Geoscience Centre of the University of Gottingen (Germany). Genus Proclinopyga Melander Proclinopyga Melander, 1928: 220. Type-species, Pro- clinopyga amplectens Melander, 1928 (original designa- tion). 86 Bradley J. Sinclair Diagnosis. Proclinopyga is distinguished from other gen- era of the Clinocerinae by its protruding face, bare eyes, labellum with pseudotracheae, broad wings with Sc evanescent, fifth tarsomere lacking extension, dorsal mar- gin of the epandrium expanded with setae and female ab- domen truncate with stout and ridged cercus. J a al Proclinopyga ulrichi sp. nov. (Figs. 1, 2, 4, 5) Type Material. Male holotype in Baltic amber, with fol- lowing label data: “HOLOTYPE/ Proclinopyga/ ulrichi/ Sinclair”; “CNC DIPTERA/ # 12248”: amber is imbed- ded in resin with following dimensions: 11 x 9 x 6 mm; Figs 1-3. 1. Proclinopyga ulrichi, habitus. 2. Proclinopyga ulrichi, male terminalia, postero-lateral view of right side. 3. Palaeo- paramesia proosti, habitus. Bonn zoological Bulletin 57 (1): 85-89 ©ZFMK Proclinopyga ulrichi sp. nov.: the first fossil aquatic dance fly of the subfamily Clinocerinae 87 included in amber block is a male chironomid (missing abdomen). The holotype is housed in the amber collec- tion of Dr. Hans Ulrich at the Zoologisches Forschungsmuseum A. Koenig (Bonn, Germany). Recognition. This species (Fig. 1) is distinguished by the lengthened, attenuated apical half of the postpedicel, con- voluted phallus with its hair-pin bend at mid-length and row of stout posteroventral setae on the male mid-femur. Description. Male. Eye bare, dichoptic; lower half of face with slight protuberance in profile. Antenna with scape slightly longer than pedicel; postpedicel pointed ovate, longer than pedicel and scape combined, strongly atten- uated and prolonged in apical half; arista-like stylus 1.5 times longer than postpedicel. Proboscis short; palpus club-shaped; labellum held in horizontal position, suck- er-like. Alternating biserial row of acrostichal setae, ending at 5 Figs 4-5. 4. Proclinopyga ulrichi, wing. 5. Proclinopyga ulrichi, male terminalia, lateral view of right side. Abbreviations: cl cere — clasping cercus; epand — epandrium; epand Ib — epandrial lobe; hypd — hypandrium; ph — phallus; sur — surstylus. Bonn zoological Bulletin 57 (1): 85—89 ©OZFMK 88 Bradley J. Sinclair prescutellar depression; 6—7 dorsocentral bristles, foremost large and positioned toward postpronotal lobe; 1 post- pronotal bristle; 1 presutural supra-alar bristle; 2 noto- pleural bristles; 2 postsutural supra-alar bristles; 1 posta- lar bristle; 4 scutellar bristles. Laterotergite with short, black setae in row. Wing (Fig. 4) (length 3 mm) broad, slightly infuscate; basal costal bristle long, reaching to near humeral crossvein; anal lobe well developed; Ry strongly diver- gent from Rs. Legs with evenly scattered setae. Mid femur with pos- teroventral row of long, stout setae, longer than width of femur. Mid tibia bearing stout, short, erect ventral setae; one antero- and one posterodorsal setae near base. Hind tibia with row of erect dorsal and anteroventral setae on distal two-thirds. Abdomen with tergite 6 with transverse row of 6 erect dark setae, length more than half width of abdomen. Termina- lia (Figs. 2, 5): Hypandrium cone-shaped, as long as epan- drium is wide. Phallus long, slender and convoluted, with strong bend at mid-length; apex slightly expanded. Epan- drium with expanded posterodorsal margin, bearing long setae and setulae; epandrial lobe continuous with expand- ed region of epandrium, bearing long setae and attenuat- ed apically. Surstylus closely associated with epandrial lobe; shallow notch proximal to slightly expanded and truncate apex. Clasping cercus digitiform, straight, longer than surstylus; bearing short setulae. Female. Unknown. Geographical Distribution and Phenology. Extant species of Proclinopyga are confined to the Nearctic, east- ern Palearctic, and Oriental regions (Sinclair 1995). This fossil species is the only known representative of this genus in Europe and the western Palearctic Region. Adults of Proclinopyga are associated with running water habi- tats, most commonly collected in streams, creeks, and cas- cades. The immature stages of this genus are unknown. Phylogenetic Relationships. Proclinopyga comprises ten described species and at least this many undescribed species (Sinclair 1995). The slender, highly convoluted phallus (Fig. 5) of the fossil species is most similar to ex- tant species in North America [e.g., P_ flavicoxa Melander, 1928 (see Sinclair 1995, fig. 13)]. Etymology. This species is dedicated to Dr. Hans Ulrich who has amassed a rich collection of little known amber species of Empidoidea. Bonn zoological Bulletin 57 (1): 85-89 DISCUSSION Proclinopyga ulrichi represents the first definitive clinocerine fossil taxon. The presence of Clinocerinae in Baltic amber was first definitively stated by Ulrich (2003, table, fig. 1) and was based on the specimen described here (H. Ulrich, Bonn, pers. comm. 2008). As in other subfam- ilies of the Empidoidea (exclusive of Dolichopodidae s.str.), Baltic amber fossils often belong to Recent genera (see discussion by Ulrich 2003). Clinocerines appear to be underrepresented in amber compared to other groups of empidoids, likely resulting from their very close asso- ciation with flowing water habitats. Adults are rarely col- lected away from emergent rocks in streams and rivers and they do not form aerial swarms and consequently rarely would they be associated with tree trunks and other sources of amber. Palaeoparamesia proosti was described on the basis of a single male specimen (No. 2118) (Meunier 1902). Meu- nier (1908) listed two additional specimens of this species (Nos. 8437, 7786) in his monograph on Baltic amber Em- pididae. Only specimen No. 8437 from these original se- ries remains in the Gottingen collection (H. Ulrich, Bonn, pers. comm. 2001). This male specimen was examined and found to be in excellent condition (Fig. 3) and conspecif- ic with P proosti. This examination revealed that Palaeoparamesia Meunier should be assigned to a group of several small genera currently found in the western Nearctic region and assigned to the Trichopezinae; 1.e., Boreodromia, Nearctic species of Apalocnemis Philipp and two winter collected species of a genus with anten- nae morphologically similar to Gloma Meigen (Sinclair 2008). Palaeoparamesia is characterized by a dichoptic male head, pair of long frontal bristles (rare in empidoids, except for Hilarini), postpedicel somewhat similar to Glo- ma, Trichopezinae-like female terminalia (based on con- specific specimen in Ulrich collection), hypandrium and epandrium fused (at least partially), erect male cerci and ear-like surstyli. On the basis of similarly shaped postpedi- cel and dichoptic male head, Palaeoparamesia and the winter species near G/oma are most closely related. These genera will be more thoroughly examined and described in a future publication by the author. Acknowledgements. Many thanks to Monica Solorzano Krae- mer (University of Bonn) for preparing the amber specimen. Hans Ulrich (Bonn, Germany) generously permitted the exam- ination of his amber collection, access to literature and person- al notes on the Meunier amber collection. Mike Reich (Univer- sity of Géttingen) kindly arranged the loan of the amber speci- men, Palaeoparamesia proosti. Henri Goulet (CNC — Canadi- an National Collection) produced the photographs of the amber inclusions. Jeff Cumming (CNC), H. Ulrich and three anony- mous reviewers provided valuable comments on earlier drafts. ©OZFMK Proclinopyga ulrichi sp. nov.: the first fossil aquatic dance fly of the subfamily Clinocerinae 89 REFERENCES Evenhuis NL (1994) Catalogue of the Fossil Flies of the World (Insecta: Diptera). Backhuys Publishers, Leiden Grimaldi D, Cumming J (1999) Brachyceran Diptera in Creta- ceous ambers and Mesozoic diversification of the Eremoneu- ra. Bulletin of the American Museum of Natural History 239: 1-124 Hong Y-C (2002a) Amber Insects of China. Beijing Science & Technology Press, Beijing Hong Y-C (2002b) Atlas of Amber Insects of China. Henan Sci- entific & Technological Publishing House, Beijin McAlpine JF (1981) Morphology and terminology — Adults. [Chapter] 2. Pp 9-63 in: McAlpine JF, Peterson BV, Shewell GE, Teskey HJ, Vockeroth JR & Wood DM (coords.) Man- ual of Nearctic Diptera, Volume |. Agriculture Canada Mono- graph 27 Melander AL (1928) Diptera, Fam. Empididae. Pp. 1-434 in: Wytsman P (ed.) Genera Insectorum 185 (1927) Meunier F (1902) Description de quelques dipteres de l’ambre. Annales de la Société scientifique de Bruxelles 26: 96-104 Meunier F (1908) Monographie des Empidae de l’ambre de la Baltique et catalogue bibliographique complet sur les dipte- res fossiles de cette résine. Annales des Sciences Naturelles Zoologie 7: 81-135, pls HI-XII Bonn zoological Bulletin 57 (1): 85—89 Sinclair BJ (1995) Generic revision of the Clinocerinae (Empi- didae), and description and phylogenetic relationships of the Trichopezinae, new status (Diptera: Empidoidea). The Cana- dian Entomologist 127: 665—752 Sinclair BJ (2008) Review of three little-known monotypic gen- era (Diptera: Empidoidea: Brachystomatidae), assigned to Tri- chopezinae. Zootaxa 1754: 52-62 Sinclair BJ, Cumming JM (2006) The morphology, higher-lev- el phylogeny and classification of the Empidoidea (Diptera). Zootaxa 1180: 1-172 Solorzano Kraemer MM, Sinclair BJ, Cumming JM (2005) Five new species of Tachydromiinae (Diptera: Empididae s.1.) from New World Tertiary ambers. Zootaxa 1010: 37-52 Stuckenberg BR (1999) Antennal evolution in the Brachycera (Diptera), with a reassessment of terminology relating to the flagellum. Studia dipterologica 6: 33-48 Ulrich H (2003) How recent are the Empidoidea of Baltic am- ber? Studia dipterologica 10: 321-327 Received: 09.01.2009 Accepted: 16.02.2009 Corresponding editor: N. Dorchin ©OZFMK Bonn zoological Bulletin | Volume 57 Issue | | pp. 91-98 Bonn, April 2010 Antennal sensilla of the rice hispa Dicladispa armigera (Olivier, 1808) (Coleoptera: Chrysomelidae) Munna Rani Saikia!; Lakshmi Kanta Hazarika!. 2; Surajit Kalita! & Ramen Lal Baishya! ‘Assam Agricultural University, Department of Entomology, Jorhat-785 013, Assam, India; 2E-mail: Ikhazarika@aau.ac.in Abstract. Number and types of sensilla on each antennal segment of male and female adult rice hispa Dicladispa armigera (Olivier) (Coleoptera: Chrysomelidae) were determined based on light and scanning electron microscopic observations. The males had a significantly greater total number (@ 1828.11) of sensilla than females (@ 1764.43). Five types of sen- silla, namely, sensilla chaetica, sensilla trichodea I, sensilla trichodea II, sensilla basiconica and pit or coeloconic sensil- la were distinguished in both sexes. Sensilla trichodea I and II were distributed over the entire length of the antenna, whereas sensilla chaetica were observed only on the apical five flagellomeres. Methoprene affected antennal morpholo- gy by producing two-clubbed antenna (additional one at the 34 flagellomere) and alteration in the sensilla of the last fla- gellomere. Key words. Rice hispa, sensilla chaetica, sensilla trichodea I, sensilla trichodea II, sensilla basiconica, methoprene. INTRODUCTION The rice hispa Dicladispa armigera (Olivier, 1808) (Coleoptera: Chrysomelidae) occurs in South East Asia and Africa, and is one of the major pests of rice in many rice growing states of India (Deka & Hazarika 1996; Palaszek et al. 2002; Islam et al. 2004; Hazarika et al. 2005). It causes considerable damage to vegetative stages of rice resulting in yield loss of 28% in India (Nath & Dutta 1997), between 20-30% in Nepal (Dhaliwal et al. 1998) and up to 52% in deepwater rice in Bangladesh (Islam 1989); however, it may be as high as 100% in the rice transplanted post flood in Assam (Hazarika 2005). In order to manage this pest, attempts were made to identi- fy pheromones in this insect, with mixed results (Deka & Hazarika 1997). Pheromones are not only used for survey and surveillance of insect pest but also used to manage them. During the last three decades, insect communication through antennal sensilla has received substantial inter- est (Rao et al. 1990; Kumar et al. 1995; Axtell 1999). Gen- erally, antennae are covered with huge numbers of sen- silla, relevant as sensory organs (Chapman 1982). Anten- nal sensilla are involved in host recognition and mate or microhabitat choice by pheromone- thermo- and hygrore- ' ception (Hazarika & Bardoloi 1998; Chen et al. 2003; Ploomi et al. 2003; Marttje et al. 2004). A number of studies has been conducted on the sensilla of other coleopteran insects like flea beetles, Phyllotetra crucifer- ae (Goeze, 1777), Psylliodes punctulata Melsheimer, 1847, P. affinis (Paykull, 1799) and Epitrix cucumeris (Harris, 1851) (Ritcey & McIver 1990); however, current- ly there is no information available describing the sensil- la of the rice hispa. Accordingly, the present study was un- dertaken in order to determine the number and types of sensilla on each antennal segment of male and female adults of D. armigera. MATERIALS AND METHODS For determination of number of sensilla, male and female antennae of field collected adults were fixed separately in carnoy-lebrun fixative for 30 min and washed for 10 min- utes in each of the 30%, 50% and 70% alcohol. The an- tennae were then allowed to remain for twenty minutes in each of the 90% and absolute alcohol, after which they were again passed through xylene and cleared in clove oil. These were then mounted in DPX. Sensilla were observed under the compound microscope at 100X, 400X and 1000X magnification in oil and were counted on each seg- ment following the method of Ramaswamy & Gupta (1981). 92 Munna Rani Saikia et al. Table 1. Estimated number (Mean+S.E.) of sensilla on each antennal segment of adult male and female D. armigera. Segment Male Female t-value Scape 25.25+0.29i 21.23+0.37i 8.95** Pedicel 28.45+0.32h 24.99+0.28h 8.77** Flagellum (FI) 1M Il 19.28+0.25k 18.02+0.29) 2.95** F] 2 22.48+0.30i 21.48+0.30! 2.48 Fl 3 33.66+40.328 26.69+0.308 15.15** Fl 4 61.67+£0.65f 56.77+0.38 6.82** Fl 6 265.20+0.414 258.36+0.494 9.7 9** FL 302.26+0.33°¢ 295.73+0.40¢ 11.99+« FI8 336.11+0.28> 332.84+0.39> 8.15+* FI9 490.84+0.534 485.07+0.464 7.A47** S- Ed) @) 0.05 0.52 C.D. 905 Ost 1.04 SE = standard error, sample size = 50. Means within columns are separated by Duncan’s Multiple Range Test (DMRT) at p < 0.05. Means followed by the same letter shown in superscript(s) are not significantly different; means within the rows followed by * and ** are significantly different at p < 0.01 and p < 0.05, respectively (Student t-test; t-values are shown against each pair; d.f. = 49). For determination of types of sensilla, the antennae were fixed for six hours in carnoy-lebrun fixative and mount- ed in DPX (Schafer & Sanchez 1976). Based on the mor- phology of the sensilla, sensilla were classified into types. Sizes of these sensilla were measured with the ocular mi- crometer and readings were converted into um. We also bleached the antennae in 2% hydrogen peroxide for 24 hr. Permeable areas of antennae were examined under bright field illumination using crystal violet method (Slifer 1960). One hundred 6—12 h old pupae were treated with 5 ppm methoprene (Altosid SE, 62.5% RS methoprene, Zoecon, Palo Alto, CA). Twenty adultoids were randomly select- ed for this study. Scanning electron microscopy studies of the antennae of the adults and adultoids were undertaken by following the method described in Hazarika & Bardoloi (1998). Anten- nae were dissected out of the head and cleaned in distilled water, which were then fixed in buffered glutaraldehyde. They were dehydrated by passing through a series of ace- tone starting from 30% to 100%. On drying, antennae were placed on stubs using double sided, scotch tapes and coat- ed with gold-palladium in a sputterer (JEOL, JFC 1100, Japan) for 5—10 min. The specimens were scanned in a scanning electron microscope (JEOL, 35-CF, Japan) at 15 KV and photographs were taken for each of the specimens. Bonn zoological Bulletin 57 (1): 91-98 RESULTS The clubbed antennae of adult specimens consisted of scape, pedicel and nine sub-segmented flagellum, each sub-segment is called flagellomere, the first flagellomere being the longest (0.30+0.02 mm) while eighth is the shortest (0.15+0.02 mm). A spine is present on the ven- tral surface of the scape though the rest of the antenna is free from such spines. Scanning electron microscopic stud- ies revealed the presence of scales on the scape, pedicel and first to sixth flagellomere. The numbers of sensilla in male and female adults of D. armigera are shown in Tab. 1. In male and female antennae, the distal segments are densely covered with sensilla. There is a distinct differ- ence in shape of the 9' flagellomere between the male and female. In each segment as well as sub-segments, the male adults had a significantly greater number of sensilla than its female counterpart (Tab. |) except on the fifth flagel- lomere, where it was reverse. The mean number of sensilla per unit area is shown in Tab. 2. Both in male and female, the maximum population was observed on the eighth and ninth flagellomere (7.11+0.03 mm and 8.89+0.03 mm, respectively) and the lowest was observed on the scape (0.37+0.01 mm? and 0.29+0.01 mm, respectively). Though density of sensilla on each segment was significantly higher in the male, but on the fifth to ninth flagellomere, it was reverse (Tab. 2). ©ZFMK Antennal sensilla of the rice hispa Dicladispa armigera (Olivier, 1808) 93 Table 2. Mean+S.E. density of sensilla (number/mm2) on each antennal segment of adult male and female D. armigera. Segment Male Female t-value Scape 0.37+0.01: 0.29+0.01) 10.28+* Pedicel 0.60+0.0 1 0.48+0.01! 9.45+** Flagellum (FI) FL 0.42+0.011 0.35+0.01) 4.35% 1 0.63+0.025 0.57+0.014 2.80* 13) 0.96+0.012 0.81+0.012 7.77 Fl 4 2.03+0.02f 2.09+0.03f -1.94NS BLS 4.49+0.02f 5.26+0.02¢ -26.47** Fl 6 5.98+0.164 6.30+0.044 -1.89NS ly 6.15+0.02¢ 7.65+0.03° -48.52** F118 6.95+0.03> 7.75+0.03° -17.18** FI 9 7.11+0.034 8.89+0.034 -42.61** S-Ed: @) 0.07 0.03 C.D. 9.95 0.14 0.07 SE = standard error, sample size = 50. Means within columns are separated by Duncan’s Multiple Range Test (DMRT) at P < 0.05. Means followed by the same letter shown in superscript(s) are not significantly different. Means within the rows are followed by * and ** are significantly different at p < 0.01 and p < 0.05, respectively (Student t-test; t-values are shown against each pair; d.f. = 49). Table 3. Size (Mean+S.E., length x width in 1m) of sensilla trichodea I on each antennal segment of adult male and female D. armigera. Segment Male Female t-value Length Width Scape 41.46+0.032 x 1.86+0.02«4 39.52+0.044 x 1.83+40.02¢4 41.07** -2.27* Pedicel 39.23+0.08° x 1.80+0.014 38.50+0.07¢ x1.74+0.02¢ 13.70** 2.46% Flagellum (F1) ell I 36.50+0.095 x 1.74 +0.02¢ 36.56+0.05f x 1.7440.02¢ -0.53NS 0.01NS Fl 2 37.18+0.078 x 1.81+0.01¢4 35.21+0.05: x 1.81+0.014 S20 ies 0.42NS BS 37.46+0.07£ x 1.84+0.02¢4 35.80+0.03" x 1.83+40.024 23.55** OLISNS Fl 4 37.84+0.12¢ x 1.81+0.01¢¢ 36.37+40.168 x 1.84+0.02¢4 9.53** -1.08NS FS 37.87+0.18¢ x 1.85+0.01¢ 38.00+0.05¢ x 1.86+0.02>¢ -0.70NS -0.55NS F16 38.00+0.074 x 1.89+0.01° 38.10+0.044e x 1.90+0.02a 1.91NS 0.31NS lei 7 38.22+0.064 x 1.90+0.02° 38.22+0.044 x 1.89+40.014 0.01NS 0.41NS F18 38.51+0.06¢ x 1.92+0.02° 38.52+0.04> x 1.99+0.01a> 26.17NS 0.82NS F19 38.54+0.07¢ x 1.96+0.014 38.63+0.03> x 1.92+0.014 24.52NS 6.56** S:Ed. ) 0.13 0.02 0.09 0.02 C.D. 0.05 0.25 0.04 0.18 0.05 SE = standard error, sample size = 50. Means within columns are separated by Duncan’s Multiple Range Test (DMRT) at p < 0.05. Means followed by the same letter shown in superscript(s) are not significantly different. Means within the rows are followed by * and ** are significantly different at p < 0.01 and p < 0.05, respectively (Student t-test; t-values are shown against each pair; df. = 49). Bonn zoological Bulletin 57 (1): 91-98 ©OZFMK 94 Munna Rani Saikia et al. Table 4. Estimated number (Mean+S.E.) of sensilla trichodea I on each antennal segment of adult male and female D. armigera. Segment Male Female t-value Scape DES EEOIS 1.88+0.42) 1.29NS Pedicel 5.81+0,.52hi 5.50+0.50h 0.46NS Flagellum (F1) FI 1 3.93+0.501 3.45+0.2 14 0.83NS F2 6.44+0.48h 5.02+0.47h 2.02NS las} 12.72+0.628 7.6940.588 7.61 ** Fl 4 21.82+0.66f 11.46+0.52£ ISI FI 5 141.77+0.66¢ 137.22+0.82¢ 3.96** FI 6 155.43+0.664 176.46+0.844 -16.56** BL, 181.34+0.79¢ 182.43+0.82¢ -1.13N8 FI8 209.44+41.97> 204.10+0.64> 2.74NS FL9 235.03+0.694 198.29+0.768 Silo S.Ed. (4) ih alkg/ 0.89 CAD enos 232 1.76 SE = standard error, sample size = 50. Means within columns are separated by Duncan’s Multiple Range Test (DMRT) at p < 0.05. Means followed by the same letter shown in superscript(s) are not significantly different. Means within the rows are followed by * and ** are significantly different at p < 0.01 and p < 0.05 probability level, respectively (Student t-test; t-values are shown against each pair; d.f. < 49). Table 5. Size (Mean4 armigera.. tS.E., length x width in pm) of sensilla trichodea II on each antennal segment of adult male and female D. Segment Male Female t-value Length Width Scape 38.11+40.234 x 1.73+0.01¢4 36.52+0.052 x 1.70+0.03¢ef 6.77** 0.87NS Pedicel 36.52+0.09> x 1.71+0.014 34.75+0.04> x1.68+0.02¢¢ 20.00+* 1-33NS Flagellum (FI) Fl 1 30.03+0.05¢ x 1.69 +0.01¢ 29.12+0.04f x 1.6740.01 11.60+* 1.10NS Fl 2 31.45+40.014 x 1.73+0.01¢4 31.04+0.064 x 1.73+0.014e 5.93** 0.31NS FI 3 32.06+0.09¢ x 1.72+0.01¢4 31.73+0.04¢ x 1.74+0.01¢4 2.83+* =1:59NS Fl4 32.15+40.10¢ x 1.7340.024 30.04+0.04¢ x 1.74+0.02¢4 22.25** -0.46NS FS 25.05+0.12i x 1.73+40.01¢4 26.51+0.065 x 1.78+0.02ab¢ -8.97** -1.64NS Fl 6 27.73+0.128 x 1.75+0.01¢ 26.83+0.078 x 1.74+0.02be4 7.44** -0.13NS Ie 28.24+0.05! x 1.7840.01 25.01+0.04' x 1.78+0.0 1abe 40.29%* -0.43NS FI8 26.17+0.074 x 1.80+0.015 24.30+0.04) x 1.79+0.01a> 26.57** 1.75NS FIL9 24.52+0.06) x 1.87+0.014 24.04+0.03* x 1.80+0.014 6.77** 8.11** SsEds@) 0.15 0.01 0.07 0.02 C.D. 9.05 0.29 0.02 0.13 0.04 SE = standard error, sample size = 50. Means within columns are separated by Duncan’s Multiple Range Test (DMRT) at p < 0.05. Means followed by the same letter shown in superscript(s) are not significantly different; means within the rows followed by * and *« are significantly different at p < 0.01 and p < 0.05, respectively (Student t-test; t-values are shown against each pair; d.f. = 49). Bonn zoological Bulletin 57 (1): 91-98 ©OZFMK Antennal sensilla of the rice hispa Dicladispa armigera (Olivier, 1808) 95 Table 6. Estimated number (Mean+S.E.) of sensilla trichodea II on each antennal segment of adult male and female D. armige- ra Segment Male Female t-value Scape 22.61+0.49! 19.3140.698 4.10** Pedicel 22.92+0.56f 19.63+0.598 4.27** Flagellum (Fl) Jell Il 15.61+0.675 14.76+0.465 0.38NS BZ 16.01+0.555 16.33+0.53h -0.44NS Jel 3} 20.10+0.538 20.10+0.628 -0.01NS Fl 4 39.88+0.61¢ 45.37+0.74! -5.55** FI 5 63.90+0.614 64.68+0.66° -0).84NS Fl 6 73.66+0.61¢ 70.81+0.624 -2.52* ell 7 83.21+0.62> 79.10+0.70¢ 3.28" F18 83.37+0.84> 90.75+0.82° Els lal ® [36975200172 156.84+0.732 -19.07+* S.Ed: (4) 0.89 0.93 C.D. 0.05 1.76 1.84 SE = standard error, sample size = 50. Means within columns are separated by Duncan’s Multiple Range Test (DMRT) at p < 0.05. Means followed by the same letter shown in superscript(s) are not significantly different; means within the rows followed by * and ** are significantly different at p < 0.01 and p < 0.05, respectively (Student t-test; t-values are shown against each pair; d.f. = 49). Table 7. Size (Mean+S.E., length x width in um) of sensilla chaetica on each antennal segment of adult male and female D. ar- migera. Segment Male Female t-value Length Width Scape — — = = Pedicel — ~ — = Flagellum (F1) Fl — _ = = FL2 — = _ = Fl 3 = = — - Fl 4 — - — - ell) 30.15+0.19¢ x 2.70+0.024 32.24+0.05¢ x 2.74+0.014 -22.17 ** Play Fl 6 33.41+0.064 x 2.86+0.02° 32.99+0.05¢4 x 2.89+0.05¢ 7.26 ** -0.64NS F17 34.11+0.06¢ x 2.94+0.02 33.50+0.04¢ x 2.85+0.02° AQ ee 3.83 ** F1 8 37.57+0.06° x 3.00+0.03° 38.40+0.04> x 2.99+0.03> -11.92 + 0.28NS FI9 3871501062 x3 122-0022 38.70+0.034 x 3.10+0.032 -6.67 ** 0.67NS S-Ed: @) 0.09 0.03 0.06 0.04 C.D. 9.95 0.18 0.06 0.12 0.08 SE = standard error, sample size = 50. Means within columns are separated by Duncan’s Multiple Range Test (DMRT) at p < 0.05. Means followed by the same letter shown in superscript(s) are not significantly different; means within the rows followed by * and ** are significantly different at p < 0.01 and p < 0.05, respectively (Student t-test; t-values are shown against each pair; d.f. = 49). Bonn zoological Bulletin 57 (1): 91-98 ©OZFMK 96 Munna Rani Saikia et al. Table 8. Estimated number (Mean+S.E.) of sensilla chaetica on each antennal segment of adult male and female D. armigera. Segment Male Female t-value Scape — - — Pedicel — _ = Flagellum (FI) Fld — — _ Fl 2 = = — las) = = = Fl 4 = = ~ ES 14.76+0.604 21.35+0.634 -6.33+* F1 6 36.58+0.61¢ 10.83+0.62¢ 29.58** FL 37.84+0.58¢ 32.34+0.61¢ 6.05** FL8 42.55+0.66> 37.99+0.68> 5.45%» ial’) 119.01+0.79 130.15+0.708 -12.06** S.Ed. (+) 0.92 0.91 C.D. 9.05 1.84 1.82 SE = standard error, sample size = 50. Means within columns are separated by Duncan’s Multiple Range Test (DMRT) at p < 0.05. Means followed by the same letter shown in superscript(s) are not significantly different; means within the rows followed by * and ** are significantly different at p < 0.01 and p < 0.05, respectively (Student t-test; t-values are shown against each pair; d.f. = 49). Based on morphology, sensilla trichodea (ST) I, ST II, sen- silla chaetica (SC), sensilla basiconica (SB) I and coelo- conic sensilla were identified in both sexes of the insect. The ST I and II were numerous and observed over the en- tire length of the antennae whereas SC were observed on- ly on the apical five flagellomeres. In addition, some sen- silla suspected to be thermoreceptors (Tr) were also ob- served on scape, pedicel and some flagellomeres. The ST I pointed distally and curved towards the anten- nal shaft. It is a slender structure, which tapers gradually into a very sharp point at the distal end. In males, the lengths of ST I varied from 36.50 + 0.09 um to 41.46+ 0.03um. Likewise, their widths varied from 1.74+ 0.02 um to 1.96+ 0.01 um. In female, the lengths varied from 35.21+ 0.05 um 39.52+ 0.04 tm and their widths varied from 1.74+ 0.02 um to 1.99+ 0.01 tm (Tab. 3). The high- est population of ST I was observed on the ninth flagel- lomere (235+ 0.03+ 0.69) and the lowest was observed on the scape (2.51+ 0.37) in males, whereas in females, the highest population was observed on the eighth flagel- lomere (204.10+ 0.64), and the lowest was observed on the scape (1.88 + 0.42) (Tab. 4). The ST II were also similar to ST I except that they were unaffected when a solution of crystal violet was applied. It might be due to non- permeability of the senisilla to crystal violet. In males, the lengths of ST II varied from 24.52+0.06 to 38.11+0.23 um (Tab. 5). Likewise their Bonn zoological Bulletin 57 (1): 91-98 widths varied from 1.69+0.01 to 1.87 + 0.01 um. In fe- males, their lengths varied from 24.04 + 0.03 to 36.52 + 0.05 um and their widths varied from 1.67+0.01 to 1.80+0.01 um. The highest number of ST II was observed on the ninth flagellomere (136.75 + 0.77) and lowest on the first fla- gellomere (15.6+0.67) in males. Likewise, in females al- so the highest number of ST II was observed on the ninth flagellomere (156.84 + 0.73) and the lowest was observed on the first flagellomere (14.76 + 0.46) (Tab. 6). The SC were pointed distally and projected outward from a socket at an approximate angle of 50°, thick-walled and longitudinally grooved. They were restricted to the termi- nal five flagellomeres and were unaffected when a solu- tion of crystal violet was applied. In males, their lengths varied from 30.15+0.09 to 38.17+0.06 um (Tab. 7), where- as in females they varied from 32.24 +0.05 to 38.70+ 0.03 um. In male and female, the highest number of SC was observed on the ninth flagellomere and the lowest was ob- served on the fifth flagellomere (Tab. 8). A few sensilla basiconica (SB) were present in different flagellomere of D. armigera. SB are smaller than ST measuring 5—6 um, wall being porous. Pit or coeloconic sensilla are also present on the antennae, however, details of their structure were not studied. ©ZFMK Antennal sensilla of the rice hispa Dicladispa armigera (Olivier, 1808) 97 After application of growth regulator, methoprene, some deformities not only on the antennal structure but also on the sensillar morphology were observed; the 4" flagellom- ere got deformed. This is very prominent on the tip of the ninth flagellomere where SC were observed to be disori- ented (Baishya 1992). DISCUSSION Densely covered distal segments of the antennae as ob- served here are also present in many other insects like Blatella germanica (Linnaeus, 1767) (Ramaswamy & Gupta 1981, Wheeler & Gupta 1986), Croesia curvala (Kearfott, 1907) (Langmaid & Seabrook 1985), Bootet- tix argentatus Bruner, 1890 (Chapman & Fraser 1989), Geotrupes auratus Motschulsky, 1858 (Inouchi et al. 1987) and Homoeosoma electellum (Hulst, 1887) (Faucheux 1995), and four hemipteran species (Usha Rani & Madhavendra 2005). Palpation conducted with the dis- tal segments of antenna may provide an explanation for this pattern. On the scape, pedicel and three flagellomeres (first to third), males had significantly greater number of sensilla per unit area than females. However, from the fourth to ninth flagellomere, females had a greater number of sen- silla per unit area than males; a similar pattern was report- ed by Ritcey & McIver (1990) in case of Psylloides punc- tulata and P. affinis. A study by Usha Rani & Madhaven- dra (2005) suggested that sensillae in scape and pedicel may not be used in sensory perception, which remains however to be tested critically. Similar to Dicladispa armigera, terminal segments of an- tennae covered with ST I and ST II were also reported in Homoeosoma electellum (Faucheux 1995), Phothori- maea operculella (Zeller, 1873) (Sharaby et al. 2002), He- licoverpa armigera (Hiibner, 1808) (Wang et al. 2002). Bromely et al. (1980) stated that the ST at the antennal tips of aphids have a contact chemosensory function, and may be involved in gustation of the plant surface, which might be the case in the insect studied here as well. The population of ST I was highest on the ninth flagel- lomere in males, whereas in the female it was on the eighth flagellomere. This kind of variation in ST I population may be associated with specific function performed by the sen- silla. Similar cases were also recorded in Trichoplusia ni (Hiibner, 1803) (Mayer et al. 1981), Grapholitha moles- ta (Busck, 1916) (George & Nagy 1984), Hypera posti- ca (Gyllenhal, 1813) (Bland 1981). However, in case of both males and females, the ST II population was high- est on the ninth flagellomere, which was also reported by Bland (1981) in Hypera postica, by Kapoor (1985) in Bonn zoological Bulletin 57 (1): 91-98 Paragnetina media (Walker, 1852) and by Ritcey & Mclv- er (1990) in flea beetles. SC is a common type usually en- countered in many insects (Ilango 2000). The presence of SC on antennae was also reported by Ritcey & Mclver (1990) in the flea beetles, P. cruciferae, P. punctulata, P. affinis and E. cucumeris. As Hazarika & Baishya (1996) showed, application of methoprene induced morphogenesis in D. armigera; how- ever, only the fourth and the last flagellomere were affect- ed. The cause of the selectivity of this effect is unclear up to now. Hormonal regulation of antennal sensilla is report- ed for many insects (Wheeler & Gupta, 1986; Yamamo- to-Kihara et al. 2004). Injection of a neurohormone, [His’]-corazonin, reduced the number of coeloconic sen- silla in Locusta migratoria. A similar study on D. armigera is also required since it could help to clarify the causal net- work in the development of the antennal sensilla pattern. Acknowledgements. This work was partly funded by the Na- tional Agricultural Technology Project, Indian Council of Agri- cultural Research, New Delhi. We are also grateful to Dr. G. B. Staal, Zoecon, Palo Alto, CA for providing the methoprene as a gift for experimental purpose. Scanning electron micrographs were taken at the Regional Sophisticated Instrumentation Cen- tre, North Eastern Hill University, Shillong, Meghalaya. REFERENCES Axtell RC (1999) Poultry integrated pest management: Status and future. Integrated Pest Management Revue 4: 53-73 Baishya RL (1992) Effect of insect growth regulators on the post embryonic development and reproduction of rice hispa, Di- cladispa armigera (Olivier) (Coleoptera: Chrysomelidae). M.Sc. (Agri.) Thesis, Assam Agricultural University, Jorhat, Assam Bland RG (1981) Antennal sensilla of the adult alfalfa weevil, Hypera postica (Gyllenhal) (Coleoptera: Curculionidae). In- ternational Journal of Insect Morphology & Embryology 10: 265-274 Bromely AK, Dunn JA, Anderson M (1980) Ultrastructure of an- tennal sensilla of aphids. II. Trichoid, chordotonal and cam- paniform sensilla. Cell and Tissue Research 205: 493-511 Chapman RF (1982) Chemoreception: The significance of re- ceptor populations. Advances in Insect Physiology 16: 247-356 Chapman RF, Fraser J (1989) The chemosensory system of the monophagous grasshopper, Bootettix argentatus Bruner (Or- thoptera: Acrididae). International Journal of Insect Morphol- ogy & Embryology 18: 111-118 Chen HH, Zhao YX, Kang LE (2003). Antennal sensilla of grasshoppers (Orthoptera: Acrididae) in relation to food pref- erences and habits. Journal of Biosciences 28: 743-752 Deka M, Hazarika LK (1996) Mating behavior of Dicladispa armigera (Coleoptera: Chrysomelidae). Annals of the Ento- mological Society of America 89: 137-141 Deka M, Hazarika LK (1997) Investigation of male sex pheromone in rice hispa, Dicladispa armigera (Oliv.) (Coleoptera: Chrysomelidae). Pestology 21: 24-25 ©OZFMK 98 Munna Rani Saikia et al. Dhaliwal GS, Arora R, Randhwa NS, Dhawan AK (1998) Eco- logical Agriculture and sustainable development. In: Proceed- ings of an International Conference on Ecological Agriculture: Towards Sustainable Development, Vol 1, Chandigarh, India, 15—17 November, 1997 Faucheux MJ (1995) Sensilla on the antennae, mouthparts, tar- si and ovipositor of the sunflower moth, Homoeosoma electel- lum (Hulster) (Lepidoptera: Pyralidae). Annales des Sciences Naturelles Zoologie et Biologie Animale 11: 121-136 George JA, Nagy BAL (1984) Morphology, distribution and ul- trastructural differences of sensilla trichodea and basiconica on the antennae of the oriental fruit moth, Grapholitha mo- lesta (Busck) (Lepidoptera: Tortricidae). International Jour- nal of Insect Morphology & Embryology 13: 157-170 Hazarika LK (2005) Rice hispa: How much we really know about it? In: Ramamurthy VV, Singh VS, Gupta GP, Paul AVN (eds), Gleanings in Entomology, Division of Entomology, In- dian Agricultural Research Institute, New Delhi, pp. 1-19 Hazarika LK, Baishya RL (1996) Effect of methoprene on rice hispa, Dicladispa armigera (Olivier) (Coleoptera: Chrysomel- idae). Pesticide Resarch Journal 8: 93-95 Hazarika LK, Bardoloi S (1998) Antennal and mouth part sen- silla of the Muga silkworm, Antheraea assama (Lepidoptera: Saturnidae). Sericologia 38: 55—63 Hazarika LK, Deka M, Bhuyan M (2005) Oviposition behav- iour of the rice hispa Dicladispa armigera (Coleoptera: Chrysomelidae). International Journal of Tropical Insect Sci- ence 25: 1-6 Hazarika LK, Dutta BC (1991) Reaction of rice cultivars to rice hispa. International Rice Research Newsletter 16: 14-15 Ilango K (2000) Morphological characteristics of the antennal flagellum and its sensilla chaetica with character displacement in the sandfly Phlebotomus argentipes Annandale and Brunet- ti Sensu lato (Diptera: Psychodidae). Journal of Biosciences 25: 163-172 Inouchi J, Shibuya T, Matsuzaki O, Hatanaka T (1987) Distri- bution and fine structure of antennal olfactory sensilla in Japanese dung beetle, Geotrupes auratus Mots. (Coleoptera: Geotrupidae) and Copris pecurius Lew. (Coleoptera: Scarabaeidae). International Journal of Insect Morphology & Embryology 16: 177-187 Islam Z (1989) Crop losses due to hispa beetle damage in deep water rice (DWR). International Rice Research Notes 14: 53 Islam Z, Heong KL, Bell M, Hazarika LK, Rajkhowa DJ, Ali S, Dutta BC, Bhuyan M (2004) Current status of rice pests and their management in Assam, India — a discussion with ex- tension experts. International Rice Research Notes 29: 89-91 Kapoor NN (1985) External morphology and distribution of the antennal sensilla of the stonefly Paragnetina media (Walker) (Plecoptera: Perlidae). International Journal of Insect Morphol- ogy & Embryology 14: 273-280 Kumar S, Prakash S, Rao KM (1995) Comparative activity of three repellents against bed-bugs, Cimex hemipterus Fabr. In- dian Journal of Medical Research 102: 20-23 Langmaid WM, Seabrook WD (1985). The micromorphology of the antennae of the blueberry leaf tier moth, Croesia cur- valana (Kft.) (Lepidoptera: Tortricidae). Canadian Journal of Zoology 63: 1189-1193 Maartje AK, Bleeker Hans M Smid, Adriaan C Van Aelst, Joop JA Van Loon, Luise EMVet (2004). Antennal sensilla of two parasitoid wasps: A comparative scanning electron mi- croscopy study. 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Indian Phytopathology 45: 35-38 Ramaswamy SB, Gupta AD (1981) Sensilla on the antennae, labial and maxillary palps of Blatella germanica (L.) Blatti- dae: Their classification and distribution. Journal of Morphol- ogy 168: 269-279 Ramaswamy SB, Gupta AP (1981) Effects of juvenile hormone on sense organs involved in mating behaviour of Blattella ger- manica (L.) (Dictyoptera: Blattellidae). Journal of Insect Phys- iology 27: 601-608 Rao KM, Prakash S, Kumar S, Suryanarayana MVS, Bhagwat MM, Gharia MM, Bhavsar RB (1990) Evaluation of repellent NN-diethyl-phenylacetamide treated fabrics against the mos- quitoes, Aedes aegypti and Culex quinquefasciatus (Diptera: Culicidae). Journal of Medical Entomology 28: 142-147 Ritcey GM, McIver SB (1990) External morphology of anten- nal sensilla of four species of adult flea beetles (Coleoptera: Chrysomelidae: Alticinae). International Journal of Insect Morphology & Embryology 19: 141-153 Schafer R, Sanchez TV (1976) The nature and development of sex attractant specificity in cockroaches of genus Periplane- ta 1. Sexual dimorphism in distribution of antennal sense or- gans in five species. Journal of Morphology 149: 139-158 Sharaby AM, Abdel Rahman HA, Moawad SS (2002) Sensors of the potato tuber moth, Phthorimaea operculella (Zell.) (Lepidoptera: Gelechiidae). Bulletin of the National Research Centre, Cairo. 27: 137-141 Slifer EH (1960) A rapid and sensitive method for identifying permeable area in the body walls of insects. Entomological News 71: 179-182 Usha Rani P, Madhavendra SS (2005) External morphology of antennal and rostral sensillae in four hemipteran insects and their possible role in host plant selection. International Jour- nal of Tropical Insect Science 25: 198-207 Wang GR, Guo YY, Wu KM (2002) Study on the ultrastructu- res of antennal sensilla in Helicoverpa armigera. Agricultu- ral Sciences in China 1: 896-899 Wheeler CM, Gupta AP (1986) Effects of exogenous juvenile hormone I on the numbers and distribution of antennal and maxillary and labial palp sensilla of male Blattella germani- ca (L.) (Dictyoptha: Blattellidae). Experientia 42: 57-58 Yamamoto-Kihara M, Hata T, Breuer M, Tanaka S (2004) Ef- fect of [His’”]-corazonin on the number of antennal sensilla in Locusta migratoria. Physiological Entomology 29: 73 Received: 03.07.2008 Accepted: 08.07.2009 Corresponding editor: M. Schmitt ©ZFMK Book Review 99 Christoph Moning, Thomas Griesohn-Pfleger & Michael Horn (2010). Grundkurs Vogelbestimmung: Eine Ein- fiihrung in die Beobachtung und Bestimmung unserer heimischen Vo6gel. 422 pp., Quelle & Meyer Verlag GmbH & Co, Wiebelsheim, ISBN-10: 3494014167, ISBN-13 978-3494014166, 19,95 €. Facilitating the access to birdwatching by a book is not particularly new. There are examples from anglophone countries as well as from Germany (e.g. “Végel beobacht- en” by E. Bezzel). However, “Grundkurs Vogelbestim- mung” (Basic course bird identification) is particular up to date and specific to today’s internet generation. The book is part of a new series of basic courses on field iden- tification of different taxonomic groups edited by the same publishing house. The book is divided into two major parts. The Chapters 1—12 set the general part; the second offers a surprising- ly extensive insight into all central European bird fami- lies. Single chapters are rather short, the text is interrupt- ed by boxes and figures and invites the reader to skim and browse. The general part includes everything from equipment (binoculars, literature sources, appropriate clothes etc.) to an introduction to avian behaviour, bioacoustics, and es- pecially to the Where, How and When of birdwatching. These chapters are the strengths of the book, and the ex- perience of the authors who regularly work as tour guides is quite obvious. They have a good sense of humour when combining a cold drink and illustrated bird books as nec- essary equipment to reflect their own observations (fig- ure on p. 21). However, a Wikipedia definition should nev- er be cited when explaining complex issues as the hand- icap principle (p. 76). The chapter on “Systematics and Taxonomy” is praisewor- thy. Contributing author Daniel Lingenhohl does not re- ally give an explanation of these terms, but explains on five and a half pages — necessarily only rudimentary — “‘bi- nary nomenclature” or the “biological species concept”. Some shortcomings can be overlooked here. The author exemplifies the changing character of this scientific field by some huge changes in avian higher taxa systematics. Sadly, his explanation on using italics for scientific names is not completely correct and it is even more regrettable that in some parts of the further text not even the correct parts of his remarks were respected (p. 133 ff. subfami- lies wrongly italicised, p.137 genus Branta not in italics). In the conservation chapter, concepts in Germany are com- pared to those in some neighbouring countries. Many as- pects the authors rise are certainly true but perhaps it would have been wiser to generalize a little bit less and Moning/Griesohn-Pflieger/Horn — Grundkurs Eine Einfiihrung zur Beobachtung und — Bestimmung unserer heimischen Végel give this issue more space. Feeding birds might be fun but its value for conservation is at least controversial. The fig- ure on page 125 is wrongly marked (it is a “Rotkopfwiirg- er”, “Rotstirnwiirger” is an African species). The second part is systematically ordered by Central Eu- ropean bird families although this order does not seem to be very consistent as non-passerines follow a generally rather conservative concept while passerines are rather or- dered by new findings. There are good, compact introduc- tions for every family and therein focus is on the most common birds so that beginners are not unnecessarily con- fused. Short texts and arrows in the pictures complement information from classical bird guides although some overlap cannot be denied. Graphics on the topography of many different birds increase the learning effect; obser- vation hints enrich the text. However, the authors forgot the huge population of Melodious warblers in Rhineland- Palatinate and more important one should never throw swifts in the air that were found on the ground. The au- thors may have to take the responsibility for some bro- ken bones in swifts. 100 Book Review So, what’s missing? Not much, maybe the unloved glos- sary but “carpal joint” (Carpalgelenk) and even “taxon” might not be that easy to understand for the beginner. A few more internet sources and examples from Austria and Switzerland would make a broader distribution easier. There is a growing attention for “citizen science” in or- ganismic biology. The participation of non-professionals has a long history in the study of birds — many important long-term monitoring schemes would be impossible with- out the help of these so called amateurs. Scientific soci- eties miss young people and there is hardly a scientist who Bonn zoological Bulletin 57 (1): 99-100 does not complain about lacking public awareness of his or her field. Therefore the book is not only recommend- ed for beginners but also to those who want to know how to teach their ornithological knowledge and for those Non- Ornithologists who look for stimulations how to prepare the popular science aspect of their organisms in an exem- plary way. 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Zoological Journal of the Linnean Society 154: 494-610 Sullivan J (1994) Bufo boreas. In: Fire Effects Information System (U.S. Department of Agriculture, Forest Service, Rocky Moun- tain Research Station, Fire Sciences Laboratory). Online at http://www. fs.fed.us/database/feis/animals/amphibian/bubo/all.ht ml last accessed on December 28, 2009 Sztencel-Jablonka A, Jones G, Bogdanowicz W (2009) Skull mor- phology of two cryptic bat species: Pipistrellus pipistrellus and P. pygmaeus — a 3D geometric morphometrics approach with land- mark reconstruction. Acta Chiropterologica 11: 113-126 Authors are kindly requested to make sure that sources listed in the References match those cited in the text. Names of persons who pro- vided unpublished information should be cited by including the ini- tial(s) and city as follows: “(JG Frommen, Bonn, pers. comm. 2009)”. Tables and Figures Tables and Figures should not duplicate infor- mation provided in the text. 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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? ‘Oo OND WU Contents ened det Sonnenberg, Rainer & Eckhard Busch: 3 Description of Callopanchax sidibei (Nothobranchiidae: Epiplateinae), a new species of killifish from southwestern Guinea, West Africa Nguyen, Quang Truong, Andreas Schmitz & Wolfgang Bohme: 15 Gekko ulikovskii Darevsky & Orlov, 1994: a junior synonym of Gekko badenii Szczerbak & Nekrasova, 1994 Krause, Patrick & Wolfgang Bohme: 19 A new chameleon of the Trioceros bitaeniatus complex from Mt. Hanang, Tanzania, East Africa (Squamata: Chamaeleonidae) Ullenbruch, Klaus, Olaf Grell & Wolfgang Bohme: 31 Reptiles from southern Benin, West Africa, with the description of a new Hemidactylus (Gekkonidae), and a country-wide checklist Wilhelm, Gertha, Hans L. Nemeschkal, John Plant & Hannes F. Paulus: 55 Fitness components in the relationship between Rhopalapion longirostre (Olivier, 1807) (Insecta: Coleoptera: Apionidae) and Alcea rosea (Linnaeus, 1758) (Malvaceae). Analysis of infestation balance of a herbivorous weevil and its host plant Shoorcheh, Hossein Rajael: 65 Life-history of Gnopharmia kasrunensis Wehrli, 1939 and G. colchidaria Lederer, 1870 (Geometridae, Ennominae) and their distribution in Iran, with first host-plant records for the genus StUning, Dieter: 75 Two new species of the Plutodes costatus-group from the Philippines and Indonesia (Lepidoptera, Geometridae, Ennominae) Sinclair, Bradley J.: 85 Proclinopyga ulrichi sp. nov.: the first fossil aquatic dance fly of the subfamily Clinocerinae (Diptera: Empididae) Saikia, Munna Rani, Lakshmi Kanta Hazarika, Surajit Kalita & Ramen Lal Baishya: 91 Antennal sensilla of the rice hispa Dicladispa armigera (Olivier, 1808) (Coleoptera: Chrysomelidae) Book Reviews Funfstlck HJ, Ebert A & Weiss | (2010) 18 Taschenlexikon der Vogel Deutschlands (Kathrin Schidelko) Moning C, Griesohn-Pfleger T & Horn M (2010) 99 Grundkurs Vogelbestimmung: Eine EinfUhrung in die Beobachtung und Bestimmung unserer heimischen Végel (Darius Stiels) Cover illustration: Callopanchax sidibei Sonnenberg & Busch (this volume, pp. 3-14) Ministerium fiir Innovation, ray . Fs Y, Wissenschaft, Forschung Lei bn 1Z Yi) und Technologie des Landes 5 <8S/_ Nordrhein-Westfalen @ Gemeinschaft