ds . A ES Phos | SC or ee oe WW soviet lice an hy ; eins tatu Pa Peete Bata fan rer TEEN na Karate Y CATA uf ae Ve ENT MANET WA Lh yeh LANA ol Lin man 7 AENA ty! ayia) N tthe VTEC aH send ey y AT KT vi ELTERN AAA RA SAR PINTS a INEA a SoA arena ARAN ar E Are) Hiern ELMER TO Ae iy 3 ’ : A N Ser : ONION ate tle mt ma NAN Co ANTONIA RETINA re y ARO NA ees et nen iat Sere Loi at Vener > x idee too ya) hi ur WITT suet yay MUSH NAN ES suede oped E mr x a Mee Pea hers in E ALEA VOUT y AS ay! N are Soe ARN SN N) ANN Rad Me a TE N mr 4 = cs 9 ‘ 1 . er =. Go Ñ = . . x we “ . . 7 ‘ ' A ‘ : ’ = | “ 5 — + . \ 1 1 ‘ ; o U i “u i i o o he i ' i ‘ ‘ YO ES de = 4 ‘ “a ' = " 5 o AR B . E a D i a . voi . “ en Bonner zoologische . Beiträge Index Volume 53 Editor-in-chief Michael Schmitt Bonn, Germany Editors Renate van den Elzen Bernhard A. Huber Gastav Peters Bradley J. Sinclair Dieter Stüning Bonn, Germany Bonner zoologische Beitráge vol. 53, Index ASPOCK, Horst & ASPOCK, Ulrike: Neuropterologische Beiträge in den Werken von Wilhelm Friedrich VON GLEICHEN, genannt KRUSSWORM ITA Ia 13-36 ASSING, Volker: On the Taxonomy and Biogeography of Stenus (s. str.) ervthrocnemus Eppelsheim and Related Species (Insecta: Coleoptera: Staph ylimidae) en... une rior 303-310 BISCHOFF, Inge, ECKELT, Esther, & KUHLMANN, Michael: On the Biology of the Ivy-Bee Colletes hederae Schmidt & Westrich, 1993 de o A A O O podias 27-36 BOHME, Wolfgang & KOHLER, Jórn: Do Endings of Adjective Flectible Species Names Affect Stability? A Final Note on the Gender of Podarcis Wagler, 1830 (Reptilia, Lacertidae)...........o.ooonnocnnonnconncoonnconncnanonnnos 293-295 DEUSCHLE, Jiirgen & GLUCK, Erich: Reproductive Types and Mobility of Carabid Assemblages: Erfects of Landuse Intensity of Extenstvely Managed Orchards siii ado ne 311-321 FRANZEN, Michael & HEINZ, Walter: Morphology, Genitalia, and Natural History Notes on the Enigmatic Tiger Beetle, Mantica horni Kolbe, 1696 (Coleoptera; Cicinidelidae) ii e 297-301 GASSMANN, Dirk: The Phylogeny of Southeast Asia and Indo-Pacific Calicnemiinae (Odonata, Platyenemidida iia ER 37-80 HOFMANN, Axel: Neue Zygaena-Taxa aus Südosteuropa, Vorder- und Zentralasien (Lepidoptera Lead a 81-97 HUBER, Bernhard, PÉREZ G., Abel, BAPTISTA, Renner L. C.: Leptopholcus (Araneae: Pholcidae) in Continental America: Rare Relicts m Low Precipitation Areas aa ee 99-107 HUNDSDOREER, Anna K.: First Record of Hyles dahlii (Geyer, 1827) (Lepidoptera: Sphingidae) Ode Arca Maida nee NANO 109-110 JASCHHOF, Mathias: Rogambara and Cabamofa, Two New Genera of Enigmatic Sciaroids from Costa Rica (secta: Diptera: Sclatolded) sita A 323-332 KUPER, Wolfgang, WAGNER, Thomas € BARTHLOTT, Wilhelm: Diversity Patterns of Plants and Phytophagous Beetles in Sub-Saharan Africa .....ooncnccconnonionnncnncnnnncananerinrano 283-289 LOURENCO, Wilson R.: A New Genus and Species of Scorpion from Afghanistan (Scorpiones, Buthidae).....ooooooonnoccnnnccinoncccconacinanonos 111-114 MEY, Wolfram & SPEIDEL, Wolfgang: Two New Species of Eoophyla Swinhoe, 1900 from Continental South East Asia (Lepidoptera: Crambidaes Acentropimae)' isc nee 115-119 MISOF, Bernhard, KLUTSCH, Cornelya F. C., NIEHUIS, Oliver & PATT, Alexandra: ©fPhenotypes:and Genotypes: Two Sides of One Coin in Taxonomy? ..........:scscccssssssesessssenesersnsazerseceses ROSLER, Herbert, ZIEGLER, Thomas, VU, Ngoc Thanh, HERRMANN, Hans-Werner & BOHME, Wolfgang: A New Lizard of the Genus Gekko Laurenti, 1768 (Squamata: Sauria: Gekkonidae) from the Phong Nha — Ke Bang National Park, Quang Binh Province, Vietnam .............ecceescseeeeeeseeeeseeeeeseeeaeens = SAUER, Klaus Peter & KULLMANN, Harald: Analyse der biologisch-ökologischen Ursachen der Evolution der gastroneuralen Metazoa — Mestense meuphylogeBetischen Hypatbese a. nen ee ea SCHINTLMEISTER, Alexander: Peridea clasnaumanni spec. nov. (Lepidoptera: Notodontidae) aus ChIMA ...oooooccnnnocccononononancnonancnononcconnoss SCHMITT, Michael: @lasıM. Naumann (26.06.1939 — 15.02.2004) - in MeMOriaM. ..........0.cccccececdeecedeccecessccaceeesdeccesecesseeeceetecess SCHUNKE, Anja C. & HUTTERER, Rainer: The variance of variation: Geographic Patterns of Coat Colouration in Anomalurops and Anomalurus (Mammalia, Rodentia, Anomaluridae).................unnennenenneeeeennnnnansenneennnnsnnnnneennnnnnnnnnneeeenn SCOBLE, Malcolm J.: Issue in Delimiting Genera in Invertebrates: an Example from the Lepidoptera (Maeanını Geometndae: EAM Ominae) .....2.cccccs.o00cc-05etceeessseseseeseececssveescoeusescsebeceedugenvassesthocdncess¥esveseecusasas SINCLAIR, Bradley J. & SAIGUSA, Toyohei: Revision of the Trichoclinocera dasyscutellum Group from East Asia MDrsas mato A A SONNENBERG, Rainer & BLUM, Thomas: Aphyosemion (Mesoaphyosemion) etsamense (Cyprinodontiformes: Aplocheiloidei: Nothobranchiidae), a New Species from the Monts de Cristal, Northwestern Gabon.......................- SPEIDEL, Wolfgang & STUNING, Dieter: Ambia naumanni sp. n., a New Species of Musotiminae from Yunnan Edie, (Camillo ac ae O A TNT UME Aer erp SPEIDEL, Wolfgang, BUCHSBAUM, Ulf & MILLER, Michael A.: A New Paracymoriza Species from Lombok (Indonesia) (Lepidoptera, Crambidae)............................... VANE-WRIGHT, Richard I. & BOPPRE, Michael: Adult Morphology and the Higher Classification of Bia Hiibner PUBS ICL Roem NINN el INAS Veg vances tects suc 0 PORRO OPPR E o WAGNER, Thomas: Revision of the vincta Species-group of Monolepta Chevrolat, 1837 from Africa, Arabia and the Near East (Coleoptera: Chrysomelidac, Galerucinae) ...ici.:.....tcassccenssedecsecsacseececsssessessesaeees ZWICK, Peter & AREFINA, Tatyana: The Net-Winged Midges (Diptera: Blephariceridae) of the Russian Far East..................n Publication dates: No. 1/2: 30 June, 2005, No. 3/4: 30 December, 2005 ¡93 149-163 165-167 169-185 187-191 193-209 211-220 7 ‘ * ’ o = ] * A . Ñ I 0 a ñ . 1 o B . j x re . =e = 1 N ; : ñ LA a ; y o 7 : 7 P . a Y A y o y 7 . o o % y . : eet er . an 5 9 mM . po ñ % A va as . 13 so. ] er y en u aa . on Ss j o . Be ry, u . EN ee ' i : 2 1 . + a Dr . ’ = ' ye \ ' a E A . ñ ñ Meo o \ T ‘ 1 ca ' a = ñ M $ \ A Be, sa : qa . , ‘ i y , 1 h \ ua 0 u . \ i ys i) 7 ’ “ ~ At : ary ye ‘ . . o ñ wt “ ‘ A oo, 7 o . . ’ © : u ' ie a s 5 A ag o 7 ou a > f ; . 7 . ef , + A Dn UN . iy y . E - o ' : ñ 2 y , ‘ ' : . ol er ys 7 7 El ‘ y u ' ñ 1 \ \ \ ñ A ' . { B ' , vs ER u: RAN, Pr E of fj y 7 Ñ u o . \ a + y ] A ui rs 5 ' . 2 7 > . ] ; ' 7 Y u og Y , Y « e u - 9 . . | . : ae \ “ 1 : iw = 5 ' ae \ ] La Re ' : en Ss ee \ \ ‘ x . 1: 1] ‘ \ ‘ \ 1 en - a = a er ice PT 7 Bonner | zoologische - Beiträge I In memoriam | Clas M. NAUMANN (1939-2004) Herausgegeben vom Zoologischen Forschungsmuseum Alexander Koenig, Bonn Band Dr, Heft 1/2 2005 2009 my Leibniz Gemeinschaft inis für Wisse ft un: Bonner zoologische Beitráge werden publiziert im Eigenverlag Zoologisches Forschungsmuseum Alexander Koenig, Adenauerallee 160, D-53113 Bonn (Germany) Die Zeitschrift erscheint mit vier Heften im Jahr, zum Preis von 11,50 € je Heft bzw. 46,- € je Band incl. Versand. Korres- pondenz betreffend Abonnement, Kauf oder Tausch bitten wir zu richten an die Bibliothek, Zoologisches Forschungsmu- seum Alexander Koenig, Adenauerallee 160, D-53113 Bonn (Germany). E-mail: d.steinebach.zfmk@uni-bonn.de. © Zoologisches Forschungsmuseum Alexander Koenig (ZFMK), Bonn, Germany ISSN 0006-7172 Druck: Druck Center Meckenheim Bonner zoologische Beitráge Editorial Board (in brackets: editorial competence) Prof.Dr. Michael SCHMITT (editor-in-chief, Coleoptera, theory), Tel./Fax: +49 228-9122 286, E-mail: m.schmitt@uni-bonn.de Dr. Renate VAN DEN ELZEN (Vertebrata, except Mammalia), Tel. +49 228-9122 231, Fax: +49 228-9122 212, E-mail: r.elzen.zfmk@uni-bonn.de Dr. Bernhard A. HUBER (Invertebrata, except Insecta), Tel. +49 228-9122 294, Fax: +49 228-9122 212, E-mail: b.huber.zfmk(@uni-bonn.de Dr. Gustav PETERS (Mammalia, bioacoustics), Tel. +49 228-9122 262, Fax: +49 228-9122 212, E-maii: g.peters.zfmk@uni-bonn.de Dr. Bradley J. SINCLAIR (Diptera, language advisor), Tel. +49 228-9122 292, Fax: +49 228-9122 212, E-mail: b.sinclair.zfmk(@uni-bonn.de Dr. Dieter STUNING (Insecta, except Coleoptera and Diptera), Tel. +49 228-9122 220, Fax: +49 228-9122 212, E-mail: d.stuening.zfmk(@uni-bonn.de Editorial office: Adenauerallee 160, D-53113 Bonn, Germany Advisory Board Prof.Dr. Miguel Angel ALONSO-ZARAZAGA, Museo nacional, Ciencias Naturales, E-28006 Madrid, E-mail: zarazaga@mnen.csic.es Prof.Dr. Ulrike ASPÓCK, Naturhistorisches Museum, 2. Zoologische Abteilung (Insekten), Postfach 417, A-1014 Wien, E-mail: ulrike.aspoeck(@nhm-wien.ac.at Prof.Dr. Paolo AUDISIO, Universita di Roma ,,La Sapienza”, Dip.Biol.Anim. e dell'Uomo (Zool.), Viale dell'Universita 32, 1-00185 Roma, Tel.: +39 6-49914744, E-mail: paolo.audisio@uniromal.it Prof.Dr. Aaron M. BAUER, Villanova University, Department of Biology, 800 Lancaster Avenue, Villanova, PA 19085-1699, USA. Tel. +1-610-519-4857, Fax +1-610-519-7863, E-mail: aaron.bauer(@villanova.edu Dr. Jiirgen HAFFER, Tommesweg 60, D-45149 Essen, Tel.: +49 201-710426, E-mail: j.haffer@web.de Prof.Dr. Andreas J. HELBIG , Ernst-Moritz-Arndt-Universität, Vogelwarte Hiddensee, D-18565 Kloster, Tel.: +49 38300-212, Fax: +49 38300-50441, E-mail: helbig@mail.uni-greifswald.de Dr. Jeremy D. HOLLOWAY, The Natural History Museum, Department of Entomology, Cromwell Road, London SW7 5BD, U.K., E-mail: j.holloway@nhm.ac.uk Dr. Marion KOTRBA, Zoologische Staatssammlung, Diptera, Miinchhausenstr. 21, D-81247 Miinchen, Tel.: +49 89-8107 147, Fax: +49 89-8107 300, E-mail: marion.kotrba@zsm.mwn.de Prof.Dr. Boris KRYSTUFEK, Slovenian Museum of Natural History, P.O.Box 290, SI-1001 Ljubljana, E-mail: boris.krystufek@uni-].si Prof.Dr. Sven O. KULLANDER, Swedish Museum of Natural History, Department of Vertebrate Zoology, P.O. Box 50007, SE-104 05 Stockholm, E-mail: sven.kullander@nrm.se Prof.Dr. Steven PERRY, Rheinische Friedrich-Wilhelms- Universitát, Institut fiir Zoologie, Poppelsdorfer Schloss, D-53115 Bonn, Tel. +49 228-73 3807, E-mail: perry@uni-bonn.de Dr. Wolfgang SCHAWALLER, Staatliches Museum für Naturkunde, Rosenstein 1, D-70191 Stuttgart, Germany, Tel. +49 711-8936 221, Fax +49 711-8936 100, E-mail: schawaller.smns@naturkundemuseum-bw.de Dr. W. David SıssoM, Dept. of Life, Earth and Environ- mental Sciences, W. Texas A. & M. University, WTAMU Box 60808, Canyon, Texas 79016, USA, E-mail: dsissom@wtamu.edu/ Dr. Miguel VENCES, University of Amsterdam, Zoological Museum, Mauritskade 61, PO Box 94766, NL-1090 GT Amsterdam, The Netherlands, Tel. +31 20-525 7319, E-mail: vences@science.uva.nl PD Dr. Heike WAGELE, Rheinische Friedrich-Wilhelms- Universität, Institut für Evolutionsbiologie und Okologie, D-53121 Bonn, Tel.: +49 228-73 5159, Fax: +49 234-322 4114, E-mail: hwaegele(@evolution.uni-bonn.de Dr. Erich WEBER, Eberhard-Karls-Universität, Zoologische Schausammlung, Sigwartstr. 3, D-72076 Tiibingen, Germany, E-mail: erich.weber@uni-tuebingen.de Bonner zoologische Beitráge | Band 53 (2004) Heft 1/2 Seiten 1-1 | Bonn, Juni 2005 | Clas M. Naumann (26.06.1939 — 15.02.2004) — in memoriam After several years of suffering from cancer, Professor Clas Michael Naumann zu Königsbrück died on Sun- day, 15" of February, 2004. He was a full professor of Systematic Zoology at the Rheinische Friedrich- Wilhelms-Universitát and director of the Zoologisches Forschungsinstitut und Museum Alexander Koenig, Bonn. Fig. 1. Professor Clas Michael Naumann zu Königsbrück, June 12, 1995 (Photo M. Schmitt) Clas Michael Naumann zu Königsbrück was born the 26th of June 1939, in Dresden as the only child of Dr. Eberhard Naumann zu Königsbrück and Freda-Irene, née Hannemann. He spent the first years of his life at the family estate (Kónigsbrúck, near Dresden) from where he could hear and see the bombing of Dresden (13th/14th of February 1945), an impression he pre- served well, as he recounted some fifty years later. He fled from the approaching Red Army with his famiiy in Spring 1945, another experience which left a lasting impression. His family found a provisional home in the village of Kirchohsen near Hameln, where Clas Naumann entered primary school. From 1949, he went to high school in Wilhelmshaven for two years, and finally in Braun- schweig, where his father, who had a doctoral degree in agriculture, found a position with the Bundesforschungs- anstalt fiir Landwirtschaft (Federal Research Institute of Agriculture). From his father, who was an ardent beetle collector, he was guided into natural history. Already during his high school years, he made contact with local entomologists, especially lepidopterists, e.g. Fritz Hart- / wieg. Certainly through these contacts, Clas Naumann’s passion for butterflies and moths developed in those years. He even wrote as his first publication a short re- port on the Lepidoptera found in the surroundings of Maria Taferl in Austria, where he had spent a summer holiday with his parents (in 1958, no. 1). In 1959, he passed the final secondary-school examinations. After three trimesters of “studium generale” at Leibniz Kol- leg, Túbingen, he studied chemistry from 1960 to 1962, and biology from 1962 to 1966 at the Eberhard-Karls- Universitat in Tiibingen. Here, his entomological inter- ests focussed on Burnet Moths, especially of the genus Zygaena. During his time in Tiibingen, he undertook his first en- tomological trip through a country of the Near East, and subsequently published his first species description: Zy- gaena problematica Naumann, 1966, from southern Turkey (no. 2). He also made first experiences as a businessman since he founded a trade of entomology supply. In his later days, he used to tell about this era of his life with great delight. In 1967, he switched to the Rheinische Friedrich Wil- helms-Universität in Bonn. Here, he received his doc- toral degree in 1970 after doing his doctoral dissertation at the Zoologisches Forschungsinstitut und Museum Alexander Koenig on the systematics and phylogeny of the holarctic Sesiidae — Insecta, Lepidoptera, under the supervision of Giinther Niethammer (published in 1971, no. 6). Subsequently, he took up an opportunity to work for three years as a university docent at Kabul Univer- sity (Afghanistan). Besides his teaching, he built up a zoological collection and a small university museum of zoology. The three years he spent in Afghanistan were most influential upon his further life and scientific ca- reer. He took more than just a fancy for the nature, cul- ture, history and people of that country. He learned the language(s) of this country and did extensive field work, even in quite remote parts of the country, such as Nuris- tan and Registan, the provinces of Badakhshan and Pak- tia, as well the Wakhan Valley in the Afghan Pamirs (Naumann 1974, nos. 11-14). His strong affection for Afghanistan and the whole region of the Middle East lasted for the rest of his life. He returned to Bonn in 1973 and received a two-year post as an assistant at the Institute for Applied Zoology (Professor Dr. Werner J. Kloft) in Bonn, followed by three years at the Ludwig-Maximilians-Universitát in München (Zoological Institute, Professor Dr. Jürgen Ja- cobs), where he received his habilitation in 1977. His Habilitationsschrift (professorial dissertation) was on the phylogeny and zoogeographical relations of the Zy- gaenini, published also in English in 1977 (no. 21). The same year he was appointed a professor at Bielefeld University, where he chaired the Department of Mor- phology and Systematics of Animals until 1989. He continued working on morphology, taxonomy, phylog- eny and zoogeography of the Zygaenidae, but extended his field of work into chemical ecology and sensory physiology. A quickly growing number of enthusiastic diploma and doctoral students soon formed a vivid and prolific team of investigators and authors. On July 1, 1989, he was appointed a full professor at the Rheinische Friedrich Wilhelms-Universitát in Bonn, and at the same time director of the Zoologisches For- schungsinstitut und Museum Alexander Koenig. Under his directorship, a corporate plan was elaborated that provided the institution with an internal structure, 1.e. departments (vertebrates, entomology, exhibition and public relations) and sections. He initiated monthly sci- entists’ meetings, weekly informal meetings of all staff members, and he made the regular meeting of the heads of the three departments an effectively working and well-accepted board of directors. His main goals were to increase the scientific output and to improve the interna- tional standing of Museum Koenig, and to enhance its attractiveness to the public. In all of these activities and throughout the whole of his decisions as Director, he strove for a consensus among the parties involved. And it caused literally anguish to him when he did not suc- ceed in reaching consensus. Clas Naumann clearly saw that the museum’s infra- structure had to be improved drastically in order to reach his goals. So, he worked inconceivably hard to acquire additional funds, be it for additional technical equipment such as a scanning electron microscope or a bone-degreasing device, or for a total renovation of the museum’s basement, or for the redesign of its public exhibition. Under his guidance, an entirely new concept for the exhibition was elaborated. With a considerable amount of “soft” money, the reorganisation of the exhibi- tion was achieved, at least partially, before his untimely death. However, most tragically, he was already too feeble to attend the opening ceremony for the first redesigned portion of the exhibition, on the 9" of October 2003. One of his most notable merits for Museum Koenig was that he succeeded in reaching an administrative decision from the state of Nordrhein-Westfalia and the German Federal Government for a new building. He would defi- nitely not have succeeded without the recruitment to ac- cept a position of full professor at the Miinchen Univer- sity and at the same time director of the Zoologische Staatssammlung Miinchen. When negotiating with both 2 Bonner zoologische Beitráge 53 (2004) the state ministries of Bavaria and Nordrhein- Westfalia, he managed to convince the government of the latter state to guarantee a new building, which would entice him to remain in Bonn. He was intensively involved in the detailed planning and invested a good deal of hope and energy in this enterprise. However, the construction of this new building began only the day after he died. As a director, Clas Naumann was always concerned about the welfare of the museum’s staff members, the countless colleagues who worked at the museum on grants, and of the students doing their theses at the mu- seum. And while he mostly did not show much of his emotions, he could be as merry as a lark when he man- aged to do something good to somebody, especially when it came as a surprise. Possibly, these were the most rewarding moments in his daily work. Clas Naumann was a dedicated academic teacher who also stimulated younger scientists to give university courses. In many cases, he announced these courses only formally, while in fact they were given by his doc- toral students. Thus, he not only provided these younger colleagues the opportunity of gaining experience in teaching, but also considerably enhanced the range of courses for students of zoology at the Bonn University. In addition, he supervised numerous diploma- and doc- toral theses (see list below), both at Bielefeld and Bonn. Systematists in Central Europe, zoologists and espe- cially museum entomologists, owe Clas Naumann a great deal for his never ending efforts for a better lobby for our field of science within Germany’s political and general scientific frame. He was one of the promoters of the Gesellschaft fiir biologische Systematik (Society for Biological Systematics) founded December 12, 1997, and organised its first annual meeting at Museum Koenig in 1998. He strove for more co-operation be- tween the German museums with vast holdings of scien- tific collections, was one of the founders of the DNFS (Direktorenkonferenz Naturwissenschaftlicher For- schungssammlungen in Deutschland), and he served for several years as a referee for the Deutsche Forschungs- gemeinschaft (German Research Foundation). Through his tireless lobbying, his own scientific papers and through the numerous diploma and doctoral theses he stimulated and supervised, Clas Naumann contributed immensely to the revitalisation and the expansion of taxonomy and systematics in Germany. Among Clas Naumann’s super-regional scientific activi- ties, his efforts for the journal Zoologischer Anzeiger played an outstanding part. He was editor-in-chief since 1995. Without his energy and optimism, this journal would certainly have ended in 1994, when Dr. Johanna Schliiter from Gustav Fischer Verlag, Jena (the publish- ers at that time) asked Professor Naumann to accept the task of saving the oldest German zoological jour- Clas M. NAUMANN: in memorian 3 nal. The present profile of the journal and the boards of editors and advisors still bear Professor Naumann’s mark. Also, Clas Naumann was very much interested in im- proving the museum’s own publication series. It was his explicit desire that Bonner zoologische Beiträge should grow into an internationally renowned, fully peer re- viewed journal of high standards in appearance and content. We, the team of editors and advisors since vol- ume 51, are proud to say that Clas Naumann was highly satisfied when he saw the newly designed issues of vol. 51. Clas Naumann worked tremendously hard for the good of Museum Koenig, for all the persons working here, for a better standing of the science region of Bonn in a wider context, for taxonomy and systematics, and for science in general. He did so finally at the cost of his private life and even his health. Who ever knew him well enough to be able to judge, and close enough to care, felt relieved when he informed us about his decision to retire ahead of schedule in April 2004, and he and we hoped he had some time to devote to his family and his hobbies. But regrettably, he passed away some two months before that date. Thus, he could neither live to see the new building being completed nor could he introduce his successor smoothly into his duties, as he wished. Two fields of interest absorbed Clas Naumann espe- cially besides his scientific and administrative work: Burnet Moths and Afghanistan (and adjacent countries). In his spare time and on several expeditions, he col- lected and investigated Zygaenidae. It was one of his highest satisfactions to discover a new species or to re- discover a lost one. He published numerous taxonomic, morphological and zoogeographical studies on that group of moths, and he enjoyed meeting other Zygaeni- dae enthusiasts (Fig. 2). Until the last weeks of his life, he was very much con- cerned about the political, social and scientific devel- opments in Afghanistan, especially at the University of Kabul. While many of his public lectures were either characterised by a certain scientific distance or by tacti- cal considerations (e.g., in order to convince his audi- ence of a certain goal), he gave an entirely different im- pression when giving a talk on countryside, people, culture and history of Afghanistan. His voice, his smile, his gesture changed. No doubt was left that this was his true passion. He was significantly involved in the revitalisation of the partnership between the Rhei- nische Friedrich-Wilhelms-Universitát Bonn and the Kabul University, and he invested not only just some thoughts in writing several proposals, but he travelled to Kabul and after returning organised campaigns for as- sistance. Fig. 2. Clas Naumann on the Isle of Skye (Scotland), during an International Zygaenidae Symposium, September 11, 1996. Left: Gerhard Tarmann (Innsbruck), right: Jean-Marie Desse (Angers) (Photo C. Esch, Miinster). Scientists, technicians, other staff members and students of the Zoologisches Forschungsmuseum Alexander Koenig are deeply indebted to Clas Naumann for his tireless efforts. The board of directors and scientific staff unanimously decided to dedicate the new building to the memory of their former director and to name it “Clas M. Naumann-Bau”. Clas Michael Naumann zu Königsbrück is survived by his wife Storai Naumann-Nawabi and his children Rox- ana and Alexander, to whom we express our deep sym- pathy. Acknowledgements. Storai Naumann-Nawabi (Wacht- berg-Pech) read the manuscript of this obituary and relin- quished a list of her husband’s publications to me, Renate Feist (Bielefeld) compiled a list of Clas Naumann’s di- ploma and doctoral students in Bielefeld, Christoph Hauser (Stuttgart) provided me with a draft of an extensive necrol- ogy meant for a lepidopterists’ journal, Christoph Esch (Miinster) gave permission to use the photograph of Figure 2, Axel Hofmann (Breisach) identified the persons showed on this photo, and Brad Sinclair (Bonn) improved the Eng- lish of my manuscript. Bonner zoologische Beiträge 53 (2004) PUBLICATIONS OF CLAS M. NAUMANN (AS OF 01.02.2005, COMPILED BY MICHAEL SCHMITT) 1 un 10 11 12 13 14 NAUMANN, C. (1958): Schmetterlinge aus der Umge- bung von Maria-Taferl (Niederósterreich). Zeitschrift der Wiener entomologischen Gesellschaft 43: 33-36. NAUMANN, C. (1966): Zygaena (Subg. Zygaena F.) problematica nov. sp. Zeitschrift der Wiener ento- mologischen Gesellschaft 51: 10-19. NAUMANN, C. (1969): Zur Kenntnis der Mesem- brynus-Arten Afghanistans (Lep., Zygaenidae). Bon- ner zoologische Beiträge 20: 266-278. NAUMANN, C. (1970): Bemerkungen zur Biologie und zur systematischen Stellung von Zygaena graslini Lederer, 1855 (Lep., Zygaenidae). Bonner zoologische Beiträge 21: 133-136. NAUMANN, C. (1971): 6. Schmetterlinge. Pp. 547-549 in: NIETHAMMER, G.: Die Fauna der Sahara und ihrer Randgebiete, Darstellung eines Naturgroßraumes, Bd. 1: Physiogeographie. Miinchen (Weltforum-Verlag). NAUMANN, C. (1971): Untersuchungen zur Systematik und Phylogenese der holarktischen Sesiiden (Insecta, Lepidoptera). Bonner zoologische Monographien 1: I— 133. NAUMANN, C. M. & NOGGE, G. (1973): Die Zoolo- gischen Projekte Afghanistans — erfolgreiches Neben- produkt einer Universitatspartnerschaft in Afghanistan. GAWI-Rundbrief 4/73: 25-28. NAUMANN, C. & NOGGE, G. (1973): Die Großsäuger Afghanistans. Zeitschrift des Kólner Zoo 16: 79-93. NAUMANN, C. & NIETHAMMER, J. (1973): Zur Sáugetierfauna des afghanischen Pamir und des Wak- han. Bonner zoologische Beiträge 24: 237-248. NAUMANN, C. M. (1973): Ein ehemaliges Wildyak- Vorkommen im afghanischen Pamir. Bonner zoolo- gische Beitráge 24: 249-253. NAUMANN, C. M. (1974): Beobachtungen úber den Verlauf einer Lepidopteren-Immigration in Afghani- stan 1972. Atalanta 5: 82-88. NAUMANN, C. M. (1974): Pamir und Wakhan — Kurz- bericht zweier Expeditionen (1971 und 1972) nebst einigen allgemeinen Bermerkungen. Afghanistan Journal 1: 91-104. NAUMANN, C. M. (1974): Neue Zygaena-Unterarten aus Afghanistan. Entomologische Zeitschrift Frankfurt am Main 84: 29-36. NAUMANN, C. & NIETHAMMER, J. (1974): Neu- nachweise von Sáugetieren aus dem nórdlichen Af- ghanistan. Säugetierkundliche Mitteilungen 22: 295— 298. NAUMANN, C. M. € NAUROZ, M.K. (1975): Be- merkungen zur Verbreitung des Markhors, Capra fal- coneri (Wagner, 1839) in Afghanistan. Sáugetierkund- liche Mitteilungen 23: 81-85. NAUMANN, C. M. & SCHULTE, A. (1977): Eine weitere neue Subspecies von Zygaena (Agrumenia) shivacola Reiss & Schulte, 1962 (Lepidoptera, Zygaenidae). En- tomologische Zeitschrift Frankfurt am Main 87: 89— 93. NAUMANN, C. M. (1977): Zygaena (Mesembrynus) halima n.sp. und einige Bemerkungen zur stammes- geschichtlichen Gliederung der Gattung .Zygaena F. (Lepidoptera: Zygaenidae). Zeitschrift der Arbeitsge- meinschaft ósterreichischer Entomologen. 29: 35-40. e 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 NAUMANN, C. M. (1977): Biologie, Verbreitung und Morphologie von Praezygaena (Epizygaenella) caschmirensis (Kollar, 1848) (Lepidoptera, Zygaeni- dae). Spixiana 1: 45-84. NAUMANN, C. M. (1977): Stammesgeschichte und tiergeographische Beziehungen der Zygaenini (Insecta, Lepidoptera, Zygaenidae). Mitteilungen der Miinchner entomologischen Gesellschaft 67: 1—25. NAUMANN, C. M. (1977): Rasterelektronenoptische Untersuchungen zur Feinstruktur von Lepidopteren- Gespinsten. Mitteilungen der Miinchner entomolo- gischen Gesellschaft 67: 27-37. NAUMANN, C. M. (1977): Studies on the Systematics and Phylogeny of Holarctic Sesiidae (Insecta, Lepi- doptera) [Untersuchungen zur Systematik und Phy- logenese der holarktischen Sesiiden (Insecta, Lepidop- tera)]. 139 pp., Amerind Publishing Co. Pvt. Ltd., New Delhi, Washington. NAUMANN, C. M., JUNGE, G. & ROSE, K. (1977): Ein weiterer Beitrag zur Zygaenen-Fauna Kappadokiens (Lep. Zygaenidae). Atalanta 8: 282-289. NAUMANN, C. M., & NAUMANN, S. (1978): Eine neue ostanatolische Unterart der Zygaena (Zygaena) loti ([Denis und Schiffermiiller], 1775) (Lep., Zygaeni- dae). Entomologische Zeitschrift Frankfurt am Main 88: 37-40. NAUMANN, C. M., Huss, H. & NAUROZ, M. K. (1978): Zur Tierwelt des "Großen Pamir". Pp. 202-213 in: DE GRANCY, R. S. & KOSTKA, R. (eds): Grosser Pamir. Verlagsanstalt, Graz. NAUMANN, C. M. & RACHELI, T. (1978): Taxonomic and ecological observations on Zygaena F. from the Province of Fars, Southern Iran (Lepidoptera, Zygae- nıdae). Atalanta 9: 208-218. NAUMANN, C.M. (1978): Zur Systematik, Verbreitung und Biologie von Zygaena pamira Sheljuzhko, 1919 (Lepidoptera, Zygaenidae). Atalanta 9: 355-372. NAUMANN, C. M. (1979. Hans Burgeffs Beitrag zur Zygaenenkunde. Atalanta 10: 85-88. RACHELI, T. & NAUMANN, C. M. (1979): On Zygaena Fabricius (Zygaenidae) from Fars, south Iran. Nota lepidopterologica 2: 53-55. NAUMANN, C. M. & BENDER, R. (1980): Eine neue Arctia intercalaris-Subspezies aus dem Hindukusch (Lepidoptera, Arctiidae). Atalanta 11: 21-28. NAUMANN, C. M. & EISNER, C. (1980): Parnassiana Nova LVII. Beitrag zur Ökologie und Taxonomie der afghanischen Parnassiidae (Lepidoptera). Zoologische Verhandelingen Leiden 178: 35 pp., 9 pls. NAUMANN, C. M. & TREMEWAN, W. G. (1980): On the biology of Zygaena (Mesembrynus) tamara Chris- toph, 1889 (Lepidoptera: Zygaenidae). Entomologist's Gazette 31: 113-121. NAUMANN, C. M. & NAUMANN, S. (1980): Neue Zy- gaenen aus den Aufsammlungen W. Eckweiler's aus dem Iran (Lepidoptera Zygaenidae). Mitteilungen der Basler entomologischen Gesellschaft N. F. 30: 47-54. NAUMANN, C. M. & NAUMANN, S. (1980): Ein Beitrag zur Kenntnis der Zygaenen-Fauna Nord- und Ost- Anatoliens (Lep., Zygaenidae). Entomofauna 1: 302— 353. NAUMANN, C. M. & SCHROEDER, D. (1980): Ein weiteres Zwillingsarten-Paar mitteleuropáischer Se- siiden: Chamaesphecia tenthrediniformis ([Denis & Schiffermúller], 1775) und Chamaesphecia empi- 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 formis Clas M. NAUMANN: in memorian (Esper, 1783) (Lepidoptera, Sesiidae). Zeitschrift der Arbeitsgemeinschaft Österreichischer Entomologen 32: 29-46. NAUMANN, C. M. & ROSE, K. (1981): Eine bisher un- bekannte peucedanoide Population der Zygaena ephialtes (Linnaeus, 1763) aus Griechenland (Lepi- doptera, Zygaenidae). Entomofauna 2: 113-124. NAUMANN, C. M. (1981): Die zoologischen Forschun- gen der letzten zwei Jahrzehnte. Pp. 105-114 in: RATHJENS, C. (ed.) Neue Forschungen in Afghanistan. Leske-Verlag + Budrich GmbH, Opladen. NAUMANN, C. M. (1982): Zur Kenntnis der von Man- fred Koch beschriebenen Taxa des Zygaena- purpuralis-Komplexes (Lepidoptera, Zygaenidae). En- tomofauna 3: 225-242, ’ NAUMANN, C. M. (1982): Zur Kenntnis der von Otto Holik beschriebenen Taxa des Zygaena-purpuralis- Komplexes. Entomofauna 3: 371—405. NAUMANN, C. M. (1982): Zur Kenntnis der von Hans Burgeff beschriebenen Taxa des Zygaena-purpuralis- Komplexes (Lepidoptera, Zygaenidae). Entomofauna 3: 411-437. NAUMANN, C. M. & TARMANN, G. (1983): Ergebnisse der tschechoslowakisch-iranischen entomologischen Expeditionen nach dem Iran 1973 und 1977 (Lepidop- tera, Zygaenidae). Acta entomologica Musei Nationa- lis Pragae 41: 273-276. NAUMANN, C. M. (1982): Zur Kenntnis der von Hugo Reiss beschriebenen Taxa des Zygaena-purpuralis- Komplexes (Lepidoptera, Zygaenidae). Entomofauna 3: 459-498. NAUMANN, C. M. (1983): Zur Kenntnis der von A. Costantini und Ubaldo Rocci beschriebenen Taxa des Zygaena purpuralis- Komplexes (Lepidoptera, Zygae- nidae). Entomofauna 4: 97-107. NAUMANN, C. M. (1983): Zur Kenntnis der von Rug- gero Verity beschriebenen Taxa des Zygaena- purpuralis-Komplexes (Lepidoptera, Zygaenidae). En- tomofauna 4: 141-155. NAUMANN, C. M. (1983): Zur Kenntnis der von A. Przegendza beschriebenen Taxa des Zvgaena- purpuralis-Komplexes (Lepidoptera, Zygaenidae). En- tomofauna 4: 173-180. NAUMANN, C. M., RICHTER, G. & WEBER, U. (1983): Spezifität und Variabilität im Zygaena-purpuralis- Komplex (Lepidoptera, Zygaenidae). Theses Zoologi- cae 2: 1-263 (Verlag J. Cramer, Braunschweig). NAUMANN, C. M. (1983): Zur Kenntnis der von O. Staudinger und H. Rebel beschriebenen Taxa des Zy- gaena-purpuralis-Komplexes (Lepidoptera, Zygaeni- dae). Entomofauna 4: 201-208. NAUMANN, C. M. (1984): Zur Kenntnis der von Leo Sheljuzhko und Otto Holik & Leo Sheljuzhko beschriebenen Taxa des Zygaena-purpuralis- Komplexes (Lepidoptera, Zygaenidae). Entomofauna 5:1-19. NAUMANN, C. M. & TREMEWAN, W. G. (1984): Zy- gaena anthyllidis Boisduval, [1828] (Insecta, Lepidop- tera): Proposed Conservation, Z. N. (S.) 2442. Bulletin of zoological Nomenclature 41: 73-76. NAUMANN, C. M. & TREMEWAN, W. G. (1984): Das Biospecies-Konzept in seiner Anwendung auf die Gat- tung Zygaena Fabricius, 1775. Spixiana 7: 161-193. NAUMANN, C. M., FEIST, R., RICHTER, G. & WEBER, U. (1984): Verbreitungsatlas der Gattung Zvgaena 51 52 53 54 33 56 57 58 59 60 61 62 63 64 65 N Fabricius, 1775 (Lepidoptera, Zygaenidae). Theses Zoologicae 5: 1-97 (Verlag J. Cramer, Braunschweig). FRANZL, S. & NAUMANN, C. M. (1984): Morphologie und Histologie der Wehrsekretbehälter erwachsener Raupen von Zygaena trifolii (Lepidoptera, Zygaeni- dae). Entomologische Abhandlungen und Berichte aus dem Staatlichen Museum für Tierkunde Dresden 118: 1-12. WITTHOHN, K. & NAUMANN, C. M. (1984): Qualita- tive and quantitative studies on the compounds of the larval defensive secretion of Zygaena trifolii (Esper, 1783) (Insecta, Lepidoptera, Zygaenidae). Compara- tive Biochemistry and Physiology 79C: 103-106. WITTHOHN, K. & NAUMANN, C. M. (1984): The dis- tribution of B-cyano-L-alanine in cyanogenic Lepidop- tera. Zeitschrift fiir Naturforschung 39C: 837-840. NAUMANN, C. M. (1984): Zur Kenntnis der von Fran- cis Dujardin beschriebenen Taxa des Zygaena- purpuralis-Komplexes (Lepidoptera, Zygaenidae). En- tomofauna 5: 343-354. PRIESNER, E., NAUMANN, C. M. & STERTENBRINK, J. (1984): Specificity of synthetic sex-attractants in Zy- gaena moths. Zeitschrift für Naturforschung 39C: 841-844. HAUSER, C. L., NAUMANN, C. M. & TREMEWAN, W. G. (1985): On the biology of Parnassius charltonius Gray, 1852 (Lepidoptera: Papilionidae). Entomolo- gist's Gazette 36: 5-13. NAUMANN, C. M. & EDELMANN, A. (1984): Insects of Southern Arabia - The life history, ecology and distri- bution of Reissita simonyi (Rebel, 1899) (Zygaenidae, Lepidoptera). Fauna of Saudi Arabia 6: 473-509. FRANZL, S. & NAUMANN, C. M. (1985): Cuticular cavities: Storage chambers for cyanoglucoside- containing defensive secretions in larvae of a zygaenid moth. Tissue & Cell 17: 267-278. NAUMANN, C. M. (1985): Zur Okologie und Biologie von Zygaena (Zygaena) christa Reiss & Schulte, 1967. Nota lepidopterologica 8: 42-50. NAUMANN, C. M. & NAUMANN, S. (1985): Zur mor- phologischen Differenzierung asiatischer Populationen des Zygaena-purpuralis-Komplexes (Lepidoptera, Zy- gaenidae). Entomofauna 6: 265-358. NAUMANN, C. M. (1985): Phylogenetische Systematik und klassisch-typologische Systematik - mit einigen Anmerkungen zu stammesgeschichtlichen Fragen bel den Zygaenidae (Lepidoptera). Mitteilungen der Miinchner entomologischen Gesellschaft 74: 1-35. FRANZL, S., NAHRSTEDT, A. & NAUMANN, C. M. (1986): Evidence for site of biosynthesis and transport of the cyanoglucosides linamarin and lotaustralin in larvae of Zygaena trifolii Insecta, Lepidoptera). Jour- nal of Insect Physiology 32: 705—709. NAUMANN, C. M. & WITTHOHN, K. (1986): Cyano- genese bei Zygaeniden (Insecta, Lepidoptera) und ihren larvalen Nahrungspflanzen: Co-Evolution oder einseitige Strategie-Optimierung? Verhandlungen der deutschen zoologischen Gesellschaft 79: 181-182. BODE, W. & NAUMANN, C. M. (1987): Structure of a newly discovered glandular organ in Neurosymploca larvae (Lepidoptera, Zygaenidae). Zoologische Jahr- biicher Anatomie 115: 319-329. NAUMANN, C. M. (1987): Distribution patterns of Zy- gaena moths in the Near and Middle East (Insecta, Lepidoptera, Zygaenidae). Beihefte zum Túbinger At- 6 66 67 68 69 70 71 72 73 74 76 77 78 79 Bonner zoologische Beitráge 53 (2004) las des Vorderen Orients, Reihe A (Naturwissen- schaften) 28 (KRUPP, F., SCHNEIDER, W. & KINZEL- BACH, R., eds., Proceedings of the Symposium on the Fauna and Zoogeography of the Middle East): 200- 212. NAUMANN, C. M. & FEIST, R. (1987): The structure and distribution of cyanoglucoside-storing cuticular cavities in Pryeria sinica Moore (Lepidoptera, Zygae- nidae). Zoologica Scripta 16: 89-93. WITTHOHN, K. & NAUMANN, C. M. (1987): Cyano- genesis — a general phenomenon in the Lepidoptera? Journal of chemical Ecology 13: 1789-1809. NAUMANN, C. M. & POVOLNY, D. (1987): Zur Lage- beziehung und Funktion von Wehrsekretbeháltern und Sinneshaaren im larvalen Integument von Zygaena (Insecta, Lepidoptera, Zygaenidae). Entomologische Abhandlungen und Berichte aus dem Staatlichen Mu- seum für Tierkunde Dresden 50: 193-198. BODE, W. & NAUMANN, C. M. (1987. On a pair of lit- tle-known accessory glands in female Zygaena moths (Lepidoptera, Zygaenidae). Zoological Journal of the Linnean Society 92: 27-42, NAUMANN, C. M. (1987): On the phylogenetic signi- ficance of two Miocene zygaenid moths Insecta, Lepidoptera). Palaeontologische Zeitschrift 61: 299— 308. WITTHOHN, K. & NAUMANN, C. M. (1987): Active cyanogenesis — a new phenomenon in arthropods. Zeitschrif ftir Naturforschung 42C: 1319-1322. NAUMANN, C. M. (1988): The internal female genita- lia of some Zygaenidae (Insecta, Lepidoptera) their morphology and remarks on their phylogenetic signifi- cance. Systematic Entomology 18: 85-99. NAUMANN, C. M. (1988): Functional morphology of the external male and female genitalia in Zygaena Fab- ricius, 1775 (Lepidoptera: Zygaenidae). Entomologica Scandinavica 18: 213-219. NAUMANN, C. M. (1987): Epizygaenella erythrosoma (Hampson, [1893]), with notes on the taxonomic treatment of the genus Epizygaenella Tremewan & Povolny, 1968 (Lepidoptera, Zygaenidae). Mitteilun- gen der Miinchner entomologischen Gesellschaft 77: 139-147. FRANZL, S., NAHRSTEDT, A. & NAUMANN, C. M. (1987): Chemische Abwehr bei Zygaena-Larven (Lepidoptera, Zygaenidae). Einfluß der Häutungsperi- odik auf Morphologie und Inhaltsstoffe kutikularer Wehrsekretbehälter. Verhandlungen der deutschen zo- ologischen Gesellschaft 80: 208-209. NAUMANN, C. M. (1988): Revision der Esper'schen Zygaena-Typen (Insecta, Lepidoptera, Zygaenidae). Mitteilungen des internationalen entomologischen Vereins 13: 1-22. NAUMANN, C. M. (1988): Zur Evolution und adap- tiven Bedeutung zweier unterschiedlicher Partner- findungsstrategien bei Zygaena trifolii (Esper, 1783) (Insecta, Lepidoptera). Verhandlungen der deutschen zoologischen Gesellschaft 81: 257-258. FÄNGER, H. & NAUMANN, C. M. (1988): Aufbau des männlichen Genitaltraktes und Spermatophoren- bildung bei Zygaena trifolii (Esper, 1783) (Insecta, Lepidoptera). Verhandlungen der deutschen zoolo- gischen Gesellschaft 81: 249-250. PRINZ, J. & NAUMANN, C. M. (1988): Optische Pa- rameter bei der Partnerfindung von Zygaena trifolii 80 81 82 83 84 85 86 87 88 89 90 91 92 filipendulae (Esper, 1783) (Insecta, Lepidoptera). Verhandlungen der deutschen zoologischen Gesellschaft 81: 258. FRANZL, S., NAUMANN, C. M. & NAHRSTEDT, A. (1988): Cyanoglucoside storing cuticle of Zygaena larvae (Insecta, Lepidoptera). Morphological and cyanoglucoside changes during the moult. Zoomor- phology 108: 183-190. TREMEWAN, W. G. & NAUMANN, C. M. (1989): The nominal taxa described by G. Grum-Grshimailo in the genus Zygaena Fabricius, 1775 (Lepidoptera: Zygae- nidae). Entomologist’s Gazette 40: 109-113, pl. 9, 10. TREMEWAN, W. G. & NAUMANN, C. M. (1989): A re- vision of Zygaena (Agrumenia) magiana Staudinger, 1889 (Lepidoptera: Zygaenidae). Entomologist’s Ga- zette 40: 114-122, pls 11, 12. NAUMANN, C. M. (1990): Stammesgeschichtliche Re- konstruktion und ökologische Beziehungen der Organ- ismen — Ansátze ftir interdisziplináre Kooperation. Verhandlungen der deutschen zoologischen Gesell- schaft 83: 291-301. OCKENFELS, P., NAUMANN, C. M., SCHMITZ, J., SCHMIDT, W. & FRANCKE, W. (1990): Antennale Reaktionen von Zygaena trifolii (Zygaenidae, Lepi- doptera) auf Blüteninhaltsstoffe von Knautia arvensis. Verhandlungen der deutschen zoologischen Gesell- schaft 83: 424. ESCH, C. B. & NAUMANN, C. M. (1990): Morpholo- gische und ethologische Differenzierung bei Zygaena (Linnaeus, 1758) und Z. lonicerae (Scheven, 1777) (Lepidoptera, Zygaenidae). Verhand- lungen der deutschen zoologischen Gesellschaft 83: 505. SCHADE, R. & NAUMANN, C. M. (1990): Morphologie und phylogenetische Bedeutung der Sieboldschen Driise einiger Nymphalidae (Lepidoptera). Verhand- lungen der deutschen zoologischen Gesellschaft 83: 513. HILLE, A., MULLER-TAPPE, S. & NAUMANN, C. M. (1990): Parameter genetischer Variabilität bei isolierten Zygaena-Populationen am nord-westlichen Arealrand (Insecta, Lepidoptera). Verhandlungen der deutschen zoologischen Gesellschaft 83: 609-610. NAUMANN, C. M., OCKENFELS, P., FRANCKE, W., SCHMIDT, F. & SCHMITZ, J. (1991): Reactions of Zy- gaena moths (Lepidoptera: Zygaenidae) to volatile compounds of Knautia arvensis. Entomologia gener- alıs 15: 255-264. NAUMANN, C. M. & TREMEWAN, W. G. (1991): A new subspecies of Zygaena (Zygaena) rhadamanthus (Esper, [1789]) from southern Portugal (Lepidoptera: Zygaenidae). Entomologist’s Gazette 42: 85-88. NAUMANN, C. M. & TREMEWAN, W. G. (1991): The type-material and type-locality of Zygaena (Agrume- nia) fausta faustina Ochsenheimer, 1808 (Lepidoptera: Zygaenidae). Entomologist's Gazette 42: 157-160. HILLE, A. & NAUMANN, C. M. (1992): Allozyme dif- ferentiation in Zygaena transalpina Esper, 1780) (Lepidoptera: Zygaenidae). Pp. 59-87 in: DUTREIX, C. et al. (eds) Recent advances in burnet moth research (Lepidoptera, Zygaenidae). Proceedings of the 4” Symposium on Zygaenidae, Nantes 11-13 September 1987. Költz, Königstein. WIPKING, W. & NAUMANN, C. M. (1992): Diapause and related phenomena in zygaenid moths. Pp. 107— 128 in: DUTREIX, C. et al. (eds) Recent advances in Clas M. NAUMANN: in memorian burnet moth research (Lepidoptera, Zygaenidae). Pro- ceedings of the 4" Symposium on Zygaenidae, Nantes 11-13 September 1987. Kóltz, KGnigstein. 93 LAMBERT, B. & NAUMANN. C. M. (1992): Zygaena (Zygaena) nevadensis Rambur, 1844 - new for Turkey (Zygaenidae, Lepidoptera). Pp. 184-186 in: DUTREIX, C. et al. (eds) Recent advances in burnet moth research (Lepidoptera, Zygaenidae). Proceedings of the 4" Symposium on Zygaenidae, Nantes 11-13 September 1987. Költz, Königstein. 94 HILLE, A., OCKENFELS, P., BÖHMER, T. & NAUMANN, C.M. (1993): Untersuchungen zur Pheromonspezifität zweier parapatrischer Zygaena Species. Verhandlun- gen der deutschen zoologischen Gesellschaft 86: 244. 95 OCKENFELS, P., NAUMANN, C. M. € FRANCKE, W. (1993): Die Rolle von Bliiteninhaltsstoffen und ihren Derivaten im Paarungsverhalten von Zygaena trifolii (Lepidoptera: Zygaenidae). Verhandlungen der deut- schen zoologischen Gesellschaft 86: 266. 96 SOBOTTA, J., OCKENFELS, P. & NAUMANN, C. M. (1993): Die Nutzung von Pyrolizidin-Alkaloiden durch Oreina elongata (Coleoptera: Chrysomelidae). Ver- handlungen der deutschen zoologischen Gesellschaft 86: 273. 97 NAUMANN, C. M. (1993): Die Russische Entomolo- gische Gesellschaft. Mitteilungen der Arbeitsgemein- schaft Rheinischer Koleopterologen (Bonn) 3: 63-65. 98 FÄNGER, H. & NAUMANN, C. M. (1993): Correlation between the mesodermal male genital system and the structure of the spermatophore of Zygaena trifolii (Es- per, 1783) (Insecta, Lepidoptera, Zygaenidae). Acta Zoologica 74: 239-246. 99 OCKENFELS, P., PRINZ, J. & NAUMANN, C. M. (1993): Reactions of Aleioides cf. assimilis (Hymenoptera: Braconidae) to the larval foodplant of its host Zygaena trifolii (Esper, 1783) (Lepidoptera: Zygaenidae). En- tomologist’s Gazette 44: 125—133. 100 NAUMANN, C. M., SCHURIAN, K. & WEISS, J.-C. (1993): Notes on the ecology and early stages of Zy- gaena (Mesembrynus) lydia Staudinger, 1887 (Lepi- doptera: Zygaenidae). Entomologist’s Gazette 44: 225-229, 1 pl. 101 HAUSER, C. L., NAUMANN, C. M. & KREUZBERG, A. V.-A. (1993): Zur taxonomischen und phyloge- netischen Bedeutung der Feinstruktur der Eischale der Parnassiinae (Lepidoptera, Papilionidae). Zoologische Mededeelingen Leiden 67: 239-264. 102 HEINE, S. & NAUMANN, C. M. (1994): Sexuelle Selek- tion und die Evolution tageszeitlich variierender Part- nerfindungsmechanismen bei Zygaena trifolii (Esper, 1783) (Insecta, Lepidoptera). Mitteilungen der deutschen Gesellschaft fiir allgemeine und angewandte Entomologie 8: 649— 658. 103 NAUMANN, C. M. & TREMEWAN, W. G. (1994): The holotype of Zygaena glasunovi Grum-Grshimailo, 1893 (Lepidoptera: Zygaenidae). Entomologist's Ga- zette 45: 123-126. 104 GASSMANN, D., HILLE, A. & NAUMANN, C. M. (1994): Morphometrische Variation in der Kontaktzone zweier Semispezies des Zygaena-transalpina-Komplexes (Lepidoptera. Zygaenidae). Verhandlungen der deut- schen zoologischen Gesellschaft 87.1: 223. 105 HILLE, A. & NAUMANN, C. M. (1994): Allel- Stufenkline und Multilocus-Kopplungsgleichgewichte von Allzymen kennzeichnen eine parapatrische Kon- taktzone zwischen Zygaena angelicae und Z. hip pocrepidis (Insecta, Lepidopt., Zygaenidae) in Süd- deutschland. Verhandlungen der deutschen zoologi- schen Gesellschaft 87.1: 225. 106 SPEIDEL, W. & NAUMANN, C. M. (1995): Phylogenetic aspects in the higher classification of the subfamily Catocalinae (Lepidoptera, Noctuidae). Beiträge zur Entomologie 45: 109-118. 107 SPEIDEL, W. & NAUMANN, C. M. (1995): Further morphological characters for a phylogenetic classifica- tion of the Noctuidae (Lepidoptera). Beitráge zur En- tomologie 45: 119-135. 108 WEYER, M. & NAUMANN, C. M. (1996): Möglich- keiten zur Integration ökologischer Ursache-Wir- kungsbeziehungen in ein naturwissenschaftliches Mu- seum zur Verbesserung von Umweltaufklárung und -erziehung. Umwelt-Bundesamt (Berlin): Texte 4/96: 423 pp. 109 SPEIDEL, W., FANGER, H. & NAUMANN, C. M. (1996): The ultrastructure of the noctuid proboscis (Lepidop- tera: Noctuidae). Zoologischer Anzeiger 234: 307 315. 110 SPEIDEL, W., FANGER, H. & NAUMANN, C. M. (1996): The phylogeny of the Noctuidae (Lepidoptera). Sys- tematic Entomology 21: 219-251. 111 NAUMANN, C.M. & TREMEWAN, W. G. T. (1996): Geographical variation in the last instar larvae of Zy- gaena (Mesembrynus) corsica Boisduval, [1828] (Lepidoptera, Zygaenidae). Linnaea belgica 15: 286- 288. 112 SPEIDEL, W., FÄNGER, H. & NAUMANN, C.M. (1997): On the systematic position of Cocytia (Lepidoptera, Noctuidae). Deutsche entomologische Zeitschrift N. F. 44: 27-31. 113 FANGER, H. & NAUMANN, C. M. (1998): Genital mor- phology and copulatory mechanism in Zygaena trifolii (Esper, 1783) (Insecta, Lepidoptera, Zygaenidae). Acta zoologica 79: 9-24, 114 NAUMANN, C. M. (1998): Biodiversity - Is there a sec- ond chance? Pp. 3-11 in: BARTHLOTT, W. & WINIGER, M. (eds): Biodiversity. A Challenge for Developmen- tal Research and Policy. Springer, Berlin, Heidelberg etc. 115 TREMEWAN, W. G. & NAUMANN, C. M. (1998): Notes on the biology and ecology of Zygaena (Mesem- brynus) alpherakyi Sheljuzhko 1936 (Lepidoptera: Zygaenidae). Stapfia 55: 107-112. 116 NAUMANN, C. M. (1998): Biogeographie und Stammesgeschichte. Prioritáten fúr die Erhaltung von Biodiversität. Pp. 46-49 in: BARTHLOTT, W. & GUT- MANN, M. (eds) Biodiversitätsforschung in Deutsch- land. Potentiale und Perspektiven. Europäische Akademie zur Erforschung von Folgen wissen- schaftlich-technischer Entwicklungen, Graue Reihe no 11. 117 HILLE, A. & NAUMANN, C. M. (1998): Allozyme dif- ferentiation in the Zygaena trasnalpina (Esper, 1780) superspecies complex (Lepidoptera, Zygaenidae). II. Genetic diversity and population taxonomy of the two semispecies Zygaena angelicae (Ochsenheimer, 1808) and Z. hippocrepidis (Húbner, [1799]). Theses zo- ologicae 30: 45-88. 118 GASSMANN, D., HILLE, A. & NAUMANN, C. M. (1998): Morphological and morphometrical investigations within a geographical contact zone between Zveaena 8 Bonner zoologische Beiträge 53 (2004) angelicae Ochsenheimer, 1808 and Zygaena transal- pina hippocrepidis (Hübner, [1799]) (Lepidoptera: Zygaenidae) in southern Germany. Theses zoologicae 30: 89-104. 119 BÖHMER, T., HILLE, A., NAUMANN, C. M. & OCK- ENFELS, P. (1998): On the specificity of female sex pheromone compounds in the Zygaena-transalpina- complex (Zygaenidae, Lepidoptera, Insecta). Theses zoologicae 30: 105-124. 120 MULLER, T., OCKENFELS, P. & NAUMANN, C. M. (1998): Olfactory discrimination between host and non-host-plants in Zygaena trifolii (Esper, 1783) and Z. transalpina (Esper, 1782) (Lepidoptera, Zygaeni- dae). Theses zoologicae 30: 125-138. 121 EscH, C. B. & NAUMANN, C. M. (1998): Host- searching and host-discrimination of a pupal parasitoid of Zygaena trifolii (Esper, 1783) (Lepidoptera, Zygae- nidae). Theses zoologicae 30: 139-158. 122 FANGER, H. & NAUMANN, C. M. (1998): Delayed in- semination in a ditrysian moth, Zygaena trifolii (Esper, 1783) (Insecta, Lepidoptera, Zygaenidae). Theses zo- ologicae 30: 159-164. 123 KARAMI, M., NAUMANN, C. M. & TREMEWAN, W. G. T. (1998): The systematics and distribution of Zygaena Fabricius, 1775 (Lepidoptera: Zygaenidae) in Iran. Iran Journal of Entomology 15(1995): 1-22, 1 [ab- stract in Farsi]. 124 KARAMI, M., NAUMANN, C. M. & TREMEWAN, W. G. T. (1998): The biology and ecology of Zygaena Fabri- cius, 1775 (Lepidoptera: Zygaenidae) in the Islamic Republic of Iran. Iran Journal of Entomology 15(1995): 2346, 3 [abstract in Farsi]. 125 EPSTEIN, M., GEERTSEMA, M., NAUMANN, C. M. & TARMANN, G. M. (1999): The Zygaenoidea. Pp. 159-180 in: KRISTENSEN, N. P. (ed.): Handbook of Zoology 4: Arthropoda: Insecta. De Gruyter, Berlin etc. 126 NAUMANN, C. M., TREMEWAN, W. G. & TARMANN, G. (1999): The western palaearctic Zygaenidae. 304 pp., Apollo Books, Stenstrup. 127 FANGER, H., YEN, S.-H., NAUMANN, C. M. (1999): External morphology of the last instar larva of Phauda mimica Strand, 1915 (Lepidoptera: Zygaenoidea, Zy- gaenidae, Phaudinae). Entomologica Scandinavica 30(1998): 429-450. 128 NAUMANN, C. M. (2000): Zygaena carniolica (Sco- poli, 1763) forma flava erstmals im Iran nachgewiesen (Lepidoptera: Zygaenidae). Entomologische Zeitschrift Frankfurt am Main 110: 57. 129 FANGER, H. & NAUMANN, C. M. (2001): The mor- phology of the last instar larva of Aglaope infausta (Lepidoptera: Zygaenidae: Chalcosiinae). European Journal of Entomology 98: 201-218. 130 NAUMANN, C. M. (2001): Biodiversitát — gibt es eine zwelte Chance? Pp. 191-195 in WINNACKER, E.-L., DICHGANS, J., ERKER, G., FRITZSCH, H., HOLLDOBLER, B., WINTERFELDT, E. & DONNER, W. (eds), Unter jedem Stein liegt ein Diamant. Struktur -Dynamik — Evolution. S. Hirzel Verlag, Stuttgart-Leipzig. 131 NAUMANN, C. M. (2001): Regenwald und Savanne. Pp. 31-41 in: TÜRKAY, M. (ed.) Leben ist Vielfalt. Kleine Senckenberg-Reihe no. 41. 132 KLiR, J. & NAUMANN, C. M. (2002): Zygaena (Me- sembrynus) purpuralis pseudorubincundus subsp. nov.: eine weitere eurosibirisch verbreitete Zygaena- Art ım Iran (Lepidoptera: Zygaenidae). Entomolo- gische Zeitschrift Frankfurt am Main 112: 233-236. 133 FÄNGER, H., OWADA, M. & NAUMANN, C. M. (2002): The pupa of Eterusia aedea (Clerck, 1759) (Lepidop- tera, Zygaenidae, Chalcosiinae): morphology and phy- logenetic implications. Deutsche entomologische Zeitschrift N. F. 49: 261-271. 134 KLUTSCH, C. F. C., MISOF, B. & NAUMANN, C. M.. (2003): Characterization of microsatellite loci for Re- issita simonyi (Rebel, 1899), (Lepidoptera, Zygaeni- dae) Molecular Ecology Notes 3: 528-531. 135 NAUMANN, C. M.. (2003): Die hóchste Zygaene der Welt. Entomologische Zeitschrift Frankfurt am Main 113: 362-371. 136 ZAGROBELNY, M., BAK, S., RASMUSSEN, A. V., JOR- GENSEN, B., NAUMANN, C. M., MOLLER, B.L. (2004): Cyanogenic glucosides and plant-insect interactions. Phytochemistry 65 (3): 293-306. 137 KLUTSCH, C. F. C., GROSSE, W.-R., NASHER, A. K., Miısor, B. & NAUMANN, C. M. (accepted): The distri- bution of the yellow-lemon tree frog Hyla savignyi (Audouin, 1827): updates and extensions of previous records (Zoology in the Middle East). 138 PATT, A., MISOF, B., WAGNER, T. & NAUMANN, C. M. (2004): Characterization of microsatellite loci in Am- phitmetus transversus (Kolbe, 1897) (Coleoptera, Cur- culionidae). Molecular Ecology Notes 4, 2: 188-190. 139 SPEIDEL, W. & NAUMANN, C. M. (2005): A survey of family-group names in noctuoid moths (Insecta: Lepi- doptera). Systematics and Biodiversity 2 (2): 191-221. 140 SPEIDEL, W. & NAUMANN, C. M. (2005): The system- atic position of the genus Cetola Walker, 1855 (Lepi- doptera: Noctuidae) Entomologische Zeitschrift 115 (1): 13-14. 141 KUZNETZOV, V. ]., NAUMANN, C. M., SPEIDEL, W. & STEKOL'NIKOV, A. A. (2005): The skeleton and muscu- lature of male and female terminalia in Oenosandra boisduvalii Newman, and the phylogenetic position of the family Oenosandridae (Lepidoptera). SHILAP, Re- vista de Lepidopterologia 32: 297-313. Patt, A., MISOF, B., WAGNER, T. & NAUMANN, C. M.: Isolation and characterization of microsatellite loci in Monolepta vincta Gerstaecker 1871 (Coleoptera, Chrysomelidae, Galerucinae). Molecular Ecology Notes (submitted). KLUTSCH, C. F. C., MISOF, B. & NAUMANN, C. M.: Distribution patterns of the Arabian burnet moth Reis- sita simonyi (Rebel, 1899) and the wall butterfly Lasiommata felix (Warnecke, 1929) in Yemen (in prep.: Fauna of Saudi Arabia). NAUMANN, C. M.: New Zygaenidae from South Af- rica (Lepidoptera, Zygaenidae) (in prep.) NAUMANN, C. M.: Notes on the biology and ecology of Zygaena huguenini Staudinger, 1888, an endemic species of the Tian Shan mountain ranges (Lepidop- tera, Zygaenidae) (in prep.). NAUMANN, C. M. & TARMANN, G. M.: New light on the phylogeny of the Zygaenidae: proposing a new cladogramm for the zygaenid subfamilies. POPULAR ARTICLES (EXTRACT) Dor, R. & NAUMANN, C. M. (1978): Die Kirghisen des afghanischen Pamir. Akademische Druck- und Verlagsanstalt, Graz. Clas M. NAUMANN: in memorian 9 NAUMANN, C. M. (1986): Zur Tierwelt Afghanistans. Pp. 89-191 in: BUCHERER-DIETSCHI, P. & JENTSCH, C. (eds): Afghanistan. Ländermonographie, bearbei- tet von der Arbeitsgemeinschaft Afghanistan. Schrif- tenreihe der Stiftung Bibliotheca Afghanica 4 (Lies- tal). NAUMANN, C. M. (1991): Apollo und Osiris. Oder: Sind wir noch zu retten? Pp. 42-45 in: Baden- Württemberg 5/1991. NAUMANN, C. M. (1991): Gift und Farbsignale im Überleben der Insekten. Tier und Museum 2: 113- 119. NAUMANN, C. M. (1992): Echolot und Signalfäl- schung — Krieg der Sterne in der Sommernacht. Pp. 9— 13 in: GAIDA, K. G. & PROKOT, S. (eds) Microchirop- tera. Falter Verlag, Wien. NAUMANN, C. M. (1995): Terra, quo vadis ? — oder: Ziele und Aufgaben des Naturkunde-Museums in un- serer Zeit. Tier und Museum 4: 55-61. NAUMANN, C. M. (1996): Biologische Vielfalt aus zo- ologischer Sicht. Veróffentlichungen der Deutsch- Chinesischen Gesellschaft e.V., Oktober 1996: 13-19. NAUMANN, C. M. (1996): Wo findet die Zukunft der Menschheit statt? TenDenZen 5: 27-36. NAUMANN, C. M. (1998): Unser blauer Planet — Leben im Netzwerk. Das Museum Koenig: Blick in die Zu- kunft. Das Museum Koenig (Schriftenreihe der Alex- ander-Koenig-Gesellschaft) No. 1: 32-35. NAUMANN, C. M. (1999): Der internationale Code fiir zoologische Nomenklatur — 4. Ausgabe. Entomolo- gische Zeitschrift Frankfurt am Main 109: 353-358. NAUMANN, C. M. (1999): Ras Fartak — 100 Jahre und ein bisschen spáter. Entomologische Zeitschrift Frank- furt am Main 109: 393-408. NAUMANN, C. M. (1999): In Bonn blüht die Biodiver- sitátsforschung. Das Museum Koenig (Schriftenreihe der Alexander-Koenig-Gesellschaft) No. 2: 4-9. NAUMANN, C. M. (1999): Das Museum Koenig auf dem Weg zu neuen Ufern. Kurzfassung einer zehn- jährigen Odyssee. Das Museum Koenig (Schriften- reihe der Alexander-Koenig-Gesellschaft) No. 2: 44— 57. NAUMANN, C. M. (2000): Zu Besuch beim Großfür- sten Nikolai Mikhailovich Romanoff. Entomologische Zeitschrift Frankfurt am Main 110: 12-17. NAUMANN, C. M. (2000): Schakuh - ein Traum wird wahr. Entomologische Zeitschrift Frankfurt am Main 110: 203211, 12 Abb. NAUMANN, C. M. 2000. Jede Art führt ein „Nischen- Dasein”. Leibniz 3/2000: 18. NAUMANN, C. M. (2001): Schattenfreies Fotografieren von präparierten Insekten. Entomologische Zeitschrift Frankfurt am Main 111: 187, 188. NAUMANN, C. M. (2001): Johann Moritz David Herold (1815): „Entwickelungsgeschichte der Schmet- terlinge“. Entomologische Zeitschrift Frankfurt am Main 111: 268-276. NAUMANN, C. M: (2001): Laudatio fiir Herrn Profes- sor Dr. sc. Bernhard Klausnitzer anlásslich der Verlei- hung der Fabricius-Medaille 2001 der DgaaE. Mit- teilungen der deutschen Gesellschaft fiir allgemeine und angewandte Entomologie 13: 7-9. NAUMANN, C.M. (2001): Ein Mullah mit Gazelle zagt in Kabul nie. Frankfurter Allgemeine Zeitung, 21. Dezember 2001. NAUMANN, C. M. (2002): Zum Gedenken an Max Korb (1851-1933) aus Anlass seines 150. Geburt- stages. Entomologische Zeitschrift Frankfurt am Main 112: 12-19. NAUMANN, C. M. (2002): Biodiversität - gibt es eine zweite Chance? Mitteilungen der deutschen Gesell- schaft für Chirurgie 1/02: 49-53. j NAUMANN, C. M. (2002): Wie lernen Fische das Schwimmen ohne Wasser? Frankfurter Allgemeine Zeitung, 21. Juni 2002. NAUMANN, C. M. (2002): Ohne die Erhaltung der Biodiversitát keine erneuerbaren Ressourcen. Pp. 119 123 in: Bayerische Akademie fiir Naturschutz und Landschaftspflege (ed.) Das Ende der Biodiversitát? Grundlagen zum Verstándnis der Artenvielfalt und seiner Bedeutung und der Maßnahmen, dem Arten- sterben entgegen zu wirken (5. Franz-Ruttner-Sym- posium). Laufener Seminarbeitrage 2/02. NAUMANN, C. M. (2002): Im Spannungsfeld — Um- weltforschung. Das Museum Koenig (Schriftenreihe der Alexander-Koenig-Gesellschaft) No. 4: 4-9. NAUMANN, C. M. (2002): BMZ? AA? GTZ? DAAD? Gender-Seminare statt Damen-Toiletten: Die absurden Folgen der deutschen Entwicklungshilfe in Afghani- stan. Frankfurter Allgemeine Zeitung, 21. November 2002, Nr. 271, S. 40. NAUMANN, C. M. (2003): Albertus Seba (1665—1736) Rerum Naturalium = Thesaurus. Entomologische Zeitschrift Frankfurt am Main 113: 355-359. LIST OF DIPLOMA- AND DOCTORAL THESES SUPERVISED BY C. M. NAUMANN Diploma Theses ALTHOFF, Arnhild (2001) „Fragmentierung von Kalk- magerrasen und die Auswirkungen auf die Bestáuber- Pflanzen-Interaktionen am Beispiel von Campanula glomerata“ (University of Bonn, jointly with Dr. M. KRAEMER). BERGFELD, Thomas (1996) ,,Dynamik von Zooplankton und Jungfischen in Altarmen der Sieg“ (University of Bonn, jointly with J. FREYHOF and Dr. K. BUSSE). BISCHOFF, Antje (1995) ,,Verteilungsmuster und Nah- rungsókologie von Jungfischen in Rauschen der Sieg” (University of Bonn). BISSCHOPINCK, Thorsten (1995) ,,Die phytophagen Insek- ten auf Birke (Betula pendula Roth) unter besonderer Beriicksichtigung der Blattminierer im Raum Bonn. Ein Standortvergleich“ (University of Bonn). BÖHMER, Thomas (1994) „Untersuchungen zur Abundanz, Dispersion und genetischen Nachbarschaftsgröße aus- gewählter Populationen des Zygaena transalpina- Superspezies-Komplexes (Insecta, Lep., Zygaenidae” (University of Bonn, jointly with Dr. A. HILLE). Cousin, Markus (1998) „Diversität baumkronenbe- wohnender Arthropoden eines Tieflandregenwaldes im westlichen Uganda (Semliki Forest)* (University of Bonn, jointly with Dr. T. WAGNER). DALBECK, Lutz (1998) „Aspekte der Lebensstrategie von Hasel Leuciscus leuciscus (Linnaeus 1758) und Döbel Leuciscus cephalus (Linnaeus 1758) in der Sieg™ (University of Bonn, jointly with Dr. J. FREYHOF). ESCH, Christoph (1989) „Untersuchungen zur etholo- gischen, genetischen und morphologischen Differen- 10 Bonner zoologische Beitráge 53 (2004) zierung von Zygaena filipendulae (Linnaeus, 1758) und Zygaena lonicerae (Scheven, 1777) (Insecta, Lepidoptera) sowie zu deren Verbreitung in SW- Europa” (University of Bielefeld). FANGER, Harald (1986) ,,Untersuchungen zum Bau des weiblichen Genitalsystems und der Spermatophore von Zygaena trifolii (Esper, 1783)“ (University of Bielefeld). FELTGEN, Kerstin (1998) ,,Bionomie der Seidenbiene Col- letes cunicularius (Linnaeus 1761) (Hymenoptera, Aculeata, Apidae)“ (University of Bonn, jointly with I. BISCHOFF). FIELENBACH, Jörg (1996) „Zeitliche und räumliche Verteilungsmuster der Nasen Chondrostoma nasus (L.) in der Sieg“ (University of Bonn, jointly with J. FREYHOF and Dr. K. BUSSE). FREUND, Wolfram (2000) “Revision of Bonesioides Laboissiere, 1925 (Coleoptera: Chrysomelidae, Galerucinae)” (University of Bonn, jointly with Dr. T. WAGNER). FREY, Susanne (1989) „Altersbedingte Veränderungen in der Feinstruktur des weiblichen Pheromonsystems von Zygaena trifolii (Esper, 1783)“ (University of Biele- feld). FREYHOF, Jörg (1992) „Zeitliche und räumliche Verteilung von Jungfischen in der Fließstrecke der Sieg“ (Univer- sity of Bonn, jointly with Dr. K. BUSSE). GRUNDMANN, Bernd (1984) „Vergleichende, ökologisch- faunistische Untersuchungen zur terricolen Käferfauna der Pflanzengesellschaften eines regenerierenden Ho- chmoores“ (University of Bielefeld). HERDER, Fabian (2000) „Zoogeographie und Okologie zentralvietnamesischer Süßwasserfische“ (University of Bonn, jointly with Dr. K. BUSSE and Dr. J. FREY- HOF). HEUSER, O. (1994) „Zur chemosensorischen Funktion der Pedipalpen bei der juvenilen Jagdspinne Cuppiennius salei (Aranei, Ctenidae)“ (Universities of Wien and Bonn). HILLE, Axel (1986) ,Morphologische und enzymelek- trophoretische Merkmalsdifferenzierung im Zygaena transalpina-Komplex (Lepidoptera, Insecta)” (Univer- sity of Bielefeld). HOLLING, Doris (1992) ,,Vergleichende Untersuchungen von Carabiden- und Curculioniden-Zónosen in gegat- terten und ungegatterten Rotbuchenwáldern im Arns- berger Wald (Sauerland)* (University of Bonn, jointly with Prof.Dr. G. KNEITZ). HOLLING, Michael (1990) „Über die Eiablage, den Repro- duktionserfolg und die Steuerung des des Lockverhal- tens von Zygaena trifolii (Esper, 1783)“ (University of Bielefeld). KLUG, Andrea (1993) „Bestäubungsökologie neotropischer Ericaceen” (University of Bonn, jointly with PD Dr. K.-L. SCHUCHMANN). KnEip, Carsten (1999) „Vergleich der baumkronenbewoh- nenden Arthropodenfauna in Montanwáldern Ostafri- kas* (University of Bonn, jointly with Dr. T. WAG- NER). KOHL, Michael (2001) „Die Arthropoden- und speziell die Käferfauna von Teclea (Rutaceae) in verschiedenen Wäldern Ostafrikas” (University of Bonn, jointly with Dr. T. WAGNER). KURTSCHEID, Agnes (2000) „Revision von Candeza Chapuis, 1879 (Galerucinae, Chrysomelidae, Coleop- tera)” (University of Bonn), jointly with Dr. T. Wag- ner. LEVERMANN, Eva-Maria (1998) „Vergleichend öko- logische Studien an Panurgus calcaratus (Scopoli 1763) und Dasypoda hirtipes (Fabricius 1793) (Hy- menoptera, Apidae) in der Wahner Heide“ (University of Bonn, jointly with I. BISCHOFF). LINDNER, Michael (1991) „Morphologische und taxono- mische Untersuchungen an den Zikaden-Ektoparasiten Heteropsyche schawerdae (Zerny, 1929)“ (University of Bielefeld). MAURER, Golo (2001) “Nestling begging behaviour and the impact of parental alarm-calls in white-browed scrubwrens, Sericornis frontalis (Vigors & Horsfield)” (University of Bonn). MEUSEMANN, Karın (2001) „Karyologische Unter-su- chungen zum Agrodiaetus-Komplex (Lycaenidae)“ (University of Bonn). MIDDELHAUVE, Jens (2000) “Revision of Afrocrania Hincks, 1949 (Coleoptera: Chrysomelidae, Galeru- cinae)” (University of Bonn, jointly with Dr. T. WAG- NER). NEEBE, Bettina (1992) „Der Einfluß von Störreizen auf die Herzschlagrate brütender Küstenseeschwalben (Sterna paradisea)* (University of Bonn). NEMITZ, Armin (1994) „Untersuchung zum Jungfisch- aufkommen in einigen Laichschongebieten der Sieg“ (University of Bonn, jointly with Dr. K. BUSSE). OCKENFELS, Peter (1989) „Untersuchungen zur Attraktion der Imagines von Zygaena trifoli an Dipsacaceen- Blüten“ (University of Bielefeld). OPPERMANN, Markus (1994) „Vergleichende Untersu- chungen zum abdominalen Tracheen-System der Ark- tiidae (Lepidoptera) (University of Bielefeld). PFEIFER, Carmen (1999) „Blumenvögel und Vogelblu- men — wechselseitige Anpassungen und Abhángig- keiten“ (University of Bonn, jointly with Dr. M. KRAEMER). PIEPER, Benno (1991) ,,Untersuchungen zum Reproduk- tionserfolg der Mánnchen bei Weibchenpaarungen zweiten Ranges im Paarungssystem von Zygaena trifo- lii (Esper, 1783) (Lepidoptera, Zygaenidae)* (Univer- sity of Bielefeld). PRINZ, Jochen (1988) „Untersuchungen zum Partner- findungsverhalten von Zygaena trifolii (Esper, 1783) — ein Beitrag zur Reproduktionsbiologie der Zygaeniden (Insecta, Lepidoptera)* (University of Bielefeld). PORSCHMANN, Melanie (1986) „Morphologische und habi- tuelle Parameter im larvalen Entwicklungszyklus von Zygaena trifolii (Esper, 1783)* (University of Biele- feld). RAMMERT, Uwe (1985) ,,Untersuchungen zur Wirksamkeit des Wehrsekretes und des aposematischen Zeich- nungsmusters von Zygaena trifolii (Esper,1783) auf unerfahrene Stare (Sturnus vulgaris, L.)“ (University of Bielefeld). SCHADE, Renate (1991) „Morphologie und phylogene- tische Bedeutung der Siebold'schen Drüse der Nym- phalidae (Lepidoptera)“ (University of Bielefeld). SCHIDELKO, Kathrin (2002) „Neozoen im Großraum Bonn unter besonderer Berücksichtigung der Entenvögel“ (University of Bonn, jointly with Dr. R. VAN DEN EL- ZEN). SCHMIDT-LOSKE, Katharina (1992) „Raumeinbindung und Biotopnutzung tagfliegender Großschmetterlinge am Clas M. NAUMANN: in memorian | | Beispiel des stadtnahen Naturschutzgebietes Rodder- berg südlich von Bonn“ (University of Bonn). SCHMITT, Katrin (2000) „Die Blütenbesucherfauna von Cucurbita pepo: Biogeographische und ethologische Argumente für die effektivsten Bestäuber“ (University of Bonn, jointly with Dr. M. KRAEMER). SCHNEIDER, M. (1994) „Synökologische Untersuchung der mit Lotus corniculatus assoziierten Insektenfauna“ (University of Bielefeld). SCHOLTEN, Matthias (1995) „Verteilungsdynamik und Nahrungsökologie von Jungfischen in Stillwasser- bereichen der Sieg“ (University of Bonn). SCHULZ, Iris (2002) „Multiple Vaterschaft im Freiland: eine molekulare Untersuchung an Zitterspinnen“ (Uni- versity of Bonn, jointly with PD Dr. G. UHL). STAPEL, Heidi (2002) ,,A molecular phylogeny of Afro- tropical Monolepta species and related Galerucinae“ (University of Bonn, jointly with Dr. T. WAGNER and Dr. B. MISOF). STEINMANN, I. (1996) „Überwinterungsbedingungen von Jungfischen in Altarmen der Sieg“ (University of Bonn, jointly with J. FREYHOF and Dr. K. BUSSE). STUHLLEMMER, Claudia (1999) „Saisonaler Einfluß auf die baumkronenbewohnende Arthropodenfauna insbeson- dere der Käfer im Budongo-Forest, Uganda“ (Univer- sity of Bonn, jointly with Dr. T. WAGNER). VELTEN, Guido (2002) „Diversity and abundance of Hy- menoptera (Insecta) in a Guineo-Congolian rainforest fragment“ (University of Bonn). VENEMA, Michael (1986) ,,Untersuchungen tiber den Ein- fluß der Photoperiode auf die Larvalentwicklung von Zygaena trifolii (Esper, 1783)“ (University of Biele- feld). WEON, Hae-Yon (1996) ,,Zeitliche und ráumliche Struk- turen der Evertebratendrift in der Sieg“ (University of Bonn, jointly with J. FREYHOF and Dr. K. BUSSE). WIEMERS, Martin (1994) „Differenzierungsmuster bei Art- bildungsprozessen: Morphologisch-biometrische Un- tersuchungen am Coenonympha arcania (Linnaeus, 1761)-Superspezies-Komplex (Lepidoptera: Nympha- lidae: Satyrinae)“ (University of Bonn). DOCTORAL DISSERTATIONS BISCHOFF, Inge (2000) „Populationsdynamik, Sammel- strategie und Nisthabitatwahl ausgewählter Wildbie- nenarten (Hymenoptera, Apidae) in der Wahner Heide“ (University of Bonn). DALBECK, Lutz (2002) „Der Uhu Bubo bubo (L.) in Deutschland — autökologische Analysen an einer wieder angesiedelten Population, Resümee eines Ar- tenschutzprojektes“ (University of Bonn). DUTREIX, Claude (1995) „Les Zygenides de la region Bourgogne, Ecologie des peuplements des Zygenides en région Bourgogne (Lepidoptera: Zygaenidae)“ (Université de Bourgogne, Dijon — jointly with collea- gues from Dijon). EDELMANN, Alois (1983) _ ,,Enzymelektrophoretische Untersuchungen auf die Frage stammesgeschichtlicher Beziehungen in der Gattung Zygaena Fabricius, 1775 (Insecta, Lepidoptera)“ (University of Bielefeld). FÄNGER, Harald (1994) „Untersuchungen zur Struktur und Funktion des männlichen Reproduktionssystems, der Spermatophore, der Kopulations-Mechanik und des Spermien-Dualismus bei Zygaena trifolii (Esper, 1783) (Insecta, Lepidoptera)“ (University of Bonn). FRANZL, Sylvia (1986) ,,Morphologische und physio- logische Untersuchungen zu Synthese, Transport und Speicherung cyanogener Verbindungen bei Zygaena trifolii (Esper, 1783) (Insecta, Lepidoptera)” (Univer- sity of Bielefeld). FREYHOF, Jórg (1997) ,,Strukturierende Faktoren in der Fischgemeinschaft der Sieg“ (University of Bonn). GRUNDMANN, Bernd (1989) ,,Untersuchungen zur Toxi- kokinetik von Blei und Cadmium im System Zea mays-Spodoptera frugiperda (Lepidoptera) (Univer- sity of Bielefeld). HILLE, Axel (1992) „Quantitativ-analytische Untersuchung zur genetischen Populatiosstruktur und geo-graphi- schen Variation in parapatrischen Kontaktbereichen des Zygaena-transalpina-Subspezies-Komplexes (In- secta, Lepidoptera, Zygaenidae)* (University of Biele- feld). KRAEMER, Manfred (1997) ,,Struktur und Dynamik der Pflanzen-Kolibri-Gemeinschaft von Bajo Calima, Ko- lumbien“ (University of Bonn, jointly with Prof. Dr. W. BARTHLOTT). KREUSEL, Boris (2000) ,,Phylogenetische Analyse der Ctenuchinae (Lepidoptera: Arctiidae)* (University of Bonn, jointly with Dr. C. HAUSER, Stuttgart). OCKENFELS, Peter (1995) „Pflanzen-Insekt-Beziehungen bei Lepidopteren — eine chemisch-dkologische Studie am Beispiel von Zygaena trifolii (Esper, 1783) (In- secta, Lepidoptera)” (University of Bonn). RAMMERT, Uwe (1993) „Morphologische Untersuchungen zur Aufdeckung der stammesgeschichtlichen Ver- haltnisse der basalen Gruppen der ditrysen Lepidop- teren (Lepidoptera, Ditrysa)“ (University of Bielefeld). SCHMITT, Ulrike (2000) „Die Pflanzen-Kolibri-Gemein- schaft im Bergregenwald der Faraloones de Cali, Re- serva Natural Hato Viejo, Kolumbien“ (University of Bonn, jointly with Dr. M. KRAEMER). WAGNER, Thomas (1996) ,,Artenmannigfaltigkeit baum- kronenbewohnender Arthropoden in zentralafri- kanischen Wäldern, unter besonderer Berúcksichti- gung der Käfer“ (University of Bonn, jointly with PD Dr. M. SCHMITT). Author’s address: Michael SCHMITT, Zoologisches Forschungsmuseum Alexander Koenig, Adenauerallee 160, D-53113 Bonn, Germany. E-mail m.schmitt@uni- bonn.de Bonner zoologische Beiträge Band 53 (2004) Heft 1/2 | Seiten 13-26 Bonn, Juni 2005 Neuropterologische Beitráge in den Werken von Wilhelm Friedrich VON GLEICHEN, genannt RUSSWORM (1717-1783)! Horst ASPOCK" und Ulrike ASPÓCK ” ' Abteilung für Medizinische Parasitologie des Klinischen Instituts für Hygiene und Medizinische Mikrobiologie der Medizinischen Universität Wien, Österreich 2 Naturhistorisches Museum Wien, Entomologische Abteilung, Wien, Österreich Abstract. Among VON GLEICHEN’s publications on various topics in zoology, botany, agriculture, and technology there are also three, each of them dealing with a single species of Neuroptera, two appeared in 1764, the third one in 1770. Due to the careful descriptions, and in particular due to the brilliant hand-coloured copper plate engravings, all three species can be identified: Chrysopa pallens (Rambur, 1838) (Chrysopidae) (eggs, larva), Chrysoperla carnea (Stephens, 1836) (Chrysopidae) (adult), and Drepanepteryx phalaenoides (Linnaeus, 1758) (Hemerobiidae) (larva, pupa, adult). None of them has been named by VON GLEICHEN with nomenclaturally available names. The illustrations in the two ar- ticles of 1764 are the first figures of Chrysopidae in the whole scientific literature which can be identified on the species level. The figures of the imago of Ch. carnea, and of the larva and adult of D. phalaenoides are the largest ones ever published. Key words. Neuroptera, early illustrations, Chrysoperla carnea, Chrysoperla pallens, Drepanepteryx phalaenoides, W.F. VON GLEICHEN-RUSSWORM. 1. EINLEITUNG UND BIOGRAPHISCHES Wilhelm Friedrich VON GLEICHEN, genannt RU8WORM, hat in der Geschichte der Biologie und nicht zuletzt auch der Medizin vor allem durch seine 1778 veröffent- lichte „Abhandlung über die Saamen- und Infusi- onsthierchen, und über die Erzeugung, nebst mikrosko- pischen Beobachtungen des Saamens der Thiere in verschiedenen Infusionen“ eine bleibende Spur hinter- lassen. Es handelt sich dabei um ein Werk, das für das Verständnis der Funktion der Spermien von grundle- gender Bedeutung war. In bibliographischen Beschreibungen zu Werken von Wilhelm Friedrich VON GLEICHEN findet man regelmä- Big die Feststellung, dass er als erster Bakterien gefärbt hat. Vermutlich gehen alle diese Behauptungen auf ei- nen tradierten Hinweis in Morton’s Medical Bibli- ography (NORMAN 1993) zurück, wo es heißt „GLEI- CHEN was probably the first to attempt to stain bacteria; he used carmine and indigo“. Es ist uns (bisher) weder gelungen, in seinen Schriften sein „Färbe-Rezept“ zu finden noch diese Behauptung zu bestätigen. Jedenfalls hat VON GLEICHEN Fütterungsversuche mit Ciliaten un- ter Verwendung wasserlöslicher Farbstoffe durchgeführt (AESCHT 2004). Wilhelm Friedrich Freiherr VON GLEICHEN wurde am 14. Jannuar 1717 in Bayreuth geboren. Sein Vater, 1 Clas Michael Naumann zu Königsbrück (26.06.1939 — 15.02. 2004) zum Gedenken Heinrich VON GLEICHEN, war „Hochfürstl.-Branden- burgisch-Kulmbachischer Geheimderath und Oberjäger- meister“ (Anonymus 1784), seine Mutter eine geborene von RUBWORM aus dem Hause Greiffenstein (Der Name wird in manchen Publikationen VON GLEICHENS mit ei- nem, in anderen mit zwei f geschrieben), zu dem das Dorf Bonnland (südlich von Hammelburg in Unterfran- ken) gehörte. „Der Knabe, bei dem Markgraf Wilhelm Pate gestanden hatte, erhielt die höfische Erziehung seiner Zeit. In seinem elften Jahr konnte er seinen Namen kaum schreiben“ (WILLNAU 1926). Möglicherweise dies und jedenfalls längere Aufenthalte bei verschiedenen aristo- kratischen Verwandten und Freunden in seiner Kinder- zeit waren auf die zerrüttete Ehe seiner Eltern zurückzu- führen. Er wurde Page am Taxisschen Hof ın Frankfurt, und in seinem 14. Lebensjahr begann seine überaus er- folgreiche militärische Laufbahn (1740, also im Alter von 23 Jahren, war er Major, 1748 Oberstleutnant), die er jedoch 1756 aus „freien Stücken“ da er „das Hofleben mit seiner Unruhe, seinem Pomp, Luxus und seiner Intrigue recht satt bekommen“ (Bei allen wörtlichen Zi- taten haben wir die ursprüngliche Schreibweise und In- terpunktion belassen) hatte (WILLNAU 1926) — beendete, um sich auf sein Schloss Greiffenstein zurückzuziehen. VON GLEICHEN muss den Mangel an geistiger Ausbil- dung in seiner Kindheit inzwischen reichlich wettge- macht haben, und geistige Aktivität kennzeichneten auch sein weiteres Leben, das immerhin noch 27 Jahre dauern sollte. 1748 war seine Mutter gestorben, wo- 14 Bonner zoologische Beitráge 53 (2004) durch ihm Schloss Greiffenstein und die umliegenden Landereien zugefallen waren; damals nahm er auch den Namen RUBWORM an. Sein Vermögen ermöglichte es ihm, sich nun ganz seinen Interessen zu widmen. Ein Schlüsselerlebnis war die Begegnung mit den „Mikro- skopischen Augen- und Gemütsergötzungen“ von Mar- tin Frobenius LEDERMÜLLER und schließlich mit dem Autor dieses bis heute berühmten Werkes selbst. W. F. VON GLEICHEN besuchte M. F. LEDERMÜLLER 1760 in Erlangen und bestellte bei ihm ein Mikroskop, und Mik- roskope sollten fortan auch zu seinen wichtigsten Ar- beitsbehelfen gehören. Im Jahre 1762 hielt sich LEDER- MÜLLER etwa vier Wochen bei VON GLEICHEN auf Schloß Greiffenstein auf, die beiden Mikroskopiker verbrachten offenbar eine glückliche Zeit, in der sie sich in gemeinsam erlebter Freude und Begeisterung ganz dem Mikroskopieren hingaben. Schon vor seiner Be- gegnung mit M. F. LEDERMÜLLER, unmittelbar nachdem er sich auf sein Schloss zurückgezogen hatte, hatte W. F. VON GLEICHEN eine rege Publikationstätigkeit be- gonnen, wobei er sich mit ganz unterschiedlichen na- turwissenschaftlichen, landwirtschaftlichen und techni- schen Fragen befasste. Dass besonders diese frühen Publikationen manchmal eine genügend kritische Be- trachtungsweise vermissen lassen, darf nicht unerwähnt bleiben. Ab 1760 wurde W. F. VON GLEICHEN ein Mikroskopiker ersten Ranges, ausgestattet mit einer ausgeprägten Beobachtungsgabe und einer ebenso aus- geprägten Lust, das Gesehene zu interpretieren, Schlüs- se zu ziehen und zu veröffentlichen. So entstanden zahl- reiche weitere Publikationen über sehr unterschiedliche naturwissenschaftliche Themen (Bibliographie: FIKEN- SCHER 1801). W. F. VON GLEICHEN starb infolge eines Schlaganfalls in seinem 67. Lebensjahr am 16. (oder 18.) Juni 1783 auf Schloss Greiffenstein. Die Bedeutung W. F. VON GLEICHENS als Naturforscher wurde schon zu seinen Lebzeiten erkannt, und schon unmittelbar nach seinem Tod erschienen Darstellungen seines Lebenswegs: WEIKARD 1783; ANONYMUS 1784; FIKENSCHER 1801. Von den späteren biographischen Publikationen über W. F. VON GLEICHEN sind vor allem die Arbeiten von WILLNAU (1926), WEIDNER (1980), GEUS (1989, 1990) und DETTNER (1997) zu erwähnen. W. F. VON GLEICHEN war nicht nur ein begabter Expe- rimentator, sondern ein ebenso talentierter Zeichner, der seine Publikationen mit vorzüglichen Abbildungen aus- stattete. Seine berühmteste und durch hervorragend ko- lorierte Kupfertafeln ausgestattete entomologische Ar- beit ist ohne Zweifel jene über die „Geschichte der gemeinen Stubenfliege“ (VON GLEICHEN 1764), in der er überdies Beobachtungsfehler von M. F. LEDERMÜL- LER aufzeigte, was bedauerlicherweise zu einer Ent- zweiung der beiden führte. Entomologische Themen werden auch in mehreren an- deren Publikationen VON GLEICHENS behandelt. In zwei Werken setzt sich VON GLEICHEN mit Neuropteren aus- einander, nämlich in dem 1764 erschienene Band über „Das Neueste aus dem Reiche der Pflanzen ...* (Abb. 1) und in seinem 1770 publizierten „Versuch einer Ge- schichte der Blatläuse und Blatlausfreßer des Ulmen- baums ...‘“ (Abb. 9). Das 1764 veröffentlichte Werk ist — wie man schon dem langen Titel (,,... und einem Anhang vermischter Beobachtungen ...““) entnehmen kann — so etwas wie ein Sammelband über verschiedene Themen. Er enthält auch zwei neuropterologische Artikel, von denen der eine („Der Blatlausfreßer und seine Eier an einem Hanfblat“) den präimaginalen Stadien einer Chrysopiden-Spezies, der andere („Die Hofdame“) einer im Winter gefundenen Imago einer Florfliege gewidmet ist. Beofancungen "geheimen Beupemahthik, de der ‘Plane in ihren Wiiten, und einigen Rerjuden von dem Re ii einem Anhang permifGter Seooagtungen, nd mit Garben nad is Natur vorgehehkt Wilhelm Gricderid) Freiberrn von Gleihen, genannt Nufmorm, en Herrn auf Greiffenttein und Bonnlanb, Orof + Creuz des Brandendurgifchen rohten Adler, Ordens umd Hodfürftl. Brandendurgio, Culmbadifhen Ocheimen Rabe. ‚ Herausgegeben, verlegt mit den/mößtigen Im ae ae un» Waminirtes Abbildungen verfchen, Sobann Chriftonh Keller, Gerüst, bd Chelan de Launoy feel. Erb, 1764, Abb. 1: W. F. VON GLEICHEN (1764), Titelblatt des Werks, das die Arbeiten über den „Blatlausfreßer ...“ (1764a) und „Die Hofdame“ (1764b) enthält (Bibl. H. & U. Aspöck) In der 1770 veröffentlichten umfangreichen Arbeit wird die Biologie einer Hemerobiiden-Spezies ausführlich dargestellt. VON GLEICHEN hat sich wenig um die kor- Horst AsPÓCK & Ulrike Aspöck: Neuropterologische Beiträge in den Werken von Wilhelm Friedrich von GLEICHEN 15 rekte Bestimmung und Benennung der von ihm unter- suchten Insekten gekúmmert, vielmehr hat er einfache deutsche Namen verwendet. Sein Augenmerk konzent- rierte sich ganz und gar auf die funktionellen Aspekte, auf Lebensweise und Entwicklung, wáhrend ihn taxo- nomische und systematisch Aspekte offenbar gar nicht interessierten. GEUS (1990) zitiert eine geradezu abfálli- ge Aussage VON GLEICHENs über jene Naturforscher, „die nur Systeme ausheckten, von nichts als Nomenkla- tur, Papillons, Konchylien u.d.g. mit einem Worte, nur immer von der Schale der Dinge schwatzten. Solche Menschen setzen meistens die Naturkunde nur zu einem blossen Gedaechtniswerke oder blos zu einer leeren Liebhaberei herab.“ 2. 1764A: „DER BLATLAUSFREBER UND SEINE EIER AN EINEM HANFBLAT“ (ABB. 2-6) In diesem Aufsatz schildert VON GLEICHEN, wie er Ende Juli 1761 an einer Hanfstaude ein Blatt fand, ,,an dessen beiden Seiten weise subtile Fäden, deren Endungen gleichfärbige kleine Knöpfchen hatten, sehr regelmäßig angeklebet waren“. VON GLEICHEN brach das Blatt ab, nahm es mit nach Hause und fand seine Vermutung, dass es sich bei den „Knöpfchen“ um Insekteneier han- delte, bestätigt, als er bei Betrachtung der Knöpfchen unter dem Mikroskop die durchscheinende Larve sah und plötzlich Zeuge des Schlüpfens der Eilarve wurde. VON GLEICHEN beschreibt Eier und Larven außerordent- lich genau und bildet sie auch in hervorragender Weise ab; seine Zeichnungen dienten Johann Christoph KEL- LER (1737-1795) als Vorlagen für die Kupferstiche (Abb. 2). Zu seinem Bedauern konnte er keine weiteren Zeichnungen anfertigen, weil die Larven nach drei Ta- gen tot und eingeschrumpft waren. VON GLEICHEN wusste zu diesem Zeitpunkt nicht, welche Insekten er vor sich hatte, was ihre Nahrung ist, ja er schreibt sogar ganz offenherzig: „Allein meine Ungewissheit ihrer Be- stimmung verleitete mich, zu glauben, dass sie wenigs- tens nicht viel größer werden würden, als sie waren, da ich eines derselben Fig. 4. vergrößert abbildete.* Erst nach dieser Untersuchung kam VON GLEICHEN das be- rühmte Werk von REAUMUR (1734-1742) in die Hände, in dessen drittem Band (1737) die Biologie von Chryso- piden beschrieben wird. VON GLEICHEN gibt auch un- umwunden zu, dass er, hätte er von REAUMURs Werk vorher Kenntnis gehabt, mit seinen „Blatlausfreßern viel sorgfältiger umgegangen seyn würde“. Erst durch REAUMUR erfuhr VON GLEICHEN , dass die „Fresszan- gen“ nicht zum Fressen, sondern zum Saugen dienen und dass sie „hohl und an den Spitzen offen sind“. Dass diese Saugzangen von größter systematischer Bedeu- tung sind und das die Ordnung Neuroptera begründende Merkmal schlechthin (also in heutiger Terminologie: die Autapomorphie) der Neuroptera sind, wusste auch REAUMUR nicht. Erst mehr als 100 Jahre später erkannte der junge Friedrich BRAUER den außerordentlichen sys- tematischen Wert des Merkmals, indem er alle Familien der Neuroptera (im heutigen Sinn) in einer Ordnung (er nannte sie damals Megaloptera) vereinte (BRAUER & LÖW 1857). a.” o...» Tabula 2, Der Blatläusfreßer und feine Cier an einem Danfbiat, I eines Dicjer Ener befibplftiger mar, Mncte fado folches an feinem ebern QYheile, und eines der defemberfien Chierrden , fo max mod) iemals ver Die agen gefsnmmen @, fam aus bemicibigen hervor. Die tangle, me und mihfame Dereegung defriden gefhattete mur, «4 abrageidmen, ia fo gar ead auspumahlen. . Die Schmale des Cies fant bicbei med) dem Neale, als fie leer wurde, fichtdartich yufamımın umd warde Diciemigen , medie Das entzüchende Dergmigen feamen, das bei Dergicichen Unterfachungen einen gigen fich 1) peri gegencinanber gefrimte Oipigen, mit Denen es Dia Schaaic Des Cies burdo har, 3) fünf im halben Zirfel fichende Pirine Mugen, recidx Das mirriere größere umgeben, und: 5) mie ich mad) ber Deutlich gefchenen Demegung des untern braunen Theiles des Kopfes (dliefe , das Maul a. © Die Begierde, mit diefen fremben Cullen genauer befemt qu werden, verentafete mich foglad, - Nahrung bit dl, fo jeager auch das fürdpterixhe Zangengebif net úl, dag es in der Reade des fícinen Ghermdiraas can chen fo febrectbares Raub thier frp mag , als ber Bar end der Wolf in der Rede größerer Tiere. Und vielleicht erfeyet Die Let, feinen Daz qu fangen, mas iden Die Natur an ber Chefähmindigfieit verfagt hat, mie mir Diefes an Dem Yimenfennluber und Udurmiömen febo, Lm meine Mbdidung Diefes Suits aber mehcers ja eridutern, wil ich cs Nüryäch befcheciben. a Der im Derglcidana mit der Geife des Körpers fchr große und breite Kepf rubet auf einem Fats a gen Hals. Oben it ex mit cinema hormiditem Helm bededtt, defen verderes heil mcisgran, Des beaters aber braun if. An den beiden verdern Seiten des Sopfes chen peel ungeghederte , basdiicheige und geatte Júblbdemer neben den Augen. Unter Diefen raget das Pürdhreriche Zangengebeö ont gelben Sree hervor, Die Magen fichen ‘glach am Anfange Des „ und teemen bier micht ale fechft grichen merden, weit bei des Dritten Giger. Der Ser i ocfärmig , aber, gama Di rah, me pe de » : Abb. 2: W. F. VON GLEICHEN (1764a), Erste Seite der Arbeit über den „Blatlausfreßer ...* Wahrscheinlich verdanken wir indes dem Umstand, dass VON GLEICHEN das Werk von REAUMUR nicht frü- her zur Hand hatte, die schön kolorierte Kupfertafel der Eier und der Erstlarve. (Die auf Fig. 5 und Fig. 6 darge- stellten leeren Eier stammen von einem späteren Fund des Gärtners VON GLEICHENS auf einer Zwetschke.) Un- seres Wissens handelt es ich dabei um die erste farbige Darstellung einer Chrysopiden-Larve mit einer Wieder- gabe des Pigmentierungsmusters in der gesamten ento- mologischen Literatur. Die Darstellung ist so genau, dass der Versuch, die Larve zu bestimmen, lohnend erschien. Die Larven fast aller in Mitteleuropa vorkom- menden Chrysopiden sind gut bekannt, zu überwiegen- dem Teil existieren auch (allerdings nicht durchwegs 16 Bonner zoologische Beitráge 53 (2004) Rohl ce pia Caller 0x Abb. 3: W. F. VON GLEICHEN (1764a), Tafel zu der Arbeit tiber den „Blatlausfreßer ...* Y Abb. 4: W. F. VON GLEICHEN (1764a), Ausschnitt der Tafel zu der Arbeit über den „Blatlausfreßer ...*: Die in (für C. pallens charakteristischen) Reihen abgelegten Eier. Jan. 3: Abb. 5: W. F. VON GLEICHEN (1764a), Ausschnitt der Tafel zu der Arbeit „Blatlausfreßer ...*: Die aus dem Ei schlüpfende Lar- ve (von C. pallens). auch vom ersten Larvenstadium) gute Abbildungen (vgl. KILLINGTON 1936-1937; PRINCIPI 1940; GEPP 1983; DIAZ-ARANDA 1992; DIAZ-ARANDA & MONSER- RAT 1995). Vergleicht man die Larve VON GLEICHENS mit den verfúgbaren Abbildungen, so findet sich die größte Übereinstimmung mit Chrysopa pallens (Ram- bur, 1838). Wir haben den derzeit vermutlich besten Kenner der práimaginalen Stadien von Chrysopiden, Herrn Prof. Dr. Peter DUELLI (Birmensdorf) um eine Beurteilung der Abbildung in VON GLEICHENS Arbeit gebeten. Nach seiner Meinung lassen die in Reihen ab- gelegten Eier sofort auf C. pallens schließen. Ebenso spricht die Pigmentierung der Kopfkapsel am ehesten für C. pallens. Weiters schreibt er: „Die langen, am En- de gebogenen Borsten lassen auf einen trash-carrier schließen. Das ist C. pallens ja nicht. Allerdings sind bei allen Chrysopiden-Arten die Borsten des ersten Larven- stadiums, relativ zur Körperlänge gesehen, viel länger als im 3. Larvenstadium. Auch tragen viele Arten dann doch noch etwas trash (debris) mit sich herum, später nicht mehr. Verglichen etwa mit Chrysoperla-Larven Horst Aspöck & Ulrike Aspöck: Neuropterologische Beiträge in den Werken von Wilhelm Friedrich von GLEICHEN sind bei C. pallens auch bei der L3 die Borsten sehr lang.“ Zusammenfassend hält Prof. DUELLI die Larven in VON GLEICHENS Arbeit mit größter Wahrscheinlich- keit für Chrysopa pallens. Abb. 6: W. F. VON GLEICHEN (1764a), Ausschnitt der Tafel zu der Arbeit über den „Blatlausfreßer ....“: Erstlarve (von C. pal- lens). Diese Art (früher als Chrysopa septempunctata Wes- mael, 1841, bekannt) ist eine extrem expansive, poly- zentrische Art mit Ausbreitungszentren in der West- und in der Ostpaläarktis und einer Verbreitung, die na- hezu ganz Europa (mit Ausnahme der nördlichen Teile, jedenfalls ganz Mitteleuropa von der Ebene bis in die kolline Stufe), NW-Afrika sowie weite (auch tropische) Teile Asiens östlich bis Japan und Taiwan umfasst (ASPOCK et al. 1980; ASPOCK et al. 2001). Mit Sicher- heit war und ist die Art im Bereich des RURWORM’schen Schlosses, wie überhaupt in Franken, häufig. (Zur Verbreitung von Ch. pallens in Deutschland siehe SAU- RE et al 2003.) Diese durch die erste kenntliche Farbabbildung einer Chrysopiden-Larve und die Beschreibung des Schlüpf- vorgangs bemerkenswerte Arbeit VON GLEICHENS ist of- fenbar weitestgehend in Vergessenheit geraten. Selbst in der an Vollständigkeit kaum überbietbaren Bibliogra- phie von OSWALD (2004) fehlt sie. Ältere und z. T. so- gar kolorierte Darstellungen von Chrysopiden-Larven finden sich u. a. bei GOEDART (1669), ALBIN (1720), FRISCH (1736), REAUMUR (1734-1742), BONNET (1745, Übersetzung: GOEZE 1773), sie sind allerdings durch- wegs grob ausgeführt und lassen jedenfalls keine Art- Bestimmung zu. 3. 1764B: „DIE HOFDAME“ (ABB. 7-8) Auch diese Arbeit mit dem ungewöhnlichen Titel fehlt in der Bibliographie von OSWALD (I. c.), obwohl sie die vermutlich größte je publizierte Abbildung einer Chry- sopide enthält. VON GLEICHEN schildert in dem der Zeit entsprechen- den narrativen Stil, wie sich „zur Zeit des Carnevals im Jenner bei später Nacht“ ein Insekt auf eine Spielkarte setzte, das durch seine Schönheit auffiel (weshalb ihm VON GLEICHEN den Namen „Hofdame“ gab). Tags darauf untersuchte VON GLEICHEN das Insekt mit frei- em Auge ebenso wie unter dem Mikroskop und führte eine kolorierte Zeichnung aus, von der J. C. KELLER einen Kupferstich anfertigte, den VON GLEICHEN kolo- rierte. Das Insekt ist von jedem Entomologen sogleich als Chrysopide zu erkennen, und jeder Neuropterologe weiß, dass es sich dabei nur um eine Chrysoperla-Art handeln kann. Die in Mitteleuropa vorkommenden Chrysoperla-Arten úberwintern als Imagines, und unter diesen nehmen manche (in Mitteleuropa als einzige Chrysopiden) im Herbst eine gelblich-bräunlich-rötliche Fárbung an. Vor 40 Jahren ware die Bestimmung vóllig problemlos erschienen. Damals kannte man nur eine einzige Chrysopiden-Spezies unter den Chrysopiden Mitteleuropas, die als Imago úberwintert und im Herbst ihre Fárbung von grún in braun verándert: Chrysoperla carnea (Stephens, 1836). Im Verlauf der vergangenen Jahrzehnte hat sich gezeigt, dass ,,die alte Chrysoperla carnea™ nur als Kollektivnahme für mehrere morpholo- gisch schwierig zu differenzierende Spezies zu betrach- ten ist, von denen mindestens 5 in Europa und z. T. auch in Mitteleuropa vorkommen (Literatur hierzu: HENRY & DUELLI 1999; HENRY et al 1996, 2002; HENRY et al 2003; JOHNSON et al 2003; ASPOCK et al. 2001): Chrysoperla carnea (Stephens, 1836) Chrysoperla lucasina (Lacroix, 1912) Chrysoperla mediterranea (Hölzel, 1972) Chrysoperla pallida (Henry, Duelli & Johnson, 2002) Chrysoperla agilis (Henry, Brooks, Duelli & Johnson, 2003). 18 Bonner zoologische Beitráge 53 (2004) dle Die Hofdame. $ des Degen vom Neaumurs (eye, weldhe er PL 32. Fi natürlichen abbilden : 5 é g- 10, der narlı 9 lies , cbfchom die meinigem, da ich fie brad, nde qua] waren ei bar eae he Kopfes biefer Figur. Shee Verrandlung habe aus fájon : Pc 2 2 H 2 2 3 g 3 2 E A Erklärung der Figuren, Fig. 1. Gin Hamfblat mit Smfeften Ein, Fig. 2, Ein foldes Ei vergrößert. Fig. 3. Der BWlatlausfrefer, rie er aus dem Gi ¡ur Welt fomt, Fig. 4. Derfelbe einen Tag fpáter beobachtet, Fig. 5. Dergleichen Eier auf einem Sriicfdyen Zmetfchendaut vergrößert, Fig. 6. Diefes in natirlicher Größe. Fig. 7. Das von der Zmetfchenhaut loggemachte untere Theil des Cierfadens , defen trichterförmige Geftait peiget, rie der Sal aus bem Fledrichten Anfaje, gleich dem Faden aus dem Spin roden, berausgefponnen if. Tabula 3. Die Hofdame. is olte ich mol micht zu emtfehuldigen fenm, Daf ich diefera fliegenden Snfefte ben Namen des daras des der Höfe beilege, da es cin fo prictiger Schmuck der Natur ift ? Und maffen fic nicht die Diumitafund Cidriner gleiches Recht an, wenn fie ihren Blumen und Früchten fogar Die prächtigen Titel der Kaifer und Könige geben ? Sie felbflen, meine gnddige Damens, werden die erften fen, mir Diefe Vergleidhung zu vergeben , fobald fie einen Whe auf die Vorflelung diese (hönen (Fliege rhum mollen. She eigen Gerviffen wird ihnen fagen, daß máx nur diefe Benennung ungemem mol getroffen fey, fondera Dag auch Diefes Smfett an Pracht und Schönheit fogar in der Vergrößerung mand Hofdamg in ihrer Mobe übertreffe, zumal ram fic bedenfen molen, Dag unter allen ihren Schönheiten mide eme eayige bem Kaufmanne und Schneider jugebdre. Nod) mehr, es gehöret diefes Sufeft gesifermaßen ju Dem Gefchlechte der Sumgfrauen, Es liebet endlid) diefe Hofdame auc das Spiel Damm eben bei dem Spiele war es, als fic fid) zur Zeit des Carmevals im Fenner bei fpiiter Nacht cinfand , und auf die Karte (este. Kaum harte fic fid) ein menig hier umgefehen, als ich he, um Des andern Tages nähere Velantfchaft mit ihr qu machen , fogleich cin Quartier bei meinem WMitroftop ammvies, Die febdnen las ümen Augen, die id) fepon ohne Vergrößerung bei Licht fabe, liefen mid) auf bie Entbedungen mod) Größe vorflelt, wird yeigen, dag id mid) wiht betrogen habe. Am Kopfe fiedt man eine erhabene Stirn, umd unter folder ¡mo von vorne länglicht runde Lippen a, Nu Des unten Lippe gehen vice ges ba gliederte Abb. 7: W. F. VON GLEICHEN (1764b), erste Seite der Arbeit über „Die Hofdame“. Diese fünf Taxa unterscheiden sich vor allem durch spezifische Vibrationsmuster, die sich durch verschie- den langes, durch unterschiedliche Intervalle und Fol- gen gekennzeichnetes Vibrieren mit dem Abdomen ergeben. Die Vibrationen werden auf das Substrat über- tragen und erreichen über dieses andere Individuen, die die spezifischen Signale mit den Subgenualorganen der Beine wahrnehmen. Beide Geschlechter zeigen dieses der Partner-Findung dienende Verhalten, und wenn zwei Phäna mit unterschiedlichen Vibrationsmustern zusammentreffen, kommt es nicht nur nicht zur Paa- rung, in vielen Fällen ist sogar eine aggressive Reaktion die Folge. Die Taxa sind Spezies mit zum Teil unter- schiedlich fortgeschrittener genetischer Isolierung. Sie werden häufig als „song-morphs“ bezeichnet. Unter ex- perimentellen Bedingungen können Paarungen zwischen verschiedenen „song-morphs“ manchmal „erzwungen“ werden. Ch. mediterranea und Ch. lucasina unterscheiden sich morphologisch hinlänglich und konstant von den ande- ren drei Spezies, deren morphologische Unterschiede gering sind. Alle genannten Chrysoperla-Arten über- wintern als Imagines, aber nur drei von ihnen (Ch. car- nea, Ch. pallida, Ch. agilis) verändern im Herbst die Färbung von grün zu rötlich-bräunlich-gelblich. Diese Färbung wird während der gesamten winterlichen Dia- pause bis zum nächsten Frühjahr beibehalten. Für die Deutung der Abbildung der „Hofdame“ (Abb. 8) VON GLEICHENS scheiden also Ch. lucasina und Ch. mediter- ranea jedenfalls aus. Auch die in verschiedenen Teilen des Mittelmeerraums nördlich bis in die Südschweiz nachgewiesene Ch. agi- lis kann ausgeklammert werden, sie ist im übrigen bis- her ın Mitteleuropa nicht nachgewiesen worden und kommt mit großer Wahrscheinlichkeit in diesem Teil Europas tatsächlich nicht vor. Von den verbleibenden zwei Arten — Ch. carnea und Ch. pallida — gilt Ch. pal- lida als „Waldart‘“, die nur selten in Häusern überwin- tert. Ch. carnea hingegen ist ein ausgeprägter Kulturfol- ger und eine in allen Städten und Dörfern Mitteleuropas häufige Florfliege. Auch in diesem Fall haben wir den führenden Chryso- perla-Spezialisten, Prof. Dr. Peter DUELLI (Birmens- dorf), um seine Beurteilung des kolorierten Kupferstichs gebeten. Seine Antwort: „Die Färbung der Hofdame ist exakt das, was ich für C. carnea s. str. erwarten würde. Auch die Tatsache, dass sie im Januar im Haus anflog, spricht für carnea, C. lucasina ist wegen der Färbung ausgeschlossen, C. pallida (Waldart) kommt selten in die Häuser und ist etwas bräunlich-gelber im Winter“. Wir können also tatsächlich davon ausgehen, dass W. F. VON GLEICHEN die erste kenntliche — und überdies far- bige — Darstellung jener Spezies veröffentlicht hat, die STEPHENS (1836) mehr als 70 Jahre später als Chrysopa carnea nomenklatorisch gültig beschrieben hat und die mit Sicherheit die in der Literatur am meisten behandel- te Neuropteren-Spezies ist. Chrysoperla carnea ist ZU- dem gewiss auch die am intensivsten und unter den ver- schiedensten Gesichtspunkten untersuchte Neuropteren- Art. VON GLEICHENs Abbildung ist — obwohl 240 Jahre alt — keineswegs die erste Abbildung einer Florfliege. Darstellungen von Chrysopiden findet man schon bei MOUFFET (1634), ALDROVANDI (1638), JONSTON (1657), GOEDART (1668), FRISCH (1736), RÖSEL VON ROSENHOF (1755), und natürlich bei REAUMUR (1734- 1742) und bei BONNET (1745) und anderen früheren Autoren. Einige Autoren (GOEDART, ALBIN, RÖSEL VON ROSENHOF) haben sogar kolorierte Abbildungen veröf- fentlicht. Allerdings ist eine sichere Identifizierung der Art in keinem Fall möglich. Man muss jedoch gerech- terweise erwähnen, dass drei Umstände die Identifizie- rung der von VON GLEICHEN dargestellten Florfliege ermöglicht haben: die genaue Kenntnis des Fundorts (Schloss Greiffenstein in Unterfranken), das Funddatum (Januar, aktiv im Haus) und die bräunliche (Ver-) Fär- bung des Tiers. Das Geäder hat VON GLEICHEN zwar sehr schön (und sogar — erstmals in der Literatur — mit Horst Aspöck & Ulrike Aspöck: Neuropterologische Beiträge in den Werken von Wilhelm Friedrich von GLEICHEN 19 doppelten Strichen), aber alles andere als richtig ge- zeichnet. Zu jener Zeit wusste man natürlich noch nichts über Geäder-Merkmale von Chrysopiden und deren sys- tematische Bedeutung, weshalb man — das gilt geradezu für alle Autoren jener Zeit — auf eine korrekte Wieder- gabe des Geäderverlaufs und der Zahl und Position der Queradern kaum achtete. Abb. 8: W. F. VON GLEICHEN (1764b), Tafel zu der Arbeit über „Die Hofdame“ (= Chrysoperla carnea). VON GLEICHEN mutmaßte, dass seine „Hofdame“ „die nemliche ist, welche nach der Beschreibung des Herrn von REAUMURs aus der seidenartigen Puppe hervor- komt“. Das ist sicher nicht richtig, aufgrund der biologi- schen Angaben von REAUMUR kann es sich nicht um ei- ne Chrysoperla-Art gehandelt haben. Auch VON GLEICHENs Vermutung, dass seine „Hofdame“ sich aus einem „entkommenen Blatlausfreßer“, der im „Zimmer eine ... anständige Speise“ gefunden habe, entwickelt habe und „dass durch die Wärme geheizter Zimmer, wie dies bei allen sich verwandelnden Insekten geschieht, das Wachstum und Auskriechen der Fliege dergestalt befördert worden ist, daß letzteres im Jenner erfolget, da es nach Herrn von REAUMUR bei den Spätlingen erst im Frühjahr geschieht..* trifft natürlich nicht zu. REAUMUR hat über eine Spezies geschrieben, die nicht als Imago úberwintert. Erstaunlich ist indes, dass VON GLEICHEN offenbar erst kurz vor der Veröffentlichung seines ‚Artikels über die „Hofdame“ (er war damals immerhin schon 47 Jahre alt) eine Florfliege beachtet hat, denn nichts in den bei- den Artikeln, die sich mit Chrysopiden befassen, deutet darauf hin, dass VON GLEICHEN diese Insekten vorher gekannt hatte. Kaum ein an der Natur und insbesondere an Insekten interessierter Mensch in Mitteleuropa be- gegnet nicht schon früh, meist irgendwann in der Kind- heit, einer Florfliege und erkennt die charakteristischen, in etwa 30 Arten in Mitteleuropa vorkommenden Neu- ropteren immer wieder. Auch und besonders an den bräunlich verfärbten Florfliegen kann man im Spät- herbst, Winter oder Vorfrühling kaum vorübergehen, weil sie in fast allen Häusern, die von Gärten oder gar Feldern umgeben sind, vorkommen; oft findet man sie — besonders in alten Häusern — zwischen Fenstern, wo sie manchmal in Ritzen überwintern. In allen größeren entomologischen Werken, die vor 1764 existierten, sind auch Chrysopiden behandelt und auch abgebildet worden. Bei aller Würdigung der Verdienste VON GLEICHENS er- staunt es doch, dass ein in einer so sorgenfreien mate- riellen Situation lebender Mensch mit einer so ausge- prägten Neugierde für die Natur nicht über die entomologischen Standard-Werke, die im 17. und 18. Jahrhundert erschienen waren, verfiigte. Er muss dem- nach zumindest in den ersten Jahren seiner naturwissen- schaftlichen Studien auf Schloss Greiffenstein nur eine bescheidene Bibliothek besessen haben. 4.1770: „VERSUCH EINER GESCHICHTE DER BLATLAUSE UND BLATLAUSFRESSER DES ULMENBAUMS“ (ABB. 9-12) Diese 1770 und in einer 2. Auflage 1787 erschienene, insgesamt 30 Seiten umfassende und mit vier kolorier- ten Kupferstichen ausgestattete Publikation zählt zu den bekanntesten Arbeiten des Autors und ist zumindest von MORTON (1910), WEIDNER (1980), GEUS (1989) und DETTNER (1997) besprochen worden. Im Vorder- grund der Sekundärliteratur steht der Nachweis des Le- benszyklus und insbesondere der Parthenogenese der Ulmenblattlaus, Byrsocrypta ulmi (L.), durch VON GLEICHEN sowie dessen Uberlegungen zu diesem Phá- nomen. Im Zuge der Befassung mit den Blattläusen stieß VON GLEICHEN auf einen der Prádatoren unter den Neuropte- ren, jene charakteristische Hemerobiiden-Spezies, die LINNAEUS (1758) als Hemerobius phalaenoides be- schrieben hat und die spáter in das Genus Drepanepte- ryx Leach transferiert wurde. 20 Bonner zoologische Beitráge 53 (2004) ciner Sejhichfe ver Blafläufe Blarlausfreffer des Ulmenbaums nebfl vier mit Farben erleudteten Kupfertafeln von Wilhelm Friederich Srenberen o. Gleichen genannt Huftvorm, $ fenftein, Bonnland und Ejelbad) , Groß» Ereus des Hobíárflid, Branden sido er Knler Ordens, und Hehfürftlih Brandenburg + Eulmbadifhen Geheimen Math. Mebt einee Dorrede des Herrn Hofraths und Prof. Delius. Gn Kupfer gebradt, und verlegt von Georg Paul Nußbiegel, Múrnberg r gedsucte bei Johann Seinridy Gottfried Bicling. 1770» Abb. 9: W. F. VON GLEICHEN (1770), Titelblatt (Bibl. H.& U. ASPOCK) Etwa sieben Seiten Text und zwei kolorierte Kupfersti- che widmet VON GLEICHEN der Beschreibung der Larve und der Imago von D. phalaenoides (Abb. 10 - 12). Die Zeichnungen hat VON GLEICHEN — wie in seinen Wer- ken durchwegs — selbst angefertigt, die Tafeln wurden von seinem Verleger, dem Kupferstecher Georg Paul NUßBIEGEL (um 1713 — 1786), gestochen. VON GLEICHEN beschreibt, wie er in dem Gewimmel der jungen Blattläuse “einige braune Würmer“ erblickt habe, ,,die sehr ämsig waren, und ihre 6. langen Beine mit ungemeiner Geschwindigkeit zu gebrauchen wusten“. Er hielt „sie sogleich für das, was sie waren, nemlich für diejenige Art Blatlausfreßer, die bei dem Herrn von REAUMUR die zwote Claße machen, und wel- chen er den Nahmen Blatlaus-Löwen, Lion de Pouce- rons, gegeben hat“, womit VON GLEICHEN recht hatte. Er beschreibt die Larve von D. phalaenoides sehr genau und gibt eine gelungene Abbildung (Abb.11: Tab. II, Fig. 21-22). Erwähnung verdient auch seine Bemerkung über den Geruch der Larven, er sei „außerordentlich stark, und dabei sehr angenehm, nichts anders als Quän- del und Thimian. Vielleicht ist er die Witterung der Blatläuse die sie locket, ihrem Feinde in der Nähe zu bleiben.“ Tatsächlich haben die Larven von D. phalae- Der Umenbaum, der, wegen feiner did belaubten Jmeige, eine Zierde der Garten it, gibt einer Are Infeleen , die wir Blatläufe nennen, auf feinen breiten Blix teen Wohnungen , in folder Menge birnförmiger Blafen, daß fid) » wenn fic in der Mitte des Vradmonate igre Dollommenbeit erlanger haben, Aefte un? Jrocige davon biegen. ; Die erfte Figur der erflen Tafel peiget cin foldes Blac, das an der Spige einige dies fer Blafen von verfhledencr Größe erage. Die erften habe ich yu Anfang des Manmonare, da fe nod) fehr flein waren, aufgemador. Id) fand cin einiges braunes, didlelbiges, ungefliigel che Meines Infeft darlunen, meldyes in allem fo viel ähnliches mic den fo genannten Blarldufen parte, daf es mir nicht rer fiel, es dasor yu erfennen. Leuwenbord, Sartfoeker, x : de Abb. 10: W. F. VON GLEICHEN (1770), erste Seite (mit der von „J.d.G.R.“ [= J. VON GLEICHEN-RUSSWORM? Ein Verwandter?] gezeichneten und von Johann NUSBIEGEL (1750-1833) gesto- chenen Vignette. noides einen charakteristischen Geruch, den FULMEK (1941) mit dem von Anis- oder Fenchelöl vergleicht. VON GLEICHEN beschreibt das Verhalten der Larven sehr treffend und widmet sich dann recht ausführlich den Mundwerkzeugen. Er lobt auch an dieser Stelle zu- nächst die Arbeiten von REAUMUR, schreibt aber dann, dass er - REAUMUR — „gleichwohl bei der Untersuchung erstgedachter Kopfteile des Blausfreßers [sic!], ohnfehl- bar aus Mangel recht brauchbarer Vergrößerungswerk- zeuge, dießmal von seiner gewöhnlichen Bahn abge- kommen“ sei. Und weiter meint er, er habe daher „viel vergebliche Zeit und Mühe angewendet, auch hier [sei- ne] Beobachtungen“ mit jenen von REAUMUR „zu ver- einigen, ohne die Oefnungen an den Spitzen der Fress- zangen des Blatlausfreßers zu finden“. Darauf zitiert er REAUMUR im Original, der völlig richtig Bau und Funk- tion der Saugzangen beschreibt. Merkwürdig, dass VON GLEICHEN bei der Beschreibung der Mundwerkzeuge von Drepanepteryx phalaenoides nicht auf seine 1764 veröffentlichte Untersuchung über den „Blatlausfreßer .. an einem Hanflblat“ und auf das Bezug nimmt, was er dort über REAUMURS Meinung über die „Fresszan- gen“ gesagt hat, nämlich, dass sie nicht zum Fressen, Horst AsPÓCK & Ulrike Aspöck: Neuropterologische Beiträge in den Werken von Wilhelm Friedrich von GLEICHEN 21 £ TAB. m. Abb. 11: W. F. VON GLEICHEN (1770), Tafel III. mit der Larve des , Ulmenblatlausfressers* (= Drepanepteryx phalaenoides). TAB.TV Abb. 12: W. F. VON GLEICHEN (1770), Tafel IV: mit Puppenko- kon (Fig. 30) und Imago (Fig. 31-33). sondern zum Saugen dienen und „hohl und an den Spit- zen offen sind“ (siehe oben). Vielmehr kam er jetzt zu dem Schluss, dass sie an den Spitzen keine Öffnungen haben, weshalb er zwischen den Saugzangen nach der Mundöffnung suchte und schließlich eine solche auch fand und sogar abbildete (Abb.11: Tab. III, Fig. 23). Dazu schreibt er: ,,Da aber das Maul dieses Insekts nur alleine zum Saugen, und nicht zum Nagen gebrauchet wird, so ist auch der genaue Zusammenschlus der Lip- pen eine natiirliche Folge dieser Bestimmung, so, wie der Unmöglichkeit, solches auf andere Art, als durch ei- nen wohl angebrachten Druck des Kopfes zu öfnen und zu sehen.“ Die akribische präparative und mikroskopi- sche Arbeit VON GLEICHENS verdient alle Bewunderung. Die Nahrung wird bei allen Neuropteren-Larven über die Saugzangen aufgenommen, die Mundöffnung ist ei- ne zwischen den Saugzangen quer verlaufende, falzartig verschlossene, kaum erkennbare Spalte ohne jede Funk- tion für die Nahrungsaufnahme. Wie hat sich VON GLEICHEN die Nahrungsaufnahme wohl wirklich vorge- stellt, da er meinte, das die „Fresszangen“ keine Öff- nung haben, das Maul aber zum Saugen dient. Es ist tat- sächlich erstaunlich, mit welcher Bestimmtheit VON GLEICHEN die richtige Deutung der Saugzangen durch REAUMUR verwirft und durch eine durchaus falsche und unlogische Behauptung ersetzt. Dann beschreibt VON GLEICHEN die Entwicklung von D. phalanoides, gibt die Larvalperiode mit 13 bis 14 Tagen an, erwähnt den „Bemerkungswürdige[n] Umstand, bey diesem Insekt, daß es dieses mit den Spinnen gemein hat, aus dem Ende oder der Spize seines Leibes, und nicht, wie andere Insekten aus dem Munde zu spinnen, und zwar nur zu der Zeit, wenn man das Gespinnste zu seiner Verwandlung anfängt, ohne zuvor jemals nur das geringste Merkmal, daß er hiezu geschickt sey, gegeben zu haben.“ VON GLEICHEN gibt eine schöne Abbildung des Kokons (Abb. 12: Tab. IV, Fig. 30) und erwähnt dann, dass zwei Imagines geschlüpft seien. Es folgt eine ausführliche und anschauliche Beschreibung der Imago, ergänzt durch die schöne kolorierte Kupfertafel. Ge- rechterweise muss man aber sagen, dass die Art zwar sofort zu erkennen ist (nicht zuletzt, weil es in Europa keine andere nur halbwegs ähnliche Hemerobiiden- Spezies gibt und die zweite in Europa vorkommende Art des Genus, D. algidus (Erichson in Middendorf, 1851), sich habituell erheblich unterscheidet), dass aber das Geäder recht fehlerhaft gezeichnet ist. Auch hier gilt das bei VON GLEICHENs Abhandlung über die „Hofda- me“ Gesagte. Im 18. Jahrhundert unterschätzte man ein- fach die Gesetzmäßigkeiten des Flügelgeäders von In- sekten erheblich und achtete bei bildlichen Wiedergaben vielmehr darauf, den Habitus möglichst typisch zu erfas- sen — was VON GLEICHEN ja auch recht gut gelungen ist. REAUMUR (1737) hat die Larve, den Kokon mit der Puppe und die Imago des 21 Jahre später von LINNAEUS 22 Bonner zoologische Beitráge 53 (2004) (1758) als Hemerobius phalaenoides giltig beschriebe- nen Insekts in Form sehr vereinfachter Zeichnungen dargestellt. Aber auch die nicht wenigen Abbildungen von Drepanepteryx phalaenoides, die nach VON GLEI- CHENs Monographie im 18. Jahrhundert erschienen, rei- chen qualitativ nicht annáhernd an die (trotz des er- wähnten Fehlers der Darstellung des Flügelgeäders) außergewöhnlich sorgfältig und vom Standpunkt der Ästhetik — damals und noch lange später — geradezu un- erreichbaren Abbildungen des Schlossherrn von Greif- fenstein heran. GEUS (1989) bemerkt zu Recht, dass es unverständlich ist, dass die Arbeit VON GLEICHENs nicht nur unter dem Gesichtspunkt ihres aphidologischen, sondern auch un- ter jenem des neuropterologischen Inhalts vergessen o- der ignoriert worden ist. Ganz stimmt dies aber doch nicht, weil sie von MORTON (1910) in seinem Aufsatz über die Biologie von D. phalaenoides ausführlich be- sprochen und von KILLINGTON (1936-1937) bei der Be- handlung der Art erwähnt wurde. 5. SCHLUSSBETRACHTUNGEN In der Geschichte der Neuropterologie nehmen VON GLEICHENS Arbeiten vor allem durch ihre prächtig kolo- rierte Kupferstiche mit den geradezu riesig dargestellten Insekten, die in allen drei Fällen eine Bestimmung der Spezies zulassen, einen besondern Platz ein. Seine sorg- faltigen Beschreibungen von morphologischen Merkma- len, Entwicklung und Verhaltensweisen der von ihm un- tersuchten Neuropteren — Chrysopa pallens (Rambur), Chrysoperla carnea (Stephens) und Drepanepteryx pha- laenoides (Linnaeus) — haben daher umso größeren Wert. Die meisten Abbildungen und Beschreibungen von Chrysopiden und Hemerobiiden durch frühere Au- toren sind — von D. phalaenoides abgesehen — keinen wirklich identifizierbaren Spezies zuzuordnen. Dennoch muss man zugeben, dass VON GLEICHENS Arbeiten kei- ne Erkenntnisse enthalten, die so grundlegend neu wá- ren, dass sie die weitere neuropterologische Forschung entscheidend und nachhaltig beeinflusst hátten. Dies ist wohl der wichtigste Grund dafiir, dass diese Publikatio- nen fast oder sogar vóllig in Vergessenheit geraten sind. Ein anderer Grund mag darin zu finden sein, dass die Bücher VON GLEICHENS durch die vielen kolorierten Ta- feln auch zur Zeit ihrer Veróffentlichung (und nicht nur heute) teuer und daher nur einem kleinen Kreis von Le- sern zugánglich waren. VON GLEICHEN hat alle drei neuropterologischen Arbei- ten nach dem Jahr 1758 veröffentlicht. Hätte er die von ihm untersuchten Chrysopiden mit lateinischen Namen versehen, so hatten diese nomenklatorische Giiltigkeit erlangt, und er ware der Autor der in der gesamten en- tomologischen und insgesamt biologischen Literatur am haufigsten behandelten Neuropteren-Art geworden — oder auch nicht, da er ja der Meinung war, dass sein „Blatlausfreßer an einem Hanfblat und seine Hofdame ein und dieselbe Art darstellten. Dass er seinen ,,Ulmen- Blatlausfreßer“ nicht dem von LINNAEUS 1758 be- schriebenen Hemerobius phalaenoides zugeordnet hat, ist erstaunlich. VON GLEICHEN legte auf die Beschreibungen neuer Ar- ten wahrscheinlich wirklich keinen Wert, doch muss man auch bedenken, dass ihm die Vorkenntnisse und wissenschaftliche Ausbildung fehlten. Es ist unwahr- scheinlich, dass er die lateinische Sprache (in der die meisten entomologischen und jedenfalls alle systemati- schen Werke damals verfasst wurden) beherrschte, hin- gegen hat er mit Sicherheit fließend Französisch ge- sprochen, weil dies die Sprache der Aristokratie des 18. Jahrhunderts war. Das in französischer Sprache ge- schriebene Werk von Rene Antoine Ferchault de REAUMUR (1683-1757) hat er daher, nachdem es ihm zu Beginn der 60er Jahre in die Hände gekommen war, oft und gerne benützt, und er hat immer wieder seine Be- wunderung für dieses Werk — das tatsächlich einen Mei- lenstein in der Geschichte der Entomologie darstellt — zum Ausdruck gebracht. VON GLEICHEN muss tatsächlich nicht nur ein von wirk- licher Neugierde, die Natur zu erforschen, durchdrunge- ner, scharfsinniger und unerhört geduldiger Beobachter und Experimentator und hochbegabter Zeichner gewe- sen sein, sondern einfach ein ungewöhnlich intelligenter Mensch, so dass er es letztlich geschafft hat, einen Platz in der Geschichte der Naturwissenschaften zu erobern, obwohl er in seiner Kindheit nicht einmal eine handfeste Schulausbildung genossen hatte und erst jenseits des 40. Lebensjahrs begonnen hatte, sich für die Natur zu inte- ressieren. Dies erklärt auch die manchmal erstaunliche Einschät- zung des Neuheitswerts seiner „Entdeckungen“ — ein Phänomen, das man auch heutzutage immer wieder fin- den kann. Eine schon in der Kindheit oder frühen Ju- gendzeit einsetzende Neugierde für die Natur (oder mu- tatis mutandis für irgendein anderes Gebiet) und eine solide und lange Ausbildung führen zunächst zur frühen Vertrautheit mit dem, was andere vorher erforscht und entdeckt haben und bewahren weitgehend vor der Mei- nung, dass alles, was man nicht selbst kennt, grundsätz- lich unbekannt ist und eine neue Entdeckung darstellt. Unter diesem Gesichtspunkt ist manche überaus selbst- sichere, unbekümmert anmutende Aussage, ist aber auch manche abfällige und geradezu hochmütig anmu- tende Feststellung VON GLEICHENs zu sehen. Die Kenntnis dessen, was andere schon längst oder erst kürzlich vor einem selbst gefunden haben, ist oder zumindest war bis vor kurzer Zeit — abgesehen von einer gründlichen Ausbildung — an eine umfangreiche Bibliothek oder zumindest an den Zugang zu einer sol- Horst AsPÓCK & Ulrike Aspöck: Neuropterologische Beiträge in den Werken von Wilhelm Friedrich von GLEICHEN = 23 chen gebunden, heute kann das mehr und mehr durch den Zugang zu elektronischen Datenbanken ersetzt wer- den. Nicht ersetzbar sind indes die möglichst früh ein- setzende Befassung mit den Objekten der Neugierde und eine intensive Ausbildung, die zwar praktischerwei- se durch Gymnasien und Universitäten vermittelt wird, aber — wie viele eindrucksvolle Beispiele der Wissen- schaftsgeschichte und sogar das des „spät berufenen“ Autodidakten Wilhelm Friedrich VON GLEICHEN zeigen — nicht notwendigerweise an diese Institutionen gebun- den ist. VON GLEICHEN ist — als schon 43jähriger — bei Martin Frobenius LEDERMÜLLER (1719-1769) „in die Lehre gegangen“, und er hat allmählich Kenntnis von den früheren Autoren erhalten. Dass er den von ihm so sehr verehrten und aufrichtig bewunderten R. A. F. DE REAUMUR voreilig und unbegründet kritisiert hat (siehe oben), ist bemerkenswert. Wie immer auch: Wilhelm Friedrich VON GLEICHEN ge- bührt jedenfalls auch in der Geschichte der Neuroptero- logie ein markanter Platz — was mit der vorliegenden Studie gezeigt und bekräftigt werden sollte. Widmung. Wir widmen diese Arbeit dem Andenken unseres Freundes Clas Naumann. Die Liebe zu den alten Büchern unserer Wissenschaft und das Interesse für die Geschichte der Entomologie haben wir mit ihm geteilt, und viele Gespräche mit ihm waren diesem Thema gewidmet. Noch zu Beginn des Jahres 2004 — wenige Wochen, bevor sein Leben zu Ende ging — haben zwei entomologisch-historische Bücher mehrere Telefonge- spräche geprägt. Zum einen befasste er sich zu Ende des Jahres 2003 mit einer Rezension des im Taschen-Verlag erschienenen und von einem Wissenschaftler-Team un- ter der Federführung von Prof. Dr. Rainer Willmann wissenschaftlich betreuten Prachtwerkes über die Tafeln der 1734-1765 erschienenen Rerum Naturalium Thesau- rı von Albertus Seba. Die Rezension (NAUMANN 2003a) erschien im Dezember 2003; er schickte uns noch ein Exemplar des Hefts der Entomologischen Zeitschrift mit persönlicher Widmung, es sollte die letzte Arbeit sein, die wir von ihm bekommen haben, und — zusammen mit dem, im selben Heft veröffentlichten spannend zu le- senden Aufsatz über „Die höchste Zygaene der Welt“ (NAUMANN 2003b) wohl auch die letzte seiner Publika- tionen, die zu seinen Lebzeiten erschienen ist. Zum zweiten erzählte er in den ersten Januartagen 2004 einem von uns (H.A.) von einem besonderen Geschenk, das er zu Weihnachten bekommen habe: ein eben er- schienenes Buch über die Augsburger Naturforscher des 18. u. 19. Jhdts. Dieses Buch (PFEUFFER 2003) hat ihm in seinen letzten Lebenswochen tiefe Freude gegeben. Wir haben es sogleich erworben und auch noch mit ihm darüber gesprochen; es ist tatsächlich für jeden, dem die Namen Jacob HÜBNER, Gottlieb Tobias WILHELM und Christian Friedrich FREYER etwas sagen, ein intellektu- eller Genuss ersten Ranges. Wir haben Clas NAUMANN erstmals im März 1964 bei der Entomologen-Tagung in München getroffen, wir kannten ihn also genau 40 Jahre. Viele gemeinsame In- teressen haben zu vielen Begegnungen und vielen sub- stantiellen Gesprächen geführt, in denen wir immer wieder seine hohe Intelligenz, seine schier unerschópfl- che Arbeitskraft, seine unnachgiebig kompromisslose Einstellung zur Mittelmäßigkeit in der Wissenschaft, seine außerordentliche Hilfsbereitschaft und seinen un- glaublichen Einsatz für die, die seiner Hilfe bedurften, bewundern konnten. Wir sind dankbar für die lange Zeit unserer Freundschaft mit ihm, und wir danken ihm für die intellektuelle und menschliche Bereicherung, die wir durch ihn erfahren haben. Danksagung. Unser besonderer Dank gilt Herrn Prof. Dr. Peter Duelli (Eidgenössische Forschungsanstalt WSL, Birmensdorf, Schweiz) für seine Beurteilungen der Abbildungen in den beiden Arbeiten VON GLEI- CHENS, die sich mit Chrysopiden befassen. Prof. Duelli ist einer der international führenden Experten sowohl für präimaginale Stadien von Chrysopiden generell als auch für das Genus Chrysoperla, dessen Arten und song-morphs. Herrn Dr. Hieronymus Dastych (Zoologi- sches Institut und Zoologisches Museum der Universität Hamburg) danken wir für die Überlassung eines Origi- nals der Arbeit von H. WEIDNER (1980). 6. ZUSAMMENFASSUNG W. F. VON GLEICHEN, ein fränkischer Edelmann, der nach einer erfolgreichen Militärlaufbahn und einem lu- xuriösen Hofleben erst um sein 40. Lebensjahr seine Leidenschaft für die Erforschung der Natur entdeckte und sich fortan mit vielen verschiedenen zoologischen, botanischen, landwirtschaftlichen und technischen Fra- gen beschäftigte und ab 1757 regelmäßig über seine Forschungen publizierte, wurde schließlich einer der herausragenden deutschen Mikroskopiker der 2. Hälfte des 18. Jahrhunderts. Er verfasste auch drei Arbeiten, die drei verschiedenen Neuropteren-Arten gewidmet waren. Alle drei Publikationen sind mit sehr schön ko- lorierten Kupferstichen ausgestattet, die Darstellungen und Beschreibungen sind so gut und jedenfalls ausrei- chend, um in allen Fällen eine Artbestimmung zuzu- lassen. VON GLEICHEN hat die von ihm untersuchten Insekten nicht mit nomenklatorisch gültigen Namen versehen, sondern ihnen deutsche Namen gegeben. Die erste Arbeit (1764) behandelt die gestielten Eier, den Schlüpfvorgang und die morphologischen Merkma- le der Erstlarve des 74 Jahre später von RAMBUR (1838) beschriebenen Hemerobius pallens, heute Chrysopa pal- lens (Chrysopidae). Die zweite Arbeit (1764) ist der Beschreibung einer im Winter im Zimmer gefundenen bräunlichen Florfliege gewidmet. Es handelt sich dabei um jene Art, die STE- 24 Bonner zoologische Beitráge 53 (2004) PHENS (1836), also 72 Jahre spáter als Chrysopa carnea, heute Chrysoperla carnea, beschrieben hat. Ch. carnea ist mit Abstand die am besten untersuchte und in unzáh- ligen Publikationen behandelte Neuropteren-Art. Die Abbildung dieser Art durch VON GLEICHEN in Form eines kolorierten Kupferstiches ist wahrscheinlich die álteste bis zur Art identifizierbare Darstellung einer - Florfliege und zugleich die größte jemals publizierte Il- lustration einer Chrysopidae. Die dritte Arbeit ist 1770 und in einer zweiten Auflage 1787 als kleine Monographie der Ulmenblattlaus (Bry- socyrpta ulmi [Linnaeus,1758]) erschienen; in ihr wer- den auch der 12 Jahre vorher von LINNAEUS (1758) als Hemerobius phalaenoides beschriebene Prádator Drepanepteryx phalaenoides (Hemerobiidae) unter morphologischen, biologischen und ethologischen Ge- sichtspunkten ausführlich beschrieben und als Larve und Imago in Form kolorierter Kupferstiche abgebildet, auch diese Illustrationen fallen durch ihre Größe aus dem Rahmen. Über diese Arbeit VON GLEICHENS gibt es wegen ihres bedeutenden aphidiologischen Inhalts Se- kundärliteratur, die beiden ersten Artikel scheinen aller- dings völlig in Vergessenheit geraten zu sein. 7.SUMMARY W. F. VON GLEICHEN, a Franconian nobleman, discove- red his deep interest and ardour for exploring nature a- round his 40th year of life — after a successful military career and luxurious life at various courts. From that time onwards he spent a lot of time with various zoolo- gical, botanical, agricultural and technical questions, published regularly on results of his research work and became one of the outstanding German microscopists of the second half of the 18th century. Among his publica- tions, there are also three, each dealing with a certain species of Neuroptera, which impress also because of the brilliant hand-coloured copper plates. The descripti- ons and illustrations are of a quality which allows an identification of the species. The first paper (1764) deals with the stalked eggs, hat- ching and morphological characters of the first instar larva of that species of Chrysopidae (Green lace-wings) which was described by RAMBUR (1838) as Hemerobius pallens, today Chrysopa pallens. This is the oldest il- lustration of a larva of a species of the family Chrysopi- dae which can be identified on the species level. The second paper (also published in 1764) deals with a reddish — brownish lacewing which was on the wings in a warm room in January. The illustrations and the cir- cumstances of the finding clearly allow to identify the species as that one which STEPHENS (1836), 72 years la- ter, described as Chrysopa (today: Chrysoperla) carnea. Ch. carnea is by far the most intensively studied lace- wing, and countless publications deal with it. VON GLEICHEN’s illustration is the first one of a Chrysopid in the whole literature which allows the definite identifica- tion of the species figured. Moreover, the illustration is apparently the largest figure of a neuropteroid insect ever published. The third publication (which appeared first in 1770 and in a second edition in 1787) is a small monograph on the elm aphid (Brysocrypta ulmi [Linnaeus 1758]), which also deals with a predator of the aphid, a neuro- pteroid species described by LINNAEUS (1758) as Heme- robius (today: Drepanepteryx) phalaenoides. Larva, co- coon with the pupae, and the adult are described from a morphological, biological, and ethological view. The larva and the imago are beautifully figured. Again, these illustrations are of an unusual large size in the whole neuropterological literature. This monograph by VON GLEICHEN has been cited and even treated in the later li- terature, mainly for its aphidiological contents on seve- ral occasions, the two other articles have, however, been forgotten entirely. LITERATURVERZEICHNIS: AESCHT, E. (2004): Lust und Last des Bezeichnens — Uber Namen aus der mikroskopischen Welt. In: ASPOCK, U. (wiss. Red): Entomologie und Parasitologie. Fest- schrift zum 65. Geburtstag von Horst ASPOCK. Denisia 13: 1-640. ALBIN, E. (1720): A Natural History of English Insects. II- lustrated with a Hundred Copper Plates, Curiously Engraven from the Life: And (for those who desire it) Exactly coloured by the Author. 100 tab. + 100 pp., William & John Innys, London. ALDROVANDUS, U. (1638): De animalibus insectis libri septem cum singulorum iconibus adviuum expressis. 767 pp + Index, Denuo impress. Bonon: Apud Cle- mentem Ferrorium. ANONYMUS (1784): Auszug aus der Lebensbeschreibung des Herrn Baron Wilhelm Friedrich VON GLEICHEN, genannt RUBWORM. Schriften der Berlinischen Gesell- schaft naturforschender Freunde 5: 491-496. Im Ver- lag der Buchhandlung der Realschule, Berlin. ASPOCK, H., ASPÖCK, U. & HÓLZEL, H. (1980): Die Neu- ropteren Europas. Eine zusammenfassende Darstellung der Systematik, Okologie und Chorologie der Neurop- teroidea (Megaloptera, Raphidioptera, Planipennia) Europas. Mit 96 Bestimmungsschlüsseln, 12 Tabellen, 913 Strichzeichnungen, 259 Fotografien, 26 Aquarel- len und 222 Verbreitungskarten. 2 Bde: 495 pp.; 355 pp. Goecke und Evers, Krefeld. ASPOCK, H., HOLZEL, H. & Aspöck, U. (2001): Kommen- tierter Katalog der Neuropterida (Raphidioptera, Me- galoptera, Neuroptera) der Westpaläarktis. Denisia 02, Biologiezentrum OO. Landesmuseums, Linz: 606 pp. +6 figs. BONNET, Ch. (1745): Traite d’insectologie, ou observations sur quelques especes de Vers d’eau douce et sur les Pucerons. 2 vol: 228, 232 pp. + 8 tab., Durand, Paris. BRAUER, F. (1857) [Unter Mitarbeitung von F. Löw]: Neu- roptera austriaca. Die im Erzherzogthum Oesterreich Horst AsPOCK & Ulrike Aspöck: Neuropterologische Beiträge in den Werken von Wilhelm Friedrich von GLEICHEN 25 bis jetzt aufgefundenen Neuropteren nach der analyti- schen Methode zusammengestellt, nebst einer kurzen Charakteristik aller europäischen Neuropteren-Gattun- gen. 80 pp. + 5 tab., Druck und Verlag von Carl Ge- rold’s Sohn, Wien. DETTNER, K. (1997): Geschichte der Entomologie im Raum Bayreuth/Nordostbayern. Mitteilungen der Deutschen Gesellschaft für allgemeine und angewand- te Entomologie 11: 33-43. DIAZ-ARANDA, L. M. (1992): Estadios larvarios de los Neurópteros ibéricos (Insecta, Neuroptera: Chrysopi- dae). 305 pp., Dissertacione, Universidad de Alcala, Facultad de Ciencias. DIAZ-ARANDA, L. M. & MONSERRAT, V. J. (1995): Aphi- dophagous Predator Diagnosis: Key to genera of Eu- ropean Chrysopid Larvae (Neur. Chrysopidae). Ento- mophaga 40: 169-181. DUELLI, P. (1999): Honigtau und stumme Gesánge: Habi- tat- und Partnersuche bei Florfliegen (Neuroptera, Chrysopidae). In: H. ASPOCK (Wiss. Red.): Neuropte- rida: Raphidioptera, Megaloptera, Neuroptera/Kamel- hálse Schlammfliegen Ameisenlówen. Stapfia 60:35- 48. FIKENSCHER, G. W. A. (1801): Gelehrtes Fiirstenthum Bai- reut oder biographische und literarische Nachrichten von allen Schriftstellern, welche in dem Fürstenthum Baireut geboren sind und in oder ausser demselben ge- lebet haben und noch leben in alphabetischer Ordnung. 3. Band: 64-76. J. J. Palm, Erlangen. FRISCH, J. L. (1736): Beschreibung von allerley Insecten in Teutschland, nebst nützlichen Anmerkungen und nöthigen Abbildungen von diesem Kriechenden und Fliegenden Inländischen Gewürme, zur Bestätigung und Fortsetzung der gründlichen Entdeckung, so eini- ge von der Natur dieser Creaturen heraus gegeben, und zur Ergäntzung und Verbesserung der andern. Vierdter Theil. XXI. Von der Gold=äugigen Stinck=Fliege. pp. 40-41, Tab. XXIII. 45 pp. + 2pl., Nicolai, Berlin. FULMEK, L. (1941): Über die Aufzucht von Drepanepteryx phalaenoides L. ex ovo. (Neuroptera: Planipennia, Hemerobiidae). Arbeiten über morphologische und ta- xonomische Entomologie 8: 127-130, 2 figs. GEPP, J. (1983): Freilanddiagnose mitteleuropäischer Chry- sopidenlarven. Mitteilungen des Naturwissenschaftli- chen Vereins für Steiermark. 113: 101-132. GEUS, A. (1989): Die Untersuchungen des Bayreuther Na- turforschers Wilhelm Friedrich VON GLEICHEN gen. RUßWORM (1717-1783) zur Fortpflanzungsbiologie der Rüsternblasenlaus, Tetraneura (= Byrsocrypta) (1) Ulmi (Linnaeus 1758) Hartig 1841. Berichte der Na- turwissenschaftlichen Gesellschaft Bayreuth 20: 55- 68. GEUS, A. (1990): Naturforscher und Naturforschung im ge- lehrten „Fürstenthum Baireuth“. Berichte der Natur- wissenschaftlichen Gesellschaft Bayreuth 21: 219-234. GOEDART, J. (1669): Metamorphosis Naturalis Pars Secun- da. 288 pp.+16 pp. Register + 50 tab., Medioburgi Apud Jacobum Fierens Biblioplam. GOEZE, J. A. E. (1773): Herrn Karl BONNETs Abhandlun- gen aus der Insektiologie. Aus dem Franzósischen übersetzt und mit einigen Zusätzen herausgegeben. 414 pp. +5 tab., J. J. Gebauer Wittwe & Joh. Jac. Ge- bauer, Halle. HENRY, C. S., BROOKS, S. J., JOHNSON, J. B. & DUELLI, P. (1996): Chrysoperla lucasina (Lacroix): a distinct spe- cies of green lacewing, confirmed by acoustical analy- sis (Neuroptera: Chrysopidae). Systematic Entomolo gy 21: 205-218. HENRY, C. S., BROOKS, S. J., DUELLI, P. & JOHNSON, J. B. (2002): Discovering the True Chrysoperla carnea (In- secta: Neuroptera: Chrysopidae) Using Song Analysis, Morphology, and Ecology. Annals of the Entomologl- cal Society of America 95: 173-191. HENRY, C. S., BROOKS, S. J., DUELLI, P. & JOHNSON, J. B. (2003): A lacewing with the wanderlust: the European song species „Maltese“, Chrysoperla agilis, sp.n., of the carnea group of Chrysoperla (Neuroptera: Chry- sopidae). Systematic Entomology 28: 131-147. JOHNSON, J. B., DUELLI, P., HENRY, C. S. & BROOKS, S. J. (2003): Recognition of cryptic species in the Chryso- perla carnea group (Neuroptera: Chrysopidae) and the quest for the “true” C. carnea. — 8" International Sym- posium on Neuropterology, Texas A&M University, Department of Entomology (Texas), 26-29 July 2003: 26-28. JONSTON, J. (1657): Historiae naturalis de insectis libri III, de serpentibus et draconibus libri II, cum aeneis figu- ris. 147 pp. + 28 tab. + 27 pp. +12 tab., Apud Joan- nem, Jacobi Fil. Schipper, Amstelodami. KILLINGTON, F. J. (1936-37): A monograph of the British Neuroptera. 2 vols.: x1 + 269 pp. (15 plates with 68 fi- gures) and xii + 306 pp., Ray Society, London. LEDERMULLER, M. F. (1760): Mikroskopische Gemiiths- und Augen-Ergótzung: Bestehend, in Ein Hundert nach der Natur gezeichneten und mit Farben erleuchte- ten Kupfertafeln, sammt deren Erklárung. 204 + 4 (Register) pp. + 100 Tafeln, Adam Wolfgang Win- terschmidt, Nürnberg. LINNAEUS, C. (1758): Systema Naturae per regna tria na- turae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Edi- tio decima reformata. Tom. I, 823 pp., L. Salvi, Hol- miae. MORTON (1910): Life-history of Drepanepteryx phalae- noides, Linn. The Entomologist’s Monthly Magazine. 2. Ser., 21: 54-59. MOUFET, Th. (1634): Insectorvm sive Minimorum Anima- lium Theatrum: Olim ab Edoardo Wottono. Conrado Gesnero. Thomaqve Pennio inchoatum: Tandem Tho. Movfeti Londinatis opera sumptibusq; maximis con- cinnatum, auctum, perfectum: Et ad vivum expressis Iconibus supra quingentis illustratum. 340 pp., Thom. Cotes, London. NAUMANN, C. M. (2003a): Schátze der entomologischen Weltliteratur 3. Albertus SEBAs (1665-1736) Rerum Naturalium Thesaurus. Entomologische Zeitschrift 12: 355-359. ANN C. M. (2003b): Die hóchste Zygaene der Welt. Entomologische Zeitschrift 12: 131-147. NORMAN, J. M. (1993): Morton’s Medical Bibliography. An annotated Check-list of Texts Illustrating the History of Medicine (Garrison and Morton). 5" editi- on, 1243 pp., Scolar Press, Gower Publishing Compa- ny, Aldershot. OSWALD, J. (2004): Bibliography of the Neuropterida. A Working Bibliography of the Literature on Extant and Fossil Neuroptera, Megaloptera, and Raphidioptera (Insecta: Neuropterida) of the World. Version 7.03. (entowww.tamu.edu/research/neuropterida/neur bib- liography/bibhome.html) NAU 26 Bonner zoologische Beitráge 53 (2004) PFEUFFER, E. (Hrsg.) (2003): Von der Natur fasziniert ... Frúhe Augsburger Naturforscher und ihre Bilder. 192 pp., Wißmer-Verlag, Augsburg. PRINCIPI, M. M. (1940): Contributi allo studio dei neurotte- ri italiani I. Chrysopa septempunctata Wesm. e Chry- sopa flavifrons Brauer. Bollettino dell’Istituto di En- tomologia della R. Universita di Bologna 12: 63-144. RAMBUR, M. P. (1838): Faune entomologique de l’Andalousie, Vol 2. Tafel 9. Arthus Bertrand, Librai- re-Editeur, Paris REAUMUR, R.A.F. de (1734-1742): Mémoires pour servir a l’Histoire des Insectes. 7 vol. Imprim. Royale, Paris | Tom. 3, 2. Partie, Bl. 32 u. 33, Text: 171-174.] ROSEL VON ROSENHOF, A. J. (1755): Der monathlich- herausgegebenen Insecten-Belustigung Dritter Teil worinnen ausser verschiedenen, zu den in den beeden ersten Theilen enthaltenen Classen, gehörigen Insec- ten, auch mancherley Arten von acht neuen Classen nach ihrem Ursprung, Verwandlung und anderen wun- derbaren Eigenschafften, aus eigener Erfahrung be- schrieben, und in sauber illuminirten Kupfern, nach dem Leben abgebildet vorgestellet werden. 4 pp. Vor- rede + 624 pp. + 8pp. Register +109 tab., J. J. Fleischmann, Niirnberg. SAURE, C. unter Mitarbeit von R. BELSTEDT, A GRUPPE, R. GUSTEN, W.ROHRICHT und E. J. TROGER (2003): Ver- zeichnis der Netzfliigler (Neuroptera) Deutschlands. In: B. KLAUSNITZER : Entomofauna Germanica, Bd. 6. Entomologische Nachrichten (Dresden), Beiheft 8: 282-291. [SEBA, A. (1734-1765)]: Das Naturalienkabinett. Locuple- tissimi rerum naturalium thesauri 1734-1765. Nach dem Original aus der Koninklijke Bibliotheck, Den Haag. 588 pp., Taschen, Köln etc. STEPHENS, J. F. (1836): Illustrations of British entomology; or, a synopsis of indigenous insects: containing their generic and specific distinctions; with an account of their metamorphoses, times of appearence, localities, food and economy, as far as practicable. Vol VI, Man- dibulata. 240 pp., Baldwin F. Craddock, London. VON GLEICHEN, W. F., genannt RUßWORM (1764): Das Neueste aus dem Reiche der Pflanzen oder Mikrosko- pische Untersuchungen und Beobachtungen der ge- heimen Zeugungstheile der Pflanzen in ihren Bliiten, und der in denselben befindlichen Insekten; nebst eini- gen Versuchen von dem Keim, und einem Anhang vermischter Beobachtungen, beschrieben, und mit Farben nach der Natur : Tabula 2, p. 3-5. 16 + 72 + 40 +8+26pp. +51 tab., J. Chr. Keller, Nürnberg. VON GLEICHEN, W. F., genannt RUBWORM (1764a): Der Blatlausfreßer und seine Eier an einem Hanfblat. In: Das Neueste aus dem Reiche der Pflanzen oder Mikro- skopische ‚Untersuchungen und Beobachtungen der geheimen Zeugungstheile der Pflanzen in ihren Blüten, und der in denselben befindlichen Insekten; nebst eini- gen Versuchen von dem Keim, und einem Anhang vermischter Beobachtungen: Tabula 2, p. 3-5. J. Chr. Keller Nürnberg. VON GLEICHEN, W. F., genannt RURWORM (1764b): Tabula 3: Die Hofdame. In: Das Neueste aus dem Reiche der Pflanzen ... Anhang vermischter Beobachtungen. Ta- bula 3, p. 5-6 J. Chr. Keller, Nürnberg. VON GLEICHEN, W. F., genannt RU8WORM (1764c): Ge- schichte der gemeinen Stubenfliege, nebst 4 mit Far- ben erleuchteten Kupfertafeln. 34 pp + 4 tab., J. G. Keller, Nürnberg. VON GLEICHEN, W. F., genannt RUBWORM (1770): Versuch einer Geschichte der Blatläuse und Blatlausfresser des Ulmenbaums nebst vier mit Farben erleuchteten Kup- fertafeln. Nebst einer Vorrede des Herrn Hofraths und Prof. DELIUS. 8 pp. Vorrede + 28 pp. + 2 pp. Erklä- rung, Georg Paul Nußbiegel Nürnberg. VON GLEICHEN, W. F., genannt RUBWORM (1778): Abhand- lung über die Saamen- und Infusionsthierchen, und über die Erzeugung, nebst mikroskopischen Beobach- tungen des Saamens der Thiere in verschiedenen Infu- sionen. Adam Wolfgang Winterschmidt, Nürnberg. WEIDNER, H. (1980): Entomologische Schriften der Zeit- genossen und Schiiler von Carolus LINNAEUS im Ar- chiv der entomologischen Sammlungen des Zoologi- schen Instituts und Zoologischen Museums der Universität Hamburg. Entomologische Mitteilungen aus dem Zoologischen Museum Hamburg 6, 108/109: 309-379. WEIKARD, M. A. (1783): Biographie der Herrn Wilhelm Friedrich VON GLEICHEN genannt RUSSWORM, Herrn auf Greifenstein, Bonnland und Ezelbach Frankfurt. 92 pp., Frankfurt am Main. WILLNAU, C. (1926): LEDERMÜLLER und VON GLEICHEN- RUBWORM. Zwei deutsche Mikroskopisten der Zopf- zeit. 24 pp., Verlag Kurt Scholtze Nachf., Leipzig. Anschrift der Autoren: Univ. Prof. Dr. Horst ASPOCK, Abteilung für Medizinische Parasitologie des Klini- schen Instituts für Hygiene und Medizinische Mikrobio- logie der Medizinischen Universität Wien, Kinderspi- talgasse 15, A-1095 Wien, Osterreich, horst.aspoeck@ meduniwien.ac.at; Univ. Prof. Dr. Ulrike ASPOCK, Na- turhistorisches Museum Wien, Entomologische Abtei- lung, A-1014 Wien, Osterreich, ulrike.aspoeck@nhm- wien.ac.at Bonner zoologische Beitráge Band 53 (2004) | Heft 1/2 | Seiten 27-36 Bonn, Juni 2005 On the Biology of the Ivy-Bee Colletes hederae Schmidt & Westrich, 1993 (Hymenoptera, Apidae)’ Inge BISCHOFF”, Esther ECKELT'? & Michael KUHLMANN” Alexander Koenig Research Institute and Museum of Zoology, Bonn, Germany Institute of Landscape Ecology, University of Miinster, Miinster, Germany Abstract. The biology of Colletes hederae Schmidt & Westrich, 1993, a bee species oligolectic on ivy (Hedera helix), was studied in late summer 2003 at a nesting’aggregation in Dirmstein near Griinstadt (Pfalz), Germany. We analysed the diurnal foraging rhythm, pollen carrying capacity, rate of reproduction, nest architecture, requirements on the nesting habitat and dependence on climatic parameters. In addition, the distribution of C. hederae is revised and represents an adriato-mediterranean distribution type. All known localities of this species fall within the distributional area of Hedera helix. Colletes hederae seems to prefer inclined and southward exposed nest sites in sand or loess soils. The females of C. hederae started their first trip on average at 9.52 h in the morning, made 5.5 provisioning trips and returned from their last trip at 17.09 h. The mean duration of all provisioning trips was 60 minutes but the first flight in the morning was significantly longer (90 min) than the subsequent trips of the day. This is interpreted as additional adult feeding on nectar after a night in the nest. The duration of provisioning trips decreased with increasing temperature. The oligolecty of C. hederae was confirmed. Colletes hederae collected on average 0.5 million grains per pollen load and the cells con- tained 2.8 million grains. Four nests were excavated; the main burrow ran between 7 and 12 cm horizontally into the steep face and the first cells occurred at a depth of 30-45 cm. Beyond a depth of 60 cm no cells were found. Between four and eight cells branched off the vertical part of the main burrow; side burrows were only found in one nest. During the extraordinarily warm and dry season of 2003, the females completed up to 18 cells. Key words. Distribution, diurnal foraging rhythm, pollen carrying capacity, reproduction rate, nest architecture 1. INTRODUCTION Colletes hederae was described as a new species only in 1993 (SCHMIDT & WESTRICH 1993), based on speci- mens collected in Germany and Croatia (Fig. 2). The species appears to be uncommon and is known from few localities. This species is also recently recorded from Great Britain, Luxembourg, The Netherlands, Bel- gium and Spain (FEITZ 2001, PEETERS et al. 1999, PETIT 1996, RATHJEN 1998). Colletes hederae was probably recognized as a distinct species so recently because all species of Colletes are morphologically very similar and it particularly resembles C. succinctus (Linnaeus) and C. halophilus Verhoeff. Furthermore, C. hederae is ac- tive in autumn when most other bees have finished their life cycle and ivy was not considered as an attractive pollen source for bees before. In the past 20 years important new data on the biology of bees were gathered and published. In Germany, Paul WESTRICH (1989) set the standard for many other sub- sequent books dealing with the biology of bees (BELL- MANN 1995, SCHMID-EGGER et al. 1995, MULLER et al. 1997). Taking a more global view, “The bees of the World” by Charles D. MICHENER (2000) set a milestone in the history of literature on bees at the beginning of the millennium. Nevertheless, exact data on the biology of many of the more than 16,000 species worldwide or 1 In commemoration of Clas Michael Naumann zu Königsbrück (26.06.1939 — 15.02.2004) about the 550 German species of solitary bees are scarce or not available. Likewise, the biology of the newly de- scribed Colletes hederae is largely unknown. Biological data are needed to support taxonomic decisions as in the case of C. hederae, C. halophilus and C. succinctus, but also to understand the mechanisms of speciation. Lastly, good knowledge of the biology and habitat requirements of the species facilitates conservation measures. The aim of the present study was therefore to investigate the biology of Colletes hederae including the diurnal for- aging rhythm, pollen carrying capacity, rate of reproduc- tion, nest architecture, requirements on the nesting habitat and dependence on climatic parameters. Furthermore the current distribution of the species is reviewed. 2. MATERIALS AND METHODS 2.1. Study area The study area is located in southwestern Germany near the city of Griinstadt (Pfalz) in a village called Dirm- stein (49°34°04°’N, 08°14°22”°E). The nest aggregation of C. hederae was discovered in 1994 (NIEHUIS pers. comm. Bonn 2003, SCHMID-EGGER 1997) and is situ- ated on a steep face of loess below a vineyard. The steep face has a total length of approximately 150 m and is exposed southwards. The climate is sub-oceanic with mild winters, a summer maximum of precipitation (59- 71 mm per m? and year) and an annual mean tempera- ture of 10.2° C. 28 Bonner zoologische Beitráge 53 (2004) 2.2. Abiotic parameters Climatic parameters were measured with data loggers (Orion Tiny Logger Manager OTLM Tinytalk©). The soil temperature was taken half-hourly at a 20 cm depth on the steep face, air temperature and atmospheric hu- midity were recorded every quarter of an hour in a port- able weather station at a height of 2 m. Data of the “Deutscher Wetterdienst — Station Mannheim” were also used. The soil type was characterized using a soil particle size analysis and we determined the soil den- sity. Furthermore we calculated the inclination of the steep face. The host plant Hedera helix LINNAEUS was mapped in a radius of 500 m around the nesting aggre- gation. 2.3. Ethological observations We observed the behaviour of 16 different females for five days during September 2003. Marking tests at the beginning of the season showed that the females were very sensitive to this interference (cf. BISCHOFF et al. 2003) and consequently we decided to mark only the nests of the respective females. Nests were marked with coloured toothpicks. We recorded the time of first de- parture, number of foraging trips, last return and digging activity. A total of 168 data sets were documented. Since the activity of most bees is extremely influenced by weather conditions (BISCHOFF et al. 2003, LARSSON 1991, LIND 1968) we created a measure for the bees’ ac- tivity independent from weather conditions (cf. MULLER 1994) which is referred to as a so-called “bee-day”(BD). One bee-day corresponds to a day with optimal weather conditions, which females could use completely for provisioning activities. The observed flying activity on days with optimal conditions was 8 hours. The mean temperature at those optimal days amounted to 17°C and the duration of sunshine was between 8 and 10 hours. 2.4. Pollen analysis For pollen analysis, 15 females of C. hederae were cap- tured and transferred in vials with 70% ethanol. To re- move the complete pollen load from the bees, all body parts (legs and sometimes thorax without wings) were sonicated in vials filled with a liquid medium (cf. BUCHMANN & SHIPMAN 1990). Furthermore we exca- vated 20 cells of C. hederae, removed the cell lining, larva and stored the pollen load in ethanol. The number of grains per pollen load or cell was determined using an electronic particle counter (Casy© cell counter and Analyser). To calculate the mean number of pollen grains per pollen load and cell, only particles between 20 and 30 um were considered since they represent the grain size of Hedera helix. All pollen load and cell sam- ples were qualitatively checked with a scanning electron microscope. The pollen samples were treated with Di- ethylether to remove pollen cement and with 1,1,1,3,3,3,-Hexamethyldisilazam to avoid deflation. 2.5. Nest excavations Only nests with observed female activity were exca- vated at the end of the season. Due to the high inclina- tion, nests could not be poured out with plaster. We ex- cavated the nests by following the main burrow from the nest entrance and removed with a spoon the soil lay- ers very carefully until the first cell appeared. 3. RESULTS 3.1. Abiotic characteristics The observed nesting aggregation extended over a breadth of 8 m of the steep face. The height of the steep face amounted to 8.65 m at the studied aggregation. The base of the steep face ran with an inclination of 130° up to a height of 2.65 m and above this height the angle was nearly 90°. At a height of 2 m a line of natural grass cover could be identified and a total of 30% of the stud- ied aggregation was covered with vegetation. The mean values of air temperature, soil temperature, humidity and hours of sunshine at the study area (onsite measurements) and from the nearest weather station are documented in Table 1. No significant differences be- tween our data and the data of the local weather station at Mannheim were found. Table 1: Mean climatic parameters measured during five field days in September 2004 (DWD = local weather station at Mann- heim) Air temp. (°C) Soil temp. (°C) Humidity Hours of sunshine On-site data 22.76 + 4.42 21.48 + 2.70 3953 = 1090 DWD 21.703,58 39.38 + 11.88 9,71 £1.06 The mean soil density amounted to 1.18 g/cm? (n = 6) and the grain size analysıs revealed a typical loess soil comprising 78% silt, 15% clay and 7% sand. 3.2. Distribution of C. hederae Colletes hederae has been recorded from the following localities in nine countries of southern and western Europe (Fig. 1): Inge BISCHOFF, Ester ECKELT, Michael KUHLMANN: On the Biology of the Ivy-Bee 29 of A y A 9 va >| rs E we SY ] = je f E zZ N ARE EE t N E \ De lo x fi \ eo) / ió Se of © A f | Y re AN RO OS dal h go > = af Oy ® 7 In = . y A = he 0 > $ ta ur I IT S6 SIS A - a ~ So ? d <& * ; 8 C ae as. & u > o oy u = | - ) 2 (Sam IO > Sar : A > MI SS, DAR BE » —” | ; © A ae 2 =$ > zo Y E > ri 7 J = | ( La Be Fr; y a te ’ E y 2 = x OS Y a = | \ \ N r >, et Ms 7 po ie A) ( | Fig. 1: Distribution area of Hedera (shaded) (after MEUSEL 1978) and records of Colletes hederae. Belgium: Bassenge (PETIT 1996); Croatia: Lopar: Varenna; Luxembourg: Remerschen, Schwebsange, Francelici/Matahlici, Rovinj: 2 localities; France: Ile- de’Oléron, Le Caylar, St. Chaptes, St. Jean-du-Gard (SCHMIDT 8 WESTRICH 1993), Bathernay, Buoux: Le Fort, Fontvieille, Lacoste, Maussane-les-Alpilles, Para- dou, Pargny, Ratieres, Rempoul: near St. Pol, Rustrel (WIERING 1999), Cherbourg Peninsula: north coast (ED- WARDS & TELFER 2001), Avelmes, Corse: Caleraggio; Germany: Karlsruhe-Durlach (SCHMIDT & WESTRICH 1993), Alsheim, Bockenheim, Dirmstein, Flórsheim, Jockgrim, Monsheim, Nackenheim, Oppenheim (SCHMID- EGGER et al. 1995), Oestrich-Winkel (TISCHENDORF 1997), Burkheim (WESTRICH & DATHE 1997), Ihringen (SCHWENNINGER, pers. com.); Great Britain: Guernsey (Channel Island): 38 localities (SCHMIDT & WESTRICH 1993, EDWARDS & TELFER 2001, BWARS data base), East Prawle, Isle of Portland, Lulworth: 3 localities, Ringstead Bay, Swanage: 6 localities, Weymouth, Worth Matravers: 3 localities (CROSS 2002, BWARS data base), Branscombe, Corfe Castle, Herm (Channel Island), Jer- sey (Channel Island), Kimmeridge, Langton Matravers, Lihou (Channel Island), Osmington Mills, Preston Wins- low, Sark (Channel Island): 4 localities, Ulwell: Godling- ston Hill, Wareham, Wool, (BWARS data base); Italy: Bolzano, Isola del Gilio, Merano (SCHMIDT & WESTRICH 1993), Lazio: Albano (WIERING 1999), Calci, Mte. Gar- gano: M.S. Angelo, Sardinia: Alghero, Sardinia: Oliena, Stadtbredimus, Wintrange (FEITZ 2001); Netherlands: Maastricht, Botanical Garden Lichtenberg (LEFEBER 1998); Slovenia: Kras: Brje pri Komnu, Kras: Gorjansko (GOGALA 1991), Hrastovlje (GOGALA 1994); Spain: Gi- rona: Les Preses (RATHJEN 1998); Switzerland: Biasca: near river Brenno, Chable-Perron: Cheseaux, Lalden: banks of river Rhone near Taleia, Peney, Russin: 2 locali- ties, Sion, Yverdon-les-Bains (AMIET et al. 1999, CSCF data base), Les Bioux: Lac de Joux (AMIET pers. com.). Fig. 2: Colletes hederae male (drawing by Florian Thiimmel). 30 Bonner zoologische Beitráge 53 (2004) The distribution of C. hederae extends from central It- aly, Sardinia and northeast Spain in the south, to south- west Germany, Netherlands and the southern parts of Great Britain in the north. The east-west range 1s from Slovenia to the English Channel. On the whole C. hederae represents an adriato-mediterranean distribution type (LATTIN 1967). All known localities of this species fall within the distribution of Hedera helix but its range is much smaller than that of its host plant (Fig. 1). Hedera helix occurs from North Africa in the south to southern Scandinavia in the north and from the Canaries and United Kingdom in the west eastwards to the coast of the Black Sea (MEUSEL 1978). 3.3. General behaviour of males and females The first males patrolling at the aggregation were ob- served on the 24" of August. The males often inter- rupted their searching flights with inspections of the nest entrances of the previous years. The first flight was repeatedly preceded by grooming at a nest entrance. On the subsequent days the males appeared successively earlier in the morning at the aggregation, e.g. on the 1* of September the first male was recorded at 8.30 h. An obvious correlation between male activity and sunshine was observed. When the sun disappeared, the activity of the males stopped immediately and recurred directly when the sun came out again. The first females were discovered on the 1* of Septem- ber, but it was a matter of only a few individuals and the ivy was not yet in bloom. Only five days later, on the 6" of September, several hundred females were recognised. An emerging female was pounced upon at once by sev- eral males trying to mate and they formed a characteris- tic copulation cluster. The copulation cluster frequently tumbled down to the base of the steep face where mat- 18:20 + Last return 17:30 + N=17 16:40 + 15.50 3 15:00 + 14:10 + 13:20 + 12:30 4 y 11:40 + 0 First departure N=27 09:10 + 08:20 4 2 10:50 4 10:00 + Time ing with one male took place. Females that had already mated were mostly ignored by patrolling males. This behaviour could be observed directly after copulation. The phase of copulation lasted only from the 6" to 10" of September. Shortly after having mated the females started to dig and provision a nest. Many females apparently chose old nests and expanded the old burrows. Other females favoured to dig a new nest, mainly at places that were hidden behind grass or other vegetation. Traces of dig- ging activity could be observed sometimes in the morn- ing at the nest entrances, which meant that the females continued with nest construction after their last foraging trip on the previous day. The females did not close the nest entrance after leaving. Usually they found the en- trance immediately when they returned from a provi- sioning trip. However, changes to the last detail (e.g. marking with a toothpick) were followed by orientation flights in the shape of increasing zigzag flights. If the next approach was not successive, the females rested in the surrounding area and searched on the ground with their antennae, or rested on neighbouring vegetation and started a new approach after a few minutes. None of the observed females constructed a second nest at the ob- served aggregation. No parasites were observed at the aggregation. 3.4. Diurnal foraging rhythm and reproduction The females started their first trip at 9.52 h in the morn- ing, after waiting and trilling with their antennae a few minutes at the nest entrance. The earliest observed flight started at 09.07 h and the latest at 12.19 h. On average C. hederae made 5.5 provisioning trips and returned from its last trip at 17.09 h. The earliest return was at 16.19 h and the latest at 17.53 h (Fig. 3). The mean du- 220 4 200 4 8 180 4 160 + 140 | 120 + 100 4 N = 168 Duration of provisioning trip [min] Fig 3: Time range of main activity data as the departure for the first trip, the last return to the nest and the duration of provisio- ning flights. Inge BISCHOFF, Ester ECKELT, Michael KUHLMANN: On the Biology of the Ivy-Bee 3 E = bee in the nest (BN) u = provisioning trip (PT) first departure V first PT 09:52 (n=26) 08:00 09:00 10:00 11:00 12:00 13:00 last vo " BN 3.5 min. (n=147) other PT (n=142) 17:09 (n=18) 14:00 15:00 16:00 17:00 18:00 19:00 Time (MET) Fig. 4: Ethogram of an average daily foraging cycle of C. hederae. ration of all provisioning trips was 60.3 + 39.9 min (n = 169). However, the first flight in the morning was sig- nificantly longer (90 min) than the other trips of the day (Wilcoxon-test: Z = -2.767, p = 0.006, n = 16). The ac- tivity in the nest after one provisioning trip amounted to 3.5 + 1.7 min (n = 165). No significant correlation be- tween the activity in the nest and the preceding provi- sioning trip could be detected. A scheme of the average foraging behaviour of C. hederae is documented in an ethogramm (Fig. 4). The estimation of the time that the bees used for provi- sioning behaviour independent of weather conditions revealed 12 complete and 12 half = 18 BDs. Assuming five provisioning trips for one cell (see below), C. Table 2 : Correlation of activity data with climatic parameters (pt = hederae could approximately complete between 12 and 18 cells during the season of 2003. 3.5. Dependence on climatic parameters We correlated the different activity data (departures, length of provisioning trips and nest activity) with cli- matic parameters. The results are presented in Table 2. A highly significant negative correlation between the duration of provisioning trips and temperature was found: the higher the temperature, the shorter the provi- sioning trip (Fig. 5). The correlation between the time of first departure and temperature represents only the increasing number of starting bees with time, paral- lel to the increasing temperature and humidity with time. provisioning trip) Temperature (2m) Soil temperature Humidity Duration of pt n 99 99 99 r -0.290 0.089 0.155 p 0.004* 0,379 0.126 Time in nest n 146 146 146 r 0.101 0.001 -0.167 p 0.223 0.986 0.044* Time of 1" trip n 26 26 21 r -0.783 -0.042 -0.894 PR <0.001* 0.492 <0.001* 32 Bonner zoologische Beitráge 53 (2004) 30 24 Temperature [°C] O 20 40 60 30 100 120 140 160 180 200 220 Duration of provisioning trips [min] Fig. 5: Correlation (Pearson) between the duration of foraging trips and temperature. 3.6. Pollen - Pollen load In all pollen loads and cells only pollen of Hedera helix 2 hee (Fig. 6) was found. The mean longitudinal diameter of $ so 7 > a the pollen grains measured 38 + 1.3 um (n = 20), the 5 A A 604 equatorial diameter was 23 + 1.3 um (n = 20). 2 £ 40 zZ 20 - 04 SR NAS A o SA EP AS Particle size [um] 180 5 160 | Cell 3 140 4 © E 1204 oO a 1004 ° 901 oO É 60 + z 404 204 04 oc i AMM m, mana DV BS DOM MG DORSO RSS ONUDI H— 10 um —— Particle size [um] Fig. 6: SEM-Photograph of a pollen grain of Hedera helix. Fig. 7: Grain size distribution of one pollen load (only parti- cles between 20 and 30 um) and one cell (all particles) of C. hederae counted with an electronic particle counter in a vol- ume of 1100 pl. Inge BISCHOFF, Ester ECKELT, Michael KUHLMANN: On the Biology of the Ivy-Bee 33 Colletes hederae collected on average 518103 + 346639 (n = 15) grains per pollen load and the cells contained 2811816 + 1843170 (n = 20) grains. Thus, the females had to make approximately 4 or 5 (5.4) provisioning trips to complete one cell. The grain size distribution of one pollen load and cell is documented in Figure 7. 3.7. Nest density and architecture Most nests were found on sunny parts of the steep face, often covered by small tufts of grass. At the vertical part of the steep face more nests were located than at the base. Most females dug their nests above a level of | m. The mean number of nests was as high as 20-nests per m? in areas of the aggregation without vegetation cover, and up to 300 nests per m? in areas covered with grass. We excavated four nests and the architecture is shown in Figure 8. In all four nests the main burrow first ran between 7 and 12 cm horizontally into the steep face. The first cells occurred at a depth of 30-45 cm and be- yond a depth of 60 cm no cells were found. Between four and six cells were attached to one nest but pre- sumably the number is higher. All cells branched off the vertical part of the main burrow. Only one of the four nests exhibited side burrows. The most conspicuous character of the architecture was the successive ar- rangement of cells (up to 4 cells) without side burrows. Fig. 8: Nest architecture reconstructed from four excavated nests of C. hederae. The cell size measured on average 1-1.5 cm long with a diameter of 0.5 cm. The cells showed a convex shape at the side orientated to the main burrow and the mem- brane projected the cell at that end which optimised the successive arrangement of the cells. The fluid mixture ~ ~ A rd : of nectar and pollen filled approximately 2/3” of the cell. 4. DISCUSSION 4.1. Distribution The distribution pattern of C. hederae can be interpreted as the result of the postglacial recolonization of parts of Europe from an Italian refuge (DE LATTIN 1967, HEW- ITT 1999). Its restricted distribution compared with the host plant may indicate a relatively recent origin of this species. The latter view is supported by the weak morphological differentiation of C. hederae and its assumed closest relatives. Numerous Colletes species are oligolectic and collect pollen on the flowers of only one plant family or even a genus (MULLER & KUHLMANN 2003). Among them €. hederae, a member of the transpalaearctic C. succinctus species group (sensu NOSKIEWICZ 1936) that comprises 12 taxa, is unique because it is the only known bee spe- cies that is a specialised visitor of ivy (Hedera helix) (SCHMIDT & WESTRICH 1993). Species of the C. suc- cinctus group show a close morphological resemblance and are often hard to identify. The phylogeny of the species within this group is not yet clear but there is some evidence that C. succinctus and C. halophilus are the closest relatives of C. hederae. Despite their close relationship, the flower visiting behaviour of the three species differs significantly. As far as known this is a unique case at least among the Old World Colletes and of special interest with respect to the understanding of evolution of oligolecty and speciation within the genus (KUHLMANN 2003). Colletes halophilus is a narrow en- demic of costal sites of the southern North Sea and Eng- lish Channel and an oligolege of Asteraceae. Colletes succinctus is widespread in the western Palaearctic and known to be an oligolege of Calluna vulgaris and Erica spp. (Ericaceae) (WESTRICH 1989), but in some popula- tions of the British Isles exclusive pollen collecting on Asteraceae is documented (PERKINS 1945, EDWARDS & TELFER 2001). Phenology of all three species is similar, a single flight period in late summer / early autumn. 4.2. Pollen The oligolecty of C. hederae was confirmed in this study. All 15 pollen loads and 20 cells contained only pollen of Hedera helix. The amount of grains per pollen load can only be compared with other species of the same genus since the pollen collecting apparatus varies much between different genera (BRAUE 1916; FRIESE 1923; GRINFELD 1962; PASTEELS & PASTEELS 1979) and such studies are rare. The only Colletes species for which such data are available is C. cunicularius (Lin- naeus) (cf. BISCHOFF et al. 2003). Colletes cunicularius 34 Bonner zoologische Beiträge 53 (2004) collects on average three times as much grains per load as C. hederae, but the latter species is much smaller than C. cunicularius. Furthermore these authors did not consider the percentage of particles out of the range of pollen grains, which must be subtracted. In the present study this percentage of “pollution” amounted to 60% even in the pollen loads. Furthermore the number of col- lected grains strongly depends on the grain size (SILVEIRA 1991; BISCHOFF et al. 2003). The grains of Hedera helix display approximately the same elliptical shape but are nearly twice as large as Salix grains. Only with volumetric calculations a comparison of the carry- ing capacity can be achieved. Therefore quantitative analyses of related species (which have the same size) and a volumetric comparison should be done. 4.3. Nest site selection and nest architecture Like other Colletes species, C. hederae seems to prefer inclined and southward exposed nesting sites in sand or loess soils (cf. BISCHOFF 2000; MADER 1999). Likewise as with other congenerics C. hederae forms large aggre- gations. The nest architecture resembles that of C. cu- nicularius insofar as the short part of the horizontal main burrow is followed by a longer nearly completely vertical part and no real side burrows (K. FELTGEN pers. comm. Bonn 1997; MALYSHEV 1936). Nevertheless these authors did not mention a successive arrangement of several cells without side burrows. Only FRIESE (1923) figured a nest of C. cunicularius in which the cells were arranged successively. The successive ar- rangement of cells is also documented for C. daviesanus (Smith) (TISCHLER 1951; HAESELER 1972; MADER 1981; WESTRICH 1989). A comparison with the nest ar- chitecture of C. halophilus and C. succinctus, which remains to be discovered, would be interesting. 4.4. Diurnal and seasonal rhythm and reproduction rate Seasonally overlapping bee species living in the same habitat often differ in their diurnal foraging rhythm (LEVERMANN et al. 2000; BISCHOFF et al. 2003). Col- letes hederae is active in late summer and autumn when interspecific competition is reduced. This may have been one reason for a host switch to a late flowering host plant. The diurnal rhythm of C. hederae allows an optimal use of days with good flying conditions. One apparent character of the life cycle of C. hederae is the beginning of female provisioning behaviour shortly af- ter mating without a resting period for oocyte matura- tion as reported in other species (cf. GEBHARDT & ROHR 1987; MUNSTER-SWENDSEN 1968; BISCHOFF 2001). Colletes hederae made five provisioning trips on average per day but on days with optimal weather con- ditions we observed several females that made up to 10 trips and were active eight hours of the day. Colletes cunicularius, a vernal species, makes on average seven trips per day and shows also flying activity between 8-9 hours on days with good flying conditions (BISCHOFF et al. 2003). Comparable to vernal species, autumnal spe- cies have to deal often with long periods of unfavour- able weather conditions. Unlike the vernal C. cunicu- larius, C. hederae started its provisioning activity at a temperature threshold of 14.5 °C. This value is between the values of vernal and summer species and is also de- pendent of size (BATRA 1980; LIND 1968; MUNSTER- SWENDSEN 1968). Colletes hederae is much smaller than C. cunicularius, one of the largest bees in Germany with a mean heating rate of 7.35 °C per min (STONE & WILLMER 1989). Furthermore the activity is not only in- fluenced by temperature but also by light intensity. KLOSTERMEYER (1969) and GERBER (1969) reported that the flight activity of bees stopped under a limit of 15000 lux. We did not measure light intensity but the activity of males and females was reduced when it was cloudy. The first trip of the females was significantiy longer than the other trips of the day. This can be interpreted as additional adult feeding on nectar after a night in the nest. Another reason may be a temperature threshold since the duration of the flights correlated negatively with temperature. The results of the quantitative pollen analyses revealed on average five provisioning trips per cell. Thus C. hederae could provision one cell on an optimal day. A total of 18 bee days was estimated during which a fe- male could provision by rule of thumb 18 cells. It has to be considered, though, that the summer and autumn were extraordinary warm and dry in 2003. Thus the fe- males could not only use more days with optimal weather conditions for nest provisioning but also work faster because of high temperatures. Summarising the results, the specialisation of C. hederae on a late blooming host plant seems to mini- mise the competition for pollen resources as well as for nest site resources. The risk of parasitism may also be reduced. These advantages may compensate the disad- vantages of bad weather periods in autumn. It would be worthwhile to study also the diurnal foraging cycle and reproduction rate of the closely related species C. halo- philus and C. succinctus, who may follow different strategies. Acknowledgements. This work is dedicated to the late Prof. Dr. C. M. Naumann. We thank Prof. Dr. W. Bóhme (ZFMK) for supporting this study and Priv. Doz. Dr. G. Welp and Evi Sillmann (Institute for Soil Science, Univer- sity of Bonn) for the analysis of the soil samples. Further we wish to thank Axel Hirschfeld for help with statistical analyses. We are grateful to Dr. Rainer Hutterer and Dr. Bradley Sinclair who reviewed the manuscript. We are very much indebted to the following persons and institu- Inge BISCHOFF, Ester ECKELT, Michael KUHLMANN: On the Biology of the Ivy-Bee 35 tions that provided us with distribution records: Felix Amiet, Solothurn, Yves Gonseth (Centre Suisse de Carto- graphie de la Faune, CSCF, Neuchatel), Willem Hogenes (Zoölogisch Museum Amsterdam), Stuart Roberts and Mike Edwards (Bees, Wasps and Ants Recording Society, BWARS) and Hans-Richard Schwenninger (Stuttgart). REFERENCES AMIET, F., A. MÜLLER € NEUMEYER, R. (1999): Apidae 2. Colletes, Dufourea, Hylaeus, Nomia, Nomioides, Rho- phitoides, Rophites, Sphecodes, Systropha. Fauna Hel- vetica 4: 1-219. BATRA, S. W. T. (1980): Ecology, behavior, pheromones, parasites and management of the sympatric vernal bees Colletes inaequalis, C. thoracicus and C. validus. Journal of the Kansas Entomological Society 53: 509- 538. BELLMANN, H. (1995): Bienen, Wespen, Ameisen: die Hautfliigler Mitteleuropas. Kosmos Verlag, Stuttgart. BISCHOFF, I. (2000): Untersuchungen zur Nisthabitatwahl von Andrena vaga und Colletes cunicularius (Apidae) in der Wahner Heide (Rheinland). Beiträge Hymenop- terologen-Tagung Stuttgart 2000: 31-36. BISCHOFF, I. (2001): Populationsdynamik, Sammelstrategie und Nisthabitatwahl ausgewáhlter Wildbienen (Hyme- noptera, Apidae) in der Wahner Heide (Rheinland). Shaker, Aachen. BISCHOFF, I., FELTGEN, K. & BRECKNER, D. (2003): Forag- ing strategy and pollen preferences of Andrena vaga Panzer and Colletes cunicularius (L.) (Hymenoptera: Apidae). Journal of Hymenoptera Research 12: 220- 237. BRAUÉ, A. (1916): Die Pollensammelapparate der bein- sammelnden Bienen. Jenaische Zeitschrift fiir Natur- wissenschaft 50 (N.F. 43): 1-96. BUCHMANN, S. L. & SHIPMAN, C. W. (1990): Pollen har- vesting rate for Apis mellifera L. on Gossypium (Mal- vaceae) flowers. Journal of the Kansas Entomological Society 63: 92-100. Cross, I. C. (2002): Colletes hedearae Schmidt & We- strich (Hym., Apidae) new to mainland Britain with notes on its ecology in Dorset. Entomologist’s Monthly Magazine 138: 201-203. DE LATTIN, G. (1967): Grundriss der Zoogeographie. Fi- scher, Stuttgart. EDWARDS, R. & TELFER, M. (eds.) (2001): Provisional atlas of the aculeate Hymenoptera of Britain and Ireland. Part 3. Biological Records Centre, Huntingdon. FEITZ, F. (2001): Découverte de Colletes hederae Schmidt & Westrich, 1993 (Hymenoptera, Colletidae) au Lux- embourg. Bulletin de la Societe des Naturalistes Lu- xembourgeois 102: 87-90. FRIESE, H. (1923): Die europáischen Bienen (Apidae). — Das Leben und Wirken unserer Blumenwespen. Wal- ter de Gruyter, Berlin and Leipzig 1923. GEBHARDT, M. & RÖHR, G. (1987): Zur Bionomie der Sand- bienen Andrena clarkella (Kirby), A. cineraria (Lin- naeus), A. fuscipes (Kirby) und ihrer Kuckucksbienen (Hymenoptera: Apoidea). Drosera 11: 89-114. GERBER, H. S. (1969): Nesting behavior of Megachile ro- tundata (Hymenoptera, Megachilidae) monitored with an event recorder. Annals of the Entomological Soci- ety of America 62: 1321-1325. GOGALA, A. (1991): Contribution to the knowledge of the bee fauna of Slovenia (Hymenoptera: Apidae). Scopo- lia 25: 1-33. GOGALA, A. (1994): Contribution to the knowledge of the bee fauna of Slovenia Il. (Hymenoptera: Apidae). Scopolia 31: 1-40. GRINFELD, E. K. (1962): Origin and development of the apparatus for pollen collection in bees Hymenoptera: Apoidea). Entomological Review 41: 37-42. HAESELER, V. (1972): Anthropogene Biotope (Kahlschlag, Kiesgrube, Stadtgarten) als Refugien für Insekten, untersucht am Beispiel der Hymenoptera, Aculeata. Zoologische Jahrbúcher Systematik 99: 133-212. Hewitt, G. M. (1999): Post-glacial re-colonization of European biota. Biological Journal of the Linnean So- ciety 68: 87-112. KLOSTERMEYER, E. C. (1969): Using alkalibees and leaf- cutting bees to pollinate alfalfa. Washington Agricul- ture Experimental Station Circular 442: 1-8. KUHLMANN, M. (2003): Zur Kenntnis paláarktischer Bie- nen der Gattung Colletes Latr. mit Beschreibung neuer Arten (Hymenoptera: Apidae: Colletinae). Linzer biologische Beitráge 35: 723-746. LARSSON, F. K. (1991): Some take it cool, some like it hot — a comparative study of male mate searching tactics in two species of Hymenoptera (Colletidae and Sphe- cidae). Journal of Thermal Biology 16: 45-51. LEFEBER, B. A. (1998): Weer aculeatennieuws uit Zuid- Limburg (Hymenoptera: Apidae). Entomologische Berichten 58: 238-240. LEVERMANN, E.-M., BISCHOFF, I. 8 WAGNER, T. (2000): Species-specific foraging strategies of the syntopical and synchronous bees Panurgus calcaratus (Scopoli, 1763) and Dasypoda hirtipes (Fabricius, 1793). Beitrage zur Entomologie 50: 179-191. LIND, H. (1968): Nest provisioning cycle and daily routine of behaviour in Dasypoda plumipes (Hym. Apidae). Entomologiske Meddeleser 36: 343-372. MADER, D. (1981): Beitráge zur Substratgebundenheit von Nestbauten der solitáren Urbiene Colletes daviesanus (Hymenoptera: Colletidae) in quartáren Sedimenten im Buntsandstein in der Südeifel. Zoologische Jahrbücher Systematik 108: 165-177. MADER, D. (1999): Geologische und biologische Entomo- ökologie der rezenten Seidenbiene Colletes. Band 1. Logabook Köln. MALYSHEV, S. L (1936): The nesting habits of solitary bees. A comparative study. Revista española de ento- mología (EOS) 11: 201-309. MEUSEL, H. (ed.) (1978): Vergleichende Chorologie der zentraleuropäischen Flora. Vol. 2, Karten. Jena. MICHENER, C. D. (2000): The bees of the world. John Hop- kins University Press, Baltimore. MULLER, A. (1994): Die Bionomie der in leeren Schneck- enháusern nistenden Biene Osmia spinulosa (Kirby 1802) (Hymenoptera, Megachilidae). Veröffentlichun- gen fúr Naturschutz und Landschaftspflege in Baden- Württemberg 68/69: 91-334. MULLER, A., KREBS, A. & AMIET, F. (1997): Bienen. Mit- teleuropáische Gattungen, Lebensweise, Beobachtung. Naturbuch Verlag. MULLER, A. & KUHLMANN, M. (2003): Narrow flower spe- cialization in two European bee species of the genus Colletes (Hymenoptera: Apoidea: Colletidae). Euro- pean Journal of Entomology 100: 631-635. 36 Bonner zoologische Beitráge 53 (2004) MUNSTER-SWENDSEN, M. (1968): On the biology of the solitary bee Panurgus banksianus (Hymenoptera, Api- dae), including some ecological aspects. Yearbook Royal Veterinary Agricultural College Copenhagen 1968: 215-241. NOSKIEWICZ, J. (1936): Die paláarktischen Colletes-Arten. Prace Naukowe Wydawnictwo Towarzystwa Nau- kowego we Lwowie 3: 1-531. ; PASTEELS, J. J. & PASTEELS, J. M. (1979): Etude au micros- cope électronique a balayage des scopas collectrices de pollen chez les Andrenidae (Hymenoptera: Apoidea: Andrenidae). Archives de Biologie Paris 90: 113-130. PEETERS, T. M. J., RAEMAKERS, I. P. & SMIT, J. (1999): Voorlopige atlas van de Nederlandse bien (Apidae). EIS-Nederland, Leiden. PERKINS, R. C. L. (1945): The aculeate Hymenoptera of a small area of Dartmoor near Lydford, Devon. Ento- mologist’s Monthly Magazine 81: 145-153. Petit, J. (1996): Sur Colletes hederae Schmidt & Westrich 1993, abeille solitaire nouvelle pour la faune belge (Hymenoptera Apoide). Lambillionea 96 :55-58. RATHJEN, H. (1998): Colletes hederae Schmidt & Westrich — eine neue Solitárbienenart fúr Spanien. Bembix 11: 32-33. SCHMID-EGGER, C. (1997): Massenauftreten von Colletes hederae Schmidt & Westrich. Bembix 9: 16-17. SCHMID-EGGER, C., RISCH, S. & NIEHUIS, O. (1995): Die Wildbienen und Wespen in Rheinland-Pfalz. — Ver- breitung, Okologie und Gefáhrdungssituation. Fauna und Flora in Rheinland-Pfalz, Beiheft 16: 1-296. SCHMIDT, K. 8 WESTRICH, P. (1993): Colletes hederae n. sp., eme bisher unbekannte, auf Efeu spezialisierte Bienenart (Hymenoptera, Apoidea). Entomologische Zeitschrift 103: 89-112. SILVEIRA da, F. (1991): Influence of pollen grain volume on the estimation of the relative importance to its source to bees. Apidologie 22: 495-502. STONE, G. N. & WILLMER, P. G. (1989): Warm-up rates and body temperatures in bees: The importance of body size, thermal regime and phylogeny. Journal of Expe- rimental Biology 147: 303-328. TISCHENDORF, S. (1997): Ergánzungen zur Stechimmen- fauna von Hessen. Bembix 8: 16-17. TISCHLER, W. (1951): Ein biozónotischer Beitrag zur Be- siedlung von Steilwánden. Verhandlungen der Deut- schen Zoologischen Gesellschaft 1951: 214-229. WESTRICH, P. (1989): Die Wildbienen Baden-Wiirttem- bergs. Ulmer, Stuttgart. WESTRICH, P. & DATHE, H. H. (1997): Die Bienenarten Deutschlands (Hymenoptera, Apidae) - Ein aktualisiertes Verzeichnis mit kritischen Anmerkungen. Mittei- lungen Entomologischer Verein Stuttgart 32: 3-34. WIERING, H. (1999): Hoe zeldzaam is Colletes hederae? Bzzz, Nieuwsbrief Sectie Hymenoptera van de Neder- landse Entomologische Vereniging No. 10: 27-30. Authors” addresses: Inge BISCHOFF (corresponding author), Esther ECKELT, Alexander Koenig Research Institute and Museum of Zoology, Adenauerallee 160, D-53113 Bonn, Germany, e-mail: i.bischoff.zfmk@uni- bonn.de; Michael KUHLMANN, Institute of Landscape Ecology, University of Münster, Robert-Koch-Str. 26, D-48149 Münster, Germany, e-mail: kuhlmmi@uni- muenster.de Bonner zoologische Beiträge Band 53 (2004) Heft 1/2 Seiten 37-80 Platycnemididae)' Dirk GASSMANN Institute of Biology, Leiden University / National Museum of Natural History, Leiden Abstract. Phylogenetic relationships of Southeast Asian and Indo-Pacific damselflies of the subfamily Calicnemiinae (Odonata: Platycnemididae) are examined by cladistic analyses using morphological characters. The strict consensus cladogram of the resulting equally most parsimonious trees supports the monophyly of the Papuan genus /diocnemis Se- lys, the Philippine genus Risiocnemis Cowley and its subgenera, but leaves the basal relationships of the African genera and the Palawan genus Asthenocnemis Lieftinck partly unresolved. A preferred phylogenetic hypothesis is presented showing a well supported 'Indo-Pacific clade' consisting of Philippine, New Guinean and Solomon island taxa, and as sister group Asthenocnemis. Risiocnemis turns out to be a sister group of Lieftinckia/Salomocnemis (Solomon Islands), the sister taxon of those being the central New Guinean Arrhenocnemis Lieftinck. Together, these form a monophyletic group with the remaining Papuan taxa. /diocnemis leonorae Lieftinck is transferred to Rhyacocnemis Lieftinck comb. nov. The possible effects of taxon sampling are discussed. Key words. Taxonomy, biogeography, morphology, Southeast Asia, phylogeny, cladistic analysis, Rhyacocnemis leon- Bonn, Juni 2005 The Phylogeny of Southeast Asian and Indo-Pacific Calicnemiinae (Odonata, I orae comb. nov. 1. INTRODUCTION The odonate subfamily Calicnemiinae Fraser, 1957 repre- sents small to medium-sized damselflies (Zygoptera) in- habiting small running waters throughout the old world. The group is almost exclusively tropical. Due to the con- finement to stable habitats and lacking dispersal capaci- ties in the rainforest environment, the majority of species shows a high degree of endemism, both at species and genus level. Therefore, the group is well suited for his- torical biogeographical analyses (GASSMANN in prep.). SELYS (1863) distinguished seven subgenera in his “le- gion Platycnemis’. Together, those subgenera, which were later given generic rank by the same author (SELYS 1886) represented a large part of the Afrotropical, Southeast Asian and Indo-Pacific Calicnemiinae known today. TILLYARD (1917) raised SELYS’ legions to sub- family level resulting among others in a subfamily Platycnemidinae (he used the incorrect name ‘Platy- cneminae’). However, the family-group name had al- ready been introduced earlier (YAKOBSON & BIANKI 1905). TILLYARD & FRASER (1938/40) finally erected the family Platycnemididae. In the most recent account on the group, MARTENS (1996) provided five diagnostic characters (at least with regard to Coenagrionidae) for the family: the comparatively long and obtuse discoidal cell (quadrilateral), the length of the pterostigma (matching no more than one wing cell), the variable po- sition of arculus (at or distal to Ax2), the course of the longitudinal veins MA and IR3 (mainly straight, only 1 In commemoration of Clas Michael Naumann zu Königsbrück (26.06.1939 — 15.02.2004) apically zigzagged), and size (small to medium-sized). Another character of Platycnemididae, long ciliae on the legs, was already mentioned by Selys (1886) for his le- gion Platycnemis; however, this trait is shared by other zygopteran families (SCHMIDT 1951b). The Platycnemididae now consist of two subfamilies, Platycnemidinae Yakobson & Bianki, 1905, and Calic- nemiinae Fraser, 1957. The subfamily Platycnemidinae FRASER (1957) was the first to define the subfamily in its present form, containing the two genera Platvcnemis Burmeister, 1839, and Copera Kirby, 1890. The dilation of the male tibiae, which is the most distinct feature Fraser used to characterize the subfamily, has turned out to be not diagnostic for the group. Several species cur- rently assigned to Platycnemidinae, for example, some Madagascan ones (SCHMIDT 1951a) have undilated tib- iae. Contrarily, one species of Calicnemiinae, Risiocne- mis atropurpurea (Brauer), does have dilated tibiae. Within the genus Platycnemis, 31 species are currently recognized (derived from MARTENS 1996 and HAMALAINEN 2003). The group exhibits a disjunct pat- tern of distribution, involving the western Palaearctic region, eastern Asia, northwestern/ western Africa as well as the Madagascan region (SCHMIDT 195la: MAR- TENS 1996). No representatives were known from Southeast Asia until recently. However, the recent dis- covery of a new species from Laos (P. phasmovolans Hämäläinen) extends the distribution of the group far into the Indo-Chinese region (HAMALAINEN 2003). 38 Bonner zoologische Beitráge 53 (2004) Within the genus Copera (Copera spec., Plate 1b) cur- rently nine species are recognized (MARTENS 1996). The group is largely replacing Platvcnemis in Southeast Asia, but there is an area of overlap in eastern Asia (MARTENS 1996). The generic borders between Copera and Platycnemis are weakly defined, and their identity _recently has been put in question by HAMALAINEN (2003). The subfamily Calicnemiinae The damselfly subfamily Calicnemiinae was erected by FRASER (1957) to accomodate species of platycnemidid damselflies which did not fit into the subfamily Platy- cnemidinae. Fraser himself stated: “This subfamily is formed mainly for convenience to separate the true Platycnemiines, and one has only to compare the genera composing it, to note how artificial it appears to be...”. Unfortunately, Fraser did not provide diagnostic charac- ters for the subfamily, except for the absence of dilated tibiae, which on the other hand is a striking feature in the second subfamily Platycnemidinae. BECHLY (1996) found no autapomorphies for the group. Later authors (MARTENS 1996) have provided additional distinguish- ing characters for Calicnemiinae, i.e. the comparatively acute discoidal cell (quadrilateral) and the length of the anal vein. The Calicnemiinae reach their highest diversity in Southeast Asia and the Indo-Pacific region. Remarka- bly, the group is absent from Wallacea (Sulawesi, Smaller Sunda Islands, Moluccas), which has been con- firmed, at least for Sulawesi, by several odonatological surveys during the last decades (VAN TOL 2000). The African genera of Calicnemiinae The African genera of the subfamily are a rather hetero- genous assemblage comprising only about a dozen spe- cies in total (SCHORR et al. 2004). The family status of some of them, in particular that of Mesocnemis Karsch, 1891 (Western Africa, 4 species) and Metacnemis Selys, 1863 (Southern Africa, Madagascar; 3 species), has been doubted (MARTENS 1996; WATERSTON 1984). Ex- cept for Allocnemis Selys, 1863 (Southern Africa, 2 species), all remaining African genera are monotypic: Leptocnemis Selys, 1886 and Paracnemis Martin, 1903 (Madagascar), Oreocnemis Pinhey, 1971 (eastern Af- rica) and Arabicnemis Waterston, 1984 (Arabian Penin- sula). Allocnemis and Arabicnemis also extend into sub- tropical areas. The Oriental genera of Calicnemiinae From the Oriental region, four calicnemiine genera have been recognized. Calicnemia Strand, 1928 is a medium- sized genus with 17 described species (SCHORR et al. 2004) which reaches its highest diversity in the Himala- yan region (LIEFTINCK 1984); however, its distributional range further extends southeast into the Indo-Chinese mainland as well as to Taiwan and Hainan. Interest- ingly, no representatives of Calicnemia are known from the Greater Sunda Islands. Indocnemis Laidlaw, 1917, is a monotypic genus; /. orang (Förster in Laidlaw, 1907) occurs in India and mainland Southeast Asia. Coeliccia Kirby, 1890 is by far the largest genus of Calicnemii- nae, currently comprising 59 species according to SCHORR et al. (2004). Its distribution area ranges from the Himalayan region via the Southeast Asian mainland into the Greater Sunda Islands and even into the western and southwestern Philippines; south to the Philippines, along the Greater Sunda Islands, the eastern border of Coeliccia matches the Wallace line, except for the is- land of Bali from where no representative of the genus is known. A taxonomic and phylogenetic revision of a presumably monophyletic subgroup of the genus, con- sisting of Sundaland and Philippine taxa, is in work (DIIKSTRA & GASSMANN, in prep.). The fourth Oriental genus, Sinocnemis, comprises two species and was es- tablished quite recently (WILSON & WEN-BAO 2000); however, it could not be studied in detail for the present analysis. The Indo-Pacific Calicnemiinae Recently, a large part of the New Guinean and Philip- pine taxa have been revised taxonomically (GASSMANN 1999, 2000; GASSMANN & HAMALAINEN 2002; HAMALAINEN 1991a/b, 2000). Based on recently col- lected specimens from different sources, several new species were recognized. The New Guinean genus Idiocnemis Selys now amounts to 19 species and thus represents the largest genus in the Papuan region, fol- lowed by Lieftinckia Kimmins (Solomon Islands; 6 spe- cies), Rhyacocnemis Lieftinck (New Guinea; 3 species, see below), Paramecocnemis Lieftinck (New Guinea, 2 species). The remaining Papuan genera Cyanocnemis Lieftinck, Lochmaeocnemis Lieftinck, Thaumatagrion Lieftinck, Torrenticnemis Lieftinck (New Guinea) and Salomocnemis Lieftinck (Solomon Islands) are all monotypic (for Arrhenocnemis see below). The Philip- pine genus Risiocnemis Cowley now comprises 36 de- scribed species, with subgenus /gneocnemis Hämäläinen (20 species) being somewhat larger than the nominal subgenus Risiocnemis Cowley (GASSMANN & HAMALAINEN 2002). Within the Philippine Calicnemii- nae, the Palawan genus Asthenocnemis Lieftinck is an outstanding representative. Originally, Lieftinck thought that the peculiar female specimen from the Martin col- lection of which he based the description of the new species, A. stephanodera, would originate from New Guinea (LIEFTINCK 1949). Later, with the arrival of ad- ditional (male) specimens from Palawan Island, he stated his error (LIEFTINCK 1971, 1974). Dirk GASSMANN: The Phylogeny of Southeast Asian and Indo-Pacific Calicnemiinae 39 Plate 1: a) Coeliccia membranipes, 3, Java; Photo: J. van Tol. b) Copera spec., $, Sarawak, Borneo; Photo: D. Paulson. c) Idiocnemis australis, $, Lake Kutubu, Papua New Guinea; Photo: J. Michalski. d) /diocnemis inaequidens, ©, Morobe Province, Papua New Guinea; Photo: D. Gassmann. e) Risiocnemis kaiseri, $, Samar, Philippines; Photo: R. A. Müller. f) Risiocnemis ser- rata, 6, Luzon, Philippines; Photo: D. Paulson. 40 Bonner zoologische Beitráge 53 (2004) Preexisting hypotheses on phylogenetic relationships within Calicnemiinae Several authors have occasionally discussed the pre- sumed relationships between different genera of Calic- nemiinae (FRASER 1932; LIEFTINCK 1958, 1963). Since these considerations were not yet influenced by the methodological tools provided by W. HENNIG'S phy- logenetic systematics (1950, 1966), they were deter- mined to remain rather vague assumptions which were not necessarily founded on synapomorphic traits (see discussion). Also FRASER's ‘reclassification of Odonata’ (1957) is not based on cladistic concepts, and up to now no phylogenetic study on platycnemidid damselflies has been available. However, BECHLY (1996) and recently REHN (2003) performed morphological-phylogenetic higher-level analyses of Odonata, including some se- lected representatives of the superfamily Coenagrion- oidea, respectively. 2. MATERIALS AND METHODS 2.1. Specimens examined The present study is based on preceding revisional work on the taxonomy of Papuan and Philippine Calicnemii- nae by GASSMANN (1999, 2000) and GASSMANN & HAMALAINEN (2002). The reader is referred to those references for detailed material lists. However, in the meantime, many additional taxa, among them the entire subgenus Risiocnemis, have been studied for a more in- clusive analysis of phylogenetic relationships between the Indo-Pacific representatives of the subfamily; those specimens are listed in Appendix 4. Despite the fact that they are not yet formally described, the male of Ri- siocnemis moroensis as well as the females of R. gracilis and Idiocnemis leonorae have been included in the analysis. 2.2. Taxon sampling Relationships between the Indo-Pacific taxa were exam- ined down to species level; all the known species were included (Appendix 1). However, for the Oriental gen- era, in particular the huge genus Coeliccia Kirby, 1890, this was not possible due to practical constraints. Thus, for both the Oriental and the African genera, only single representatives were included in the analysis. Thau- matagrion funereum from New Guinea was examined, but finally not included in the analysis, because its cur- rent status within Platycnemididae should be seriously doubted (see results and discussion). On the other hand, the author did not see any reason not to include the two known Arrhenocnemis species; their habitus and wing venational characters - especially the distinctly crenulate wing margin - strongly resembles the Calicnemiinae (al- though Lieftinck considered them to belong to the Megapodagrionidae (LIEFTINCK 1965, 1971; see discus- sion)). Due to the lack (or limited availability) of mate- rial, the African genera Metacnemis Selys, 1863 and Oreocnemis Pinhey, 1971 as well as the comparatively recently described Oriental genus Sinocnemis Wilson and Wen-bao, 2000 had to be excluded from the analy- sis. It should be noted that currently also phylogenetic revisions of Calicnemia Strand, 1928 (GASSMANN, in prep.) and the Philippine and Bornean representatives of Asthenocnemis and Coeliccia (DIJKSTRA & GASSMANN, in prep.) are in work, eventually providing a better un- derstanding of the West Malesian calicnemiine fauna. 2.3. Outgroup choice The second subfamily of Platycnemididae and presumed sister group of Calicnemiinae, the Platycnemidinae, were chosen as outgroup for the analysis. Unfortunately, it was not possible to include more taxa than one repre- sentative for each of the two genera Platycnemis and Copera;, however, a literature survey revealed that the majority of species within Platycnemidinae are at least structurally very similar (e.g. SCHMIDT 195 1a). 2.4. Scanning electron microscopy (SEM) The male ligulae of almost all species and the male ap- pendages and female prothoracices of selected species were examined by scanning electron microscopy (SEM). The scanning electron micrographs were taken by the author, mainly using a JEOL SEM 6400 microscope. The objects were cleaned in 70% ethanol, air-dried and sub- sequently sputtered with gold for 1.5 - 2.0 min at 20 mA before examination. Figures 5-6, 9-10, 17-18, 19-20, 23- 24 and 35-36 were made using a JEOL SEM 840A micro- scope. 2.5. Terminology Although the modified Tillyard-Fraser system of odo- nate wing venation (cf. WATSON & O'FARRELL 1991) is controversial (loc. cit.), it is further applied here, be- cause it has already been used in preceding revisional work on Calicnemiinae (GASSMANN 1999, 2000; GASSMANN & HAMALAINEN 2002). 2.6. Measurements The measurements of wings and abdomens were taken within a precision of 0.5 mm. Measurements of the ad- ditional specimens examined for the present paper are not given here but can be obtained from the author. In a few cases, measurements had to be taken from the lit- erature. 2.7. Character coding In several cases, coding of character states for the cladistic analysis was problematic, because an unam- bigious (i.e. objective) delimitation of discrete states was not possible. However, the attempt was made to di- vide, for instance, a gradually changing character into a Dirk GASSMANN: The Phylogeny of Southeast Asian and Indo-Pacific Calicnemiinae +] ylogeny Ax1 Ax2 Sn Pt Ac / postquadrilateral IR antenodal cells 3 IR2 l cm Px N Arc | een Postquadrilateral b (discoidal cell) aa analiveis EA : | Fig. 1: Left forewing of Calicnemia eximia, 4 (a) and Platycnemis pennipes, © (b). Abbreviations: Ac = antenodal crossvein: Arc = arculus; Ax = antenodal vein; N = nodus; Px = postnodal vein; Pt = pterostigma; (I)R = (intercalary) radius; Sn = subnodal vein. Fig. 2: Wing apex of Risiocnemis (Igneocnemis) tendipes, Ö, (a), Risiocnemis (Risiocnemis) laguna, E, (b). 42 Bonner zoologische Beitráge 53 (2004) series of discrete intervals which were chosen as small and precise as possible (cf. character 56). For the treat- ment of continuous morphometrical traits: cf. characters 17 and 55. Polymorphisms were entered as separate character states (scaled coding; different methods reviewed in WIENS 1999 and KORNET & TURNER 1999), but left unordered. However, in a few cases in which character states were not unambigiously codable (e.g. colour traits of deterio- rated specimens), the states concerned were entered as polymorphies while applying the ‘uncertainty’ option in PAUP (SWOFFORD & BEGLE 1993) to minimize the po- tential effect of erroneously delimitated character states on the analysis. 2.8. Cladistic analysis Morphological characters of imagines were compiled in a data matrix which consisted of 88 binary and multi- state characters (listed below); due to our scarce knowl- edge of larval forms and the probably doubtful assign- ment of larval to adult specimens in the literature, larval characters were not considered for the present analysis. Since the female sex 1s unknown in several species, only three female characters were used to keep the amount of missing values in the data matrix low. In cases of char- acters exhibiting more than nine states, the consecutive states were labelled with Roman capital letters in the data matrix (Appendix 2); however, in the character list below, all states are numbered numerically for better readability. Four uninformative characters were ex- cluded from the analysis. A parsimony analysis using PAUP* 4.0b4a (SWOFFORD 2002) was applied to exam- ine relationships of 84 taxa in total. The resulting topol- ogy was rooted a posteriori by designating 2 taxa (see above) as outgroup. The heuristic search settings were chosen as follows: all characters unordered and equally weighted, starting tree(s) obtained via stepwise addition, random addition sequence with 100 replicates; branch- swapping algorithm: tree-bisection-reconnection (TBR). Character optimization was performed by accelerated transformation (ACCTRAN); character evolution was traced and examined by applying MacClade 3.04 (MAD- DISON & MADDISON, 1993): see Appendix 3. Analyses testing clade stability were performed using Autodecay 4.0 (ERIKSSON 1998) and PAUP* 4.0b4a (SWOFFORD 2002), the latter run under the same settings as above, but with only 10 replicates.The trees were rooted by ap- plying the paraphyletic rooting option to avoid assign- ment of many ambiguous character states to the ingroup node while using only two outgroup taxa. 2.9. List of characters Male characters: Wings 1. Apical wing margin: shape. (0) not crenulate (Fig. la/b); (1) smoothly sinuous (Fig. 2a); (2) distinctly crenulate (Fig. 2b). All Papuan representatives of Calicnemiinae (but see be- low for Thaumatagrion funereum) as well as the species of subgenus Risiocnemis have a distinctly crenulate apical wing margin: from the level of the pterostigma up to about the distal third of posterior margin. The wing apex is furnished with concave inlets between the vein endings (Fig. 2b). Contrarily, the wing apex of /gneocnemis spe- cies is smoothly sinuous ('smoothly crenulate' in GASS- MANN 1999, 2000 and GASSMANN & HAMALAINEN 2002): the posterior half of the apical wing margin is di- vided into shallowly deepened or, occasionally, straight areas, divided by more or less distinct emarginations at the levels of veins R>, R3, Ry and - more proximal and less distinct - at CuP (Fig. 2a). This feature is largely shared by Leptocnemis cyanops from the Seychelles; however, it is slightly less distinct in that species and lacking the emargination at Ro. 2. Wing petiolation. (0) between Axl and Ax2 (Fig. la/b); (1) proximal to Axl. In most species examined, the basal petiolation of the wings reaches up to a level just distal to the first an- tenodal crossvein (Axl). Only in Arabicnemis caerulea and Mesocnemis singularis, the level of petiolation ends proximally to the level of Axl. 3. Position of anal crossvein (Ac). (0) distinctly distal to Ab (Fig. la/b); (1) at or slightly distal/proximal to Ab (about halfway between Axl and Ax2); (2) somewhat proximal to Ab (closer to Ax] than to Ax2); (3) far proximal to Ab. The position of the anal crossvein (Ac) relative to the insertion of the anal bridge (Ab) can be readily classi- fied in distinct character states; however, there is some variation in those taxa showing condition (0): in Mesocnemis singularis and Paracnemis alluaudi, Ac is situated about 2.5 to 3.5 times its own length distal to Ab, whilst in other taxa as in Cyanocnemis aureofrons, the distance amounts to just about the length of Ac. Lieftinckia species have Ac situated far proximal to Ab, i. e. about halfway between wing base and Ax2. Dirk GASSMANN: The Phylogeny of Southeast Asian and Indo-Pacific Calicnemiinae 43 4. Anal vein: length. (0) short, reaching no further than up to about the level of Px6 (Fig. 1b); (1) long, reaching further than the level of Px6 (Fig. la). The condition (0) usually means that R3 inserts at the level where the anal vein ends: a character originally used to dicriminate between the two subfamilies of Platycnemididae (MARTENS 1996; see also Fig. 1). However, the relation between these two veins cannot be applied to taxa where R3 and IR2 appear to be moved much more distally, as in Risiocnemis and some species of Lieftinckia. Therefore, in the present study the length of the anal vein is put in relation to the corre- sponding number of Px. 5. Arculus: position. (0) proximal to Ax2; (1) at or slightly proximal to Ax2; (2) at Ax2 (Fig.la); (3) at or distal to Ax2; (4) distal to Ax2. The position of the arculus relative to the second an- tenodal vein (Ax2) has often been used as a taxonomic character in platyenemidid classification; however, the degree of variation differs between different genera. 6. Medio-anal link. (0) straight or nearly straight (Fig.1b); (1) straight or slightly hooked; (2) slightly hooked (Fig. la); (3) dis- tinctly hooked. The shape of the medio-anal link corresponds with the shape of the quadrilateral: the posterior distal angle of the latter can be either nearly right, or very acute; con- sequently, the medio-anal link appears either straight or broken. 7. Third postquadrangular cell (forewing). (0) absent (Fig. 1b); (1) absent or present; (2) present (Fig. la). Both in forewing and hindwing, the majority of the ex- amined species has two cells between quadrilateral and the vein descending from the subnodal vein. However, in some taxa the number of postquadrilateral cells can amount to three or four cells. 8. Fourth postquadrangular cell (forewing). (0) absent; (1) absent or present; (2) present. 9. Third postquadrangular cell (hindwing). (0) absent; (1) absent or present; (2) present. 10. Fourth postquadrangular cell (hindwing). (0) absent; (1) present. 11. Position of R4 (forewing). (0) distinctly proximal to Sn (Fig. la/b); (1) slightly proximal to Sn; (2) at Sn; (3) at or distal to Sn; (4) distal to Sn. f 12. Position of IR3 (forewing). (0) at Sn (Fig. la/b); (1) slightly distal to Sn; (2) be- tween Px1 and Px2; (3) at about the level of Px2; (4) be- tween Px3 and Px7. 13. R4 and IR3: fused/separate. (0) separate; (1) fused at base. In Paramecocnemis, the R4 and IR3 veins are fused from shortly distal to wing base up to about the level of Px1 to Px2. In all remaining taxa, the two veins are dis- tinctly separate. 14. Pterostigma (forewing): length. (0) matching one cell or less (Fig. 1b); (1) reaching well beyond one cell (Fig. la). The length of the pterostigma was examined in relation to the posteriorly neighbouring cell. 15. Pterostigma (forewing): colour. (0) brown; (1) white. 16. Wing colour. (0) entirely clear; (1) with a slight brownish breath; (2) brownish at base; (3) entirely brownish. 17. Hind wing length. (0) short (mean value = 17.0); (1) medium-sized (mean values = 19.6-29.4); (2) long (mean value = 30.7); (3) very long (mean value = 32.2). To transform the measurement values into discrete states, gap-coding as described by ARCHIE (1985) was applied. The pooled within-group standard deviation (sp) was calculated, and the group means (of species) were arranged by size; subsequently, every sub-row of means which leaves a gap greater than s, (= 1.0) to neigbouring mean values, is assigned an integer score, respectively. Arrhenocnemis sinuatipennis was not con- sidered, because only one measurement value (from the literature) was available. Head 18. Labrum: ground colour. (0) black; (1) dark brown: (2) reddish-brown: (3) yel- low-orange; (4) yellow or yellowish-white; (5) bright red; (6) blue; (7) bluish-grey; (8) greenish-blue. 44 Bonner zoologische Beitráge 53 (2004) 19. Anteclypeus: ground colour. (0) black; (1) dark brown to black; (2) dark brown; (3) medium to dark brown; (4) medium brown; (5) light to medium brown; (6) light brown; (7) reddish-brown; (8) reddish-brown to black; (9) orange-brown; (10) yellow- orange; (11) yellowish or yellowish-white; (12) blue; (13) bluish-grey; (14) green. 20. Postclypeus: ground colour. (0) black; (1) black, basal area orange; (2) dark brown to black; (3) dark brown. (4) medium to dark brown; (5) medium brown; (6) red- dish-brown; (7) orange-brown; (8) orange; (9) blue; (10) bluish-grey; (11) green. 21. Genae: colour. (0) black; (1) dark brown; (2) reddish-brown; (3) light to medium brown; (4) yellow-orange; (5) greenish- yellow; (6) greenish-blue; (7) bluish-grey; (8) blue. 22. Postclypeus: shape (in lateral view). (0) bulgy, edge rounded; (1) slightly curved, edge bulgy; (2) slightly curved, edge semi-sharp; (3) angu- late, edge sharp. 23. Frontal stripe. (0) present; (1) absent. The anterior part of the head may show several kinds of distinct or diffuse markings. A comparatively distinct feature 1s the frontal stripe which, 1f present, runs from eye to eye (for an exception see below), covering the frons and the parts lateral to it. 24. Frontal stripe: extension. (0) reaching to a level just anterior to antennae, not in- cluding antennal sockets; (1) up to and including anten- nal sockets; (2) up to and including antennal scapus. In Asthenocnemis stephanodera, as an exception, the frontal stripe 1s not reaching the eye margin, respec- tively, but instead is only slightly exceeding the width of postclypeus. 25. Frontal stripe: shape. (0) more or less complete, not interrupted; (1) inter- rupted in the center of frons. 26. Frontal stripe: colour. (0) reddish-brown; (1) blue; (2) greenish-blue; (3) greenish-yellow; (4) bluish-grey to violet; (5) orange; (6) pale orange; (7) chrome. 27. Brown stripes lateral to postclypeus. (0) present, reaching from postclypeus to eye margin; (1) short, rudimentary, reaching only halfway the dis- tance from postclypeus to eye margin; (2) absent. The character state is coded as unknown for those taxa in which the lower anterior part of the head is black throughout, so that no discrete stripes are recognizable. 28. Vertical stripe at the level of ocellar area. (0) present; (1) reduced to a pair of transverse elongate spots; (2) reduced to a pair of longitudinal elongate spots; (3) reduced to diffuse stripe; (4) absent. 29. Vertex: ground colour. (0) black; (1) black to reddish-brown; (2) reddish-brown to bright red; (3) bluish-grey. 30. Antennal scapus: ground colour. (0) black; (1) dark brown; (2) reddish-brown; (3) light brown; (4) orange; (5) bright red; (6) yellow-white; (7) bluish-grey; (8) blue. 31. Antennal pedicellus: ground colour. (0) black; (1) dark brown; (2) reddish-brown; (3) light brown; (4) orange; (5) bright red; (6) yellow-white; (7) bluish-grey; (8) blue. 32. Length ratio antennal pedicellus/scapus. (0) subequal in length; (1) about 1.25 times the length; (2) about 1.5 times the length; (3) about twice the length; (4) about 2.5 times the length; (5) about 3 times the length. 33. Subquadrangular marking around ocelli. (0) absent; (1) present. Except for 7. nigriventris, the vertex of species of the Idiocnemis bidentata species-group 1s marked by a sub- quadrangular black marking (cf. GASSMANN 2000, fig. 31): 34. Postocellar spots on vertex. (0) absent; (1) small, circular, framed; (2) large, subtri- angular; (3) very large, subtriangular. 35. Distinct elongate spots on occiput. (0) present; (1) absent. 36. Brown spots lateral to ocelli. (0) absent; (1) present. Thorax Dirk GASSMANN: The Phylogeny of Southeast Asian and Indo-Pacific Calicnemiinae 45 37. (Male) pronotal tubercles: size. (0) flat or only slightly raised; (1) distinct; (2) very dis- tinct. Males of most taxa studied have the right and left half of the median lobe either not raised or only slightly rounded in lateral view. However, they can be raised to distinct protrusions, reaching their extreme in As- thenocnemis stephanodera. 38. Prothorax: ground colour. (0) black or brown-black; (1) brown-black; (2) light, reddish- or medium brown. j 39. Legs: coloration. (0) Black throughout or mainly black; (1) mainly black or dark brown; (2) light to medium brown, darkened to a varying extent; (3) yellow-light brown with black markings; (4) yellow-brown or -black; (5) red-black; (6) bright orange; (7) orange-red; (8) black-bluish(-white). 40. Legs: shape of tibiae. (0) distinctly dilated; (1) not distinctly dilated. 41. Synthorax: robust mesonotal processes. (0) absent; (1) present. Arrhenocnemis amphidactylis shows a conspicuous, finger-like outgrowth at either side of dorsal carina of anterior part of synthorax. 42. Synthorax: black line/marking on dorsal carina. (0) narrow; (1) broad; (2) dorsal surface of synthorax black or brown-black throughout (except for ante- humeral stripes, if present). The extension of black coloration on synthorax is either restricted to the dorsal carina, or it fills the entire upper surface of synthorax (occasionally reaching beyond the humeral suture). In /diocnemis dagnyae, as an excep- tion, the black carinal stripe varies in width from exactly matching the median carina up to reaching the width of the carinal forks. 43. Antehumeral stripe. (0) complete stripe; (1) distinct, interrupted; (2) half stripe; (3) short anterior stripe, joining humeral suture (4) distinct, reduced to anterior spot, joining carinal fork; (5) rudimentary, interrupted; (6) diffuse stripe; (7) absent. 44. Metepisternal stripe. (0) absent; (1) rudimentary; (2) present. 45. Metepisternal stripe: colour. (0) blue; (1) turquoise; (2) diffuse light brown; (3) grey- ish-purple; (4) orange; (5) bluish-grey; (6) green, inter- mingled with orange; (7) yellowish. 46. Distinct metepimeral marking. / (0) not clearly defined; (1) short, subquadrangular. This feature is included here to characterize some spe- cies of subgenus Risiocnemis of which the metepimera are furnished with distinct subquadrangular colour markings (see HAMALAINEN 1991a). In the remaining taxa, the metepimeron is either entirely or largely uni- form in coloration. 47. Meso- and metathoracic markings. (0) absent; (1) present. In some species of the inornata species-group of Idiocnemis, the mes- and metepisternal fossae at poste- rior ends of first and second lateral sutures are covered by an elongate black spot, respectively. This character was examined and scored only for Idiocnemis species. 48. Pruinescence (0) absent; (1) parts of head, pro- and synthorax and legs pruinescent; (2) pruinescent throughout. 49. Underside of synthorax: bristles. (0) absent or only minute; (1) a well-delimited patch of robust spines present. Abdomen 50. Abdomen: basic colour pattern. (0) rather uniform (often with more or less distinct an- nules), but lacking a clear separation in two different colours; (1) abdomen distinctly separated in anterior (brown-)black and posterior reddish part. 51. Posterior dorsal abdominal marking: colour. (0) blue to violet/purple; (1) yellow-orange or yellow- white; (2) blue-whitish-pruinescent. A variably shaped, mostly subquadrangular colour marking covers the dorsal surface of last abdominal segments in several taxa studied. 52. Dorsal marking on segment 9. (0) absent; (1) present. 53. Dorsal marking on segment 10. (0) absent; (1) present. 54. Terminal abdominal segments: shape. (0) not or only slightly inflated; (1) distinctly inflated. Bonner zoologische Beitráge 53 (2004) Figs. 3-6: Scale bars Ligulae of 44 in ventral (3, 5) and lateral view (4, 6): 3-4, Asthenocnemis stephanodera; 5-6, Calicnemia 100 um eximia. Dirk GASSMANN: The Phylogeny of Southeast Asian and Indo-Pacific Calicnemiinae 47 Figs. 7-10: Ligulae of 94 in ventral (7, 9) and lateral view (8, 10): 7-8, Coeliccia membranipes: 9-10, Copera marginipes. Scale bars 100 um. 48 Bonner zoologische Beitráge 53 (2004) N N N | NES 12 a 14 Figs. 11-14: Ligulae of 44 in ventral (11, 13) and lateral view (12, 14): 11-12, Cyanocnemis aureofrons; 13-14, Idiocnemis pol- hemi (from alt. 700 m). Scale bars 100 um. Dirk GASSMANN: The Phylogeny of Southeast Asian and Indo-Pacific Calicnemiinae 49 16 Figs. 15-18: Ligulae of 94 in ventral (15, 17) and lateral view (16, 18): 15-16, Idiocnemis zebrina sufficiens; 17-18, Indocnemis orang. Scale bars 100 um. 50 Bonner zoologische Beitráge 53 (2004) ASS 22 20 Figs. 19-22: Ligulae of 44 in ventral (19, 21) and lateral view (20, 22): 19-20, Leptocnemis cyanops; 21-22, Lochmaeocnemis malacodora. Scale bars 100 um. Dirk GASSMANN: The Phylogeny of Southeast Asian and Indo-Pacific Calicnemiinae 5 Figs. 23-26: Ligulae of $4 in ventral (23, 2 erythrostigma. Scale bars 100 um. 26 25) and lateral view (24, 26): 23-24, Mesocnemis singularis; 25-26, Paramecocnemis Nn 159) Bonner zoologische Beitráge 53 (2004) 28 WO — 30 Figs. 27-30: Ligulae of 44 in ventral (27, 29) and lateral view (28, 30): 27-28, Rhyacocnemis leonorae; 29-30, Risiocnemis at- ropurpurea. Scale bars 100 um. Dirk GASSMANN: The Phylogeny of Southeast Asian and Indo-Pacific Calicnemiinae 53 Figs. 31-34: Ligulae of 44 in ventral (31, 33) and lateral view (32, 34): 31-32, Risiocnemis appendiculata; 33-34, Risiocnemis gracilis. Scale bars 100 um. 54 Bonner zoologische Beitráge 53 (2004) 36 e 38 Figs. 35-38: Ligulae of 44 in ventral (35, 37) and lateral view (36, 38): 35-36, Stenocnemis pachystigma; 37-38, Thaumatagrion funereum. Scale bars 100 um. Dirk GASSMANN: The Phylogeny of Southeast Asian and Indo-Pacific Calicnemiinae 33 Figs. 39-40: Ligulae of ‘¢ in ventral (39) and lateral view (40): 39-40, Torrenticnemis filicornis, Scale bars 100 um. In the taxa concerned, the caudal end of abdomen is in- flating dorsoventrally and/or laterally from segment 7 or segment 8 onwards. 55. Length ratio abdomen/hindwing. (iia 1.55.(2).1:65 (3) 1.734) 1:85) 1.936) 2-1. The length ratio of wing length and abdomen length is considerably constant within species; in cases where some variation occured, mean values were calculated. Anal appendages 56. Length ratio inferior/superior anal appendages. (0) about 3.0 times: (1) about 1.5 times: (2) 0.9 to 1.0 times; (3) 0.7 to 0.8 times; (4) 0.5 to 0.6 times: (5) 0.3 to 0.4 times; (6) 0.1 to 0.2 times. For measuring the length of both inferior and superior appendages, curvation of appendages was taken into ac- count as far as possible. 56 Bonner zoologische Beitráge 53 (2004) 57. Length ratio superior appendages/abdominal seg- ment 10. (0) about half the length; (1) subequal; (2) about 1.3 times the length; (3) about 1.5 times the length; (4) about 1.75 times the length; (5) twice the length; (6) about 3 times the length. To avoid measurement errors, the character states were only scored for those species with straight superior ap- pendages. 58. Subbasal process. (0) absent; (1) present. In most of the taxa studied, there is a process situated more or less close to the base of superior appendage (Figs. 41-42, 44-45); it varies considerably in shape and length among the species (see below). 59. Subbasal process: shape. (0) elongate, slender; (1) blunt, subtriangular; (2) acute, short; (3) hook-like, short; (4) long, flattened; (5) broad, flattened; (6) robust, rounded (Figs. 44-45); (7) protrud- ing into a large extension; (8) short, slender; (9) notched squarish process; (10) broadly rounded. 60. Subbasal process: length. (0) long; (1) of medium length; (2) short. This character and the next were scored for Risiocnemis (Igneocnemis) species only because of both their good recognizability and discriminatory value in those spe- cies, 61. Subbasal process: position (in lateral view). (0) Partly covered by posterior margin of abdominal segment 10 (in lateral view); (1) just distal to posterior margin; (2) moved further distally. 62. Inner median process. (0) absent; (1) present. The superior appendage of four species of the /diocne- mis bidentata species-group is furnished with an inner median process which is situated between subbasal and subdistal process; in /diocnemis bidentata, the median process is almost fused with the subdistal one (cf. GASSMANN 2000: figs. 14-17). 63. Inner subdistal process. (0) absent (Figs. 44-45); (1) small hook; (2) small tuber- cle; (3) finger-shaped hook; (4) slender hook; (5) blunt medium-sized hook; (6) tubercular protuberance, di- rected inwards; (7) subtriangular, hooked (Figs. 41-42); (8) large bulge. 64. Subdistal process: base. (0) not prolonged; (1) moderately prolonged; (2) exten- sively prolonged. In some (mainly Lieftinckia) species, the base of the subdistal process (hook) is considerably prolonged. Alternatively, the resulting elongate protuberance could also be interpreted and thus homologized as a dis- tally moved subbasal process. However, its specific quality and position renders this alternative less plausi- ble. 65. Superior appendage: second minute spine proximal to subdistal process. (0) absent; (1) present. 66. Superior appendage: additional spines. (0) absent; (1) present. With an additional inner dorsal spine and a second inner baso-ventral spine, the superior appendage cf Loch- maeocnemis malacodora is of far more complex built than that of other Calicnemiinae. 67. Superior appendage: curvature in lateral view. (0) curved somewhat dorsad; (1) more or less straight; (2) only apically curved downwards; (3) sickle-shaped; (4) halfway incurved to right angle. 68. Superior appendage: curvature in dorsal view. (0) apically incurved, tips converging; (1) sickle- shaped; (2) moderately incurved; (3) slightly incurved; (4) more or less straight. 69. Superior appendage: shape inner side (in dorsal view). (0) halfway kinked outwards; (1) base bulgy, otherwise convex; (2) bulged out; (3) more or less straight; (4) only apically convex; (5) no such modifications. 70. Superior appendage: shape outer side (in dorsal view). (0) slightly concave, kinked; (1) slightly convex; (2) bulged out; (3) more or less straight; (4) straight, distal part bent outwards; (5) no such modifications. 71. Superior appendage: shape of apex. (0) rounded; (1) broadly rounded, somewhat flattened (Figs. 41-43; 44-46); (2) sharp; (3) complex, with inner plate; (4) notched; (5) subquadrangular. 72. Inferior appendage: shape in lateral view. (0) straight, or slightly curved up; (1) strongly curved dorsad; (2) nearly plate-like, straight. Dirk GASSMANN: The Phylogeny of Southeast Asian and Indo-Pacific Calicnemiinae 5 73. Inferior appendage: shape of apex. (0) distinctly kinked inwards (Figs. 41-43); (1) slightly bent inwards; (2) not kinked inwards (Figs. 44-46). In Coeliccia membranipes and Indocnemis orang, the apex of inferior appendage is distinctly kinked inwards as well as slightly bent downwards. Calicnemia eximia and Allocnemis leucosticta show the same feature, but less distinct. 74. Superior appendages: colour. (0) black; (1) mainly black, with some yellow areas; (2) reddish; (3) reddish-brown; (4) yellow-brown, (5) yel- low-orange; (6) yellow-white; (7) blue. Ligula 75. Ligula: lateral hooks (cf. GASSMANN 2000). (0) absent; (1) present (Fig. 12). Cyanocnemis aureofrons and the species of the Idiocnemis bidentata - group exhibit more or less dis- tinct lateral emarginations at the ligula head. 76. Ligula: distal median cleft (cf. GASSMANN 1999). (0) absent; (1) present; (2) only slightly developed. 77. Ligula: flagella at ligula head. (0) present (Figs. 5-6, 7-8, 17-18, 23-24, 33-34); (1) ab- sent. 78. Ligula: terminal lobes. (0) long, broad, apically subrectangular (Figs. 13-14); (1) of median length and width (Figs. 33-34, 39-40); (2) reduced (Figs. 11-12); (3) short, spatulate (Figs.15-16; Figs. 31-32); (4) long, broad, apically rounded; (5) long, very broad (Figs. 29-30); (6) very long, semi-acute (Figs. 35-36); (7) long, broad, apically hooked (Figs.19- 20); (8) broad, of median length, subrectangular (Figs. 25-26, 27-28); (9) leaf-shaped, of median length (Figs. 21-22); (10) very broad, of median length; (11) very long, slender. 79. Ligula head: degree of folding. (0) not folded; (1) slightly folded; (2) moderately folded; (3) strongly folded. 80. Ligula head: width. (0) broad (Figs. 7-8, 15-16, 31-32, 33-34); (1) elongate, narrow (Figs. 13-14); (2) stout, as wide as long (Figs. 29-30); (3) elongate, subtriangular, widening distally. 81. Ligula: medio-lateral protrusion. (0) absent; (1) present (Figs. 8, 28, 40). 82. Ligula head: inner protrusion. (0) absent; (1) present (Figs. 4, 22, 28). 83. Ligula: apical ridge/plate. (0) absent; (1) intermediate ridge (Figs. 15-16); (2) ridge (Figs. 33-34); (3) plate (Figs. 7-8). 84. Ligula: inner lamina/flagellum. (0) lamina; (1) intermediate between lamina and flagel- lum (Fig. 16); (2) flagellum (Figs. 22, 40); (3) reduced to short process. 85. Ligula: incision between terminal lobes: shape. (0) rounded (Figs. 5-6, 11-12, 13-14, 15-16, 17-18, 19- 20, 21-22, 25-26, 27-28, 31-32, 33-34, 39-40); (1) arc- shaped; (2) slightly arc-shaped (Figs. 29-30); (3) sharp: (4) very sharp (Figs. 7-8, 23-24); (5) straight. Female characters: 86. Female pronotal tubercles. (0) distinct, high (Fig. 50); (1) distinct, of moderate height; (2) only sightly raised (Fig. 48), or bulgy. The female pronotal tubercles can be flat up to slightly raised (Fig. 48), or extended into high conical processes (Fig. 50) which are mostly pointed apically. 87. Female posterior pronotal lobe. (0) trilobate; (1) undivided. All species of genus Risiocnemis have the posterior pronotal lobe divided into a broad median lobe and a pair of narrow lateral lobes (Fig. 49). However, the character state 'trilobate' also circumscribes a condition where either the median lobe is completely lost or at least distinctly reduced. 88. Female posterior pronotal lobe: elevation. 5 (0) directed anteriorly; (1) strongly elevated; (2) dis- tinctly elevated (Fig. 50); (3) halfway elevated; (4) not or only slightly elevated (Fig. 48). In case of species with trilobate posterior pronotal lobe, only the elevation of the median part (lobe) was deter- mined. 58 Bonner zoologische Beitráge 53 (2004) Figs. 41-43: Anal appendages of Coeliccia membranipes, Í, in lateral (41), posterior (42) and dorsal (43) view. Figs. 44-46: Anal appendages of Risiocnemis elegans, Ö, in lateral (44), posterior lateral (45) and dorsal (46) view. Scale bars 300 um (44, 45), 100 um (41-43, 46). Dirk GASSMANN: The Phylogeny of Southeast Asian and Indo-Pacific Calicnemiinae 59 Figs. 47-50: Prothorax of Idiocnemis mertoni, Y (47, 48) and Risiocnemis elegans, Y (49, 50) in dorsal (47, 49) and lateral (48, 50) view. Scale bars 300 um (47, 49), 100 um (48, 50). 3. RESULTS 3.1. Monophyly of Platyenemididae and Calicnemiinae Within the scope of the present study, it was not possi- ble to thoroughly test the hypothesis of monophyly for the family Platyenemididae and/or the subfamily Calic- nemiinae; several families of Coenagrionoidea and even other superfamilies would have to be included in such an analysis. However, the morphological studies re- vealed putative synapomorphies which apparently have not been mentioned by other authors, and which could, however, be considered in future studies of higher-level phylogeny. For the Platycnemididae these are the only moderately kinked arculus (which is not nearly right- angled, as in at least some coenagrionides), and the comparatively long legs (femora of hind legs distinctly reaching beyond posterior margin of synthorax). In the present analysis, the Calicnemiinae turn out to be mo- nophyletic with regard to the Platycnemidinae (Copera and Platycnemis); however, all apomorphic character states at the ingroup node experience one or more sub- 60 Bonner zoologische Beitráge 53 (2004) sequent reversals or transformations in the ingroup; thus, there is no synapomorphy uniting all calicnemiines based on the present data set. 3.2. Phylogeny The parsimony analysis resulted in 732 equally parsi- monious cladograms (721 steps; consistency index: 0.406, retention index: 0.717, rescaled c.i.: 0.292), of which the strict consensus is shown in Figure 51. The strict consensus cladogram is largely resolved at the species level for the New Guinean /diocnemis and the Philippine Risiocnemis. The monophyly of both genera as well as that of the smaller Papuan genera Ar- rhenocnemis and Paramecocnemis ıs confirmed. Rhyacocnemis forms a monophyletic group together with Idiocnemis leonorae which therefore is transferred to that genus (see below). The three representative spe- cies of the Oriental genera Calicnemia, Coeliccia (C. membranipes, Plate la) and /ndocnemis also cluster to- gether in the strict consensus. At the base of the clado- gram, we find an unresolved assemblage consisting of all Afrotropical genera examined. The ingroup node is well supported (d=3). We find Arrhenocnemis (New Guinea) in a reasonably well supported but unresolved clade (d=2) together with the genera Risiocnemis (Phil- ippines), and single species of Lieftinckia and Salo- mocnemis (Solomon Islands); at a higher level, this clade is part of a polytomy comprising Asthenocnemis, the smaller Papuan genera and the /diocnemis clade. At the next higher level, the African taxa join into a poly- tomy also involving the Coeliccia-/Indocnemis- /Calicnemia - clade. Idiocnemis Selys The relationships among the /diocnemis species turn out to be nearly fully resolved in the strict consensus clado- gram. The /. inornata species-group (GASSMANN 1999) is monophyletic, leaving the /. bidentata - group (GASSMANN 2000) paraphyletic. The latter one shows a largely pectinate topology except for the two sister spe- cies /. inaequidens Lieftinck (Plate Id) and /. pruines- cens. Not surprisingly, /. dagnvae represents the “link” between the two species-groups; this species combines traits of both species-groups. The /. inornata - group is well supported (d=3); its subclades only partly reflect the preliminary subgroupings recognized by GASSMANN (1999): leaving aside group I which only consists of one single species, /. inornata, we find that group I (north- ern New Guinea) is distributed across two clades which are, however, not sister clades. Instead, most members of group III (eastern Papuan archipelagos) are united with the western representatives of the northern New Guinea species in one clade, while the northeastern New Guinea species, together with a single southern species, I. australis Gassmann (Plate Ic), are the sister group to the remaining clade. Risiocnemis Cowley The Philippine genus Risiocnemis as well as both of its subgenera turn out to be monophyletic. While subgenus Risiocnemis is reasonably well supported (d=2), the basal node of subgenus /gneocnemis is even more strongly supported with the highest decay index (d=5) of all clades. Within subgenus Risiocnemis, the mainly northern Philippine R. serrata - species-group (Plate If: R. serrata) is well supported (d=3). The sister group re- lationship of R. rolandmuelleri and R. seidenschwarzi, distributed in the West Visayans, is confirmed and thus the monophyly of the R. rolandmuelleri - species-group, although only weakly supported (d=1). Independent from the still uncertain position of R. erythrura and R. praeusta, the predominantly southernly distributed R. appendiculata - group can be considered clearly para- phyletic, with R. kiautai representing the sister taxon of the R. serrata - group, and R. confusa being the sister taxon of the R. rolandmuelleri - group. R. arator, the only representative of the fourth species-group defined by HAMALAINEN (1991a), 1s the most basal taxon of the group. In subgenus /gneocnemis, R. atripes is the most basal species, followed by R. rubricercus; both species are from Mindanao. Proceeding further on, we find two dis- tinct clades which are each, however, weakly supported (d=1). The first one comprises only species from the eastern Philippines (e.g. R. kaiseri, Plate le), except for R. fuligifrons which is distributed across northwestern Mindanao (Zamboanga Peninsula). The second one mainly contains a mixed assemblage of species from Mindanao and Luzon, but also from the West and East Visayan islands. There is little hierarchical structure and only weak support within both these clades, except for the well supported sister group relationships of R. cal- ceata/R. siniae, and R. flammea/R. odobeni. 3.3. Preferred phylogenetic hypothesis A survey of all equally parsimonious cladograms re- vealed that there are two different positions As- thenocnemis takes: either basal to a clade containing all Papuan representatives, or just in between the Papuan taxa. Here, the latter alternative is considered less fa- vourable, because it would mean that reversals in at least four important structural characters would have taken place in Asthenocnemis: from a denticulate wing margin back to a smooth one, the reversal from an inner flagellum to a lamina as well as the increase in postquadrilateral cells in both fore- and hindwing. If we filter the (resolved) trees showing Asthenocnemis in a basal position out of the entire set of trees, we obtain 70 trees which only differ in minor details of inner-generic relationships. To reconstruct character evolution, one tree out of this subset was chosen as the preferred phy- logenetic hypothesis (Fig. 52). According to this clado- Dirk GASSMANN: The Phylogeny of Southeast Asian and Indo-Pacific Calicnemiinae 6] 1 A. (Risiocnemis) appendiculata R. (Risiocnemis) moroensis 1 R. (Risiocnemis) gracilis 1 R. (Risiocnemis) laguna 3 R. (Risiocnemis) pulchra R. (Risiocnemis) serrata R. (Risiocnemis) elegans 3 R. (Risiocnemis) varians RA. (Risiocnemis) asahinai R. (Risiocnemis) kiautai 2 R. (Risiocnemis) erythrura R. (Risiocnemis) praeusta 1 R. (Risiocnemis) confusa 1 R. (Risiocnemis) seidenschwarzi 2 R. (Risiocnemis) rolandmuelleri R. (Risiocnemis) arator R. (Igneocnemis) atripes 5 R. (Igneocnemis) rubricercus R. (Igneocnemis) atropurpurea 1 R. (Igneocnemis) antoniae ] 1 R.(/gneocnemis) fuligifrons 1 1 1 R. (Igneocnemis) melanops 1 R. (Igneocnemis) kaiseri 1 1 R. (Igneocnemis) haematopus R. (Igneocnemis) polliloensis R. (Igneocnemis) nigra 3 R. (Igneocnemis) calceata 1 R. (Igneocnemis) siniae 2 1 R. (Igneocnemis) pistor 1 R. (Igneocnemis) tendipes 1 R. (Igneocnemis) rubripes 1 R. (Igneocnemis) plebeja 1 R. (Igneocnemis) incisa 1 R. (Igneocnemis) ignea 2 R. (Igneocnemis) flammea R. (Igneocnemis) odobeni 2 Arrhenocnemis amphidactylis Arrhenocnemis sinuatipennis Salomocnemis gerdae Lieftinckia isabellae Lieftinckia kimminsi Lieftinckia lairdi Lieftinckia malaitae Lieftinckia ramosa Lieftinckia salomonis 1 Idiocnemis inornata 3 Idiocnemis fissidens 1 Idiocnemis strumidens 1 Idiocnemis chloropleura 1 Idiocnemis leonardi 1 3} L2 Idiocnemis louisiadensis Idiocnemis zebrina 1 Idiocnemis australis 3 1 Idiocnemis adelbertensis 2 Idiocnemis huonensis 2 Idiocnemis kimminsi 1 Idiocnemis dagnyae 1 Idiocnemis nigriventris Idiocnemis obliterata 1 Idiocnemis mertoni 1 Idiocnemis bidentata Idiocnemis polhemi 1 Idiocnemis inaequidens Idiocnemis pruinescens 3 Paramecocnemis erythrostigma 3 Paramecocnemis stilla-cruoris Cyanocnemis aureofrons Torrenticnemis filicornis 2 Rhyacocnemis leonorae comb. nov. 3—— Ahyacocnemis prothoracica Rhyacocnemis sufficiens Lochmaeocnemis malacodora Asthenocnemis stephanodera 2 Calicnemia eximia 4 Coeliccia membranipes Indocnemis orang Paracnemis alluaudi Mesocnemis singularis Stenocnemis pachystigma Arabicnemis caerulea Leptocnemis cyanops Allocnemis leucosticta Platycnemis pennipes Copera marginipes Fig. 51: Strict consensus cladogram derived from all 732 equally parsimonious trees. Decay indices (Bremer support values) are indicated on the branches. 62 Bonner zoologische Beitráge 53 (2004) ———— 4 105 od 101 107 6 —.. 106 7. 104 109 du el 100: 103 18- 125) Rn 19- 118 «o 1 . (Risiocnemis) appendiculata .(Risiocnemis) moroensis .(Risiocnemis) asahinai .(Risiocnemis) elegans .(Risiocnemis) varians . (Risiocnemis) serrata (Risiocnemis) gracilis .(Risiocnemis) laguna . (Risiocnemis) pulchra .(Risiocnemis) kiautai .(Risiocnemis) erythrura . (Risiocnemis) praeusta .(Risiocnemis) confusa . (Risiocnemis) rolandmuelleri . (Risiocnemis) seidenschwarzi > > >» DDD DO DDD DDD» D .(Risiocnemis) arator . (Igneocnemis) antoniae . (Igneocnemis) fuligifrons . (Igneocnemis) melanops . (Igneocnemis) kaiseri . (lgneocnemis) haematopus . (Igneocnemis) polliloensis . (Igneocnemis) nigra . (Igneocnemis) atropurpurea .(Igneocnemis) calceata . (Igneocnemis) siniae . (Igneocnemis) flammea . (Igneocnemis) odobeni . (Igneocnemis) ignea . (Igneocnemis) incisa . (Igneocnemis) plebej a . (Igneocnemis) rubripes . (Igneocnemis) tendipes . (Igneocnemis) pistor . Ugneocnemis) rubricercus > DDD DD D DDD DD DD DD DD DD D . (Igneocnemis) atripes Salomocnemis gerdae Lieftinckia lairdi Lieftinckia malaitae Lieftinckia salomonis Lieftinckia ramosa Lieftinckia isabellae Lieftinckia kimminsi Arrhenocnemis amphidactylis Arrhenocnemis sinuatipennis Dirk GASSMANN: The Phylogeny of Southeast Asian and Indo-Pacific Calienemiinae 63 Idiocnemis adel bertensis Idiocnemis huonensis Idiocnemis kimminsi Idiocnemis australis Idiocnemis chloropleura / Idiocnemis fissidens Idiocnemis strumidens Idiocnemis inornata Idiocnemis leonar dí Idiocnemis louisiadensis Idiocnemis zebrina Idiocnemis dagnyae Idiocnemis nigriventris Idiocnemis obliterata Idiocnemis mertoni Idiocnemis bidentata Idiocnemis polhemi Idiocnemis inaequi dens Idiocnemis pruinescens Paramecocnemis erythrostigma Paramecocnemis stilla-cruoris Rhyacocnemis leonorae n. comb. Rhyacocnemis prothoracica Rhyacocnemis sufficiens Torrenticnemis filicornis Cyanocnemis aureofrons Lochmaeocnemis malacodora Asthenocnemis stephanodera Paracnemis alluaudi Coeliccia membranipes Indocnemis orang Calicnemia eximia Leptocnemis cyanops Un 4 fav m >> < om Mesocnemis singularis > on a Stenocnemis pachystigma Allocnemis leucosticta Arabicnemis caerulea Platycnemis penni pes VUM N SS Nu NP a? ás > 2 > wo Copera marginipes Fig. 52: Preferred tree out of 70 equally parsimonious trees showing Asthenocnemis in comparatively basal position. Branch numbers refer to the character changes listed in Appendix 3. Paraphyly of the outgroup taxa is due to the rooting method. Abbre- viations: AFR = Africa, ARAB = Arabian Peninsula; EUR = Europe; MAD = Madagascar; NG = New Guinea; PAL = Palawan: PHIL = Philippines; SEA = (Mainland) Southeast Asia, Sundaland; SEY = Seychelles; SOL = Solomon Islands. gram, a sister group relationship between the Philippine Risiocnemis and the Solomon-clade (Lieftinckia- Salomocnemis) is proposed, based on two synapomor- phies: the absence of an antehumeral stripe and the shape of the terminal abdominal segments which are not inflated. This clade shares two synapomorphies with its sister group, Arrhenocnemis: the reduction of postquad- rilateral cells in forewing from three to just two, as well as the reduction of a dorsal marking on segment 9; fur- thermore, two characters change along branch 96: the colour of superior appendages (from black to yellow- brown; strongly homoplasious character), and the shape of the terminal lobes of the ligula (from the reduced state to short, spatulate lobes). At the next higher level, that clade described above is united with all Papuan taxa in a monophyletic group, resulting in an ‘Indo-Pacific clade'; the latter comprises all Papuan and Philippine taxa and is characterized by the reduction of the third postquadrangular cell in the hindwing (synapomorphy of all representatives) as well as the absence of a trans- verse stripe on vertex (with a reduction to an only dif- fuse stripe in Arrhenocnemis). The most remarkable 64 Bonner zoologische Beitráge 53 (2004) apomorphic trait of the 'Indo-Pacific clade', however, is the distinctly crenulate wing margin; however, it is re- duced to a smoothly sinuous condition in /gneocnemis; thus, the distinctly crenulate wing margin turns out to be plesiomorphic in subgenus Risiocnemis. The New Gui- nean clade (excluding Arrhenocnemis) is characterized by an basal apomorphic change (branch 140) from an inner lamina to an inner flagellum of the ligula, how- ever, it cannot be considered an autapomorphy of the whole clade because two independent subsequent rever- sals into an intermediate state take place, as well as one into a rudimentary short process. Five further character changes occur at the base of the New Guinea clade, but all of them experience subsequent reversals. The large Indo-Pacific (sub)genera and their group- defining characters are as follows: /gneocnemis species firstly are characterized by the smoothly sinuous wing apex. Other synapomorphies of /gneocnemis species are the subequal length of antennal pedicellus and scapus, the pointed (sharp) apex of the superior appendage and the stout ligula head. Thus, in total, /gneocnemis species are united by four synapomorphies. Subgenus Risiocnemis 1s characterized by three synapomorphies, 1.e. the distal po- sition of the arculus, a slight brownish breath on wings as well as the trilobate female posterior pronotal lobe. A pair of brown spots lateral to the ocelli is an autapomorphy of the most recent common ancestor of sg. Risiocnemis and is maintained in all species except for R. appendiculata. Compared to Risiocnemis and its subgenera, the mono- phyly of /diocnemis is weakly supported (see above). However, the genus is characterized by the following au- tapomorphies ( = synapomorphies of its species): colora- tion of legs (light to medium brown, darkened to a vary- ing extent) and the shape of the terminal abdominal segments (not or only slightly inflated). 3.4. Consequences for classification Idiocnemis leonorae Lieftinck is transferred to Rhyacocnemis Lieftinck: Rhyacocnemis leonorae (Lieftinck, 1956) comb. nov. The genus now comprises three species, characterized by four synapomorphies, most of them referring to the peculiar shape of the male appendages: superior ap- pendage apically incurved, with tips converging; infe- rior appendages rudimentary (0.1 to 0.2 times the length of superiors); apex of superior appendage sharp (homo- plasious character). A distinct apomorphic trait of the ligula in Rhyacocnemis is the inner protrusion of the ligula head (Fig. 28); however, the condition in R. pro- thoracica 1s not known. Since the position of Salomocnemis within one clade to- gether with Lieftinckia (Fig. 52) is not stable (cf. con- senus cladogram: Fig. 51), synonymization of the for- mer genus with the latter is not considered here. 4. DISCUSSION 4.1. General The present study should be considered a first approach to the phylogeny of Calicnemiinae. However, some conclusions can be drawn with relative certainty, 1. e. the confirmed monophyly of the larger Indo-Pacific genera, especially Risiocnemis; there can also be no doubt about the monophyly of each of its subgenera. Unfortunately, basal relationships within Calicnemiinae are not unambigiously resolved, a fact which might be due to taxon sampling in the Oriental and African taxa, as well as to the fact that genera of doubtful family status (e.g. Mesocnemis) have been included; these fac- tors potentially introduce homoplasy. Generally it holds that a large number of taxa keeps the consistency index low (KITCHING et al., 1998); with a taxa/character ratio of 84/88, the comparatively low actual c. 1. ( = 0.406) was to be expected. However, it was the intention of the author to include as many representatives of Calicnemi- inae as possible to thoroughly cover the (morphological) diversity of the group. The monophyly of the subfamily Calicnemiinae in its present form remains doubtful; no unique synapomorphy for all species currently included has been found in the analysis with regard to Platycne- midinae; however, REHN (2003) found one synapomor- phic trait supporting his Coeliccia-Indocnemis/Ri- siocnemis - Clade: the absence of a large, proximal hornlike sclerite on the posterior articular plate. Unfor- tunately, this character could no longer considered (and examined) here after REHN's paper had come out. Al- though REHN's taxon sampling is rather scarce, certainly structural traits of the wing articulation appear to be im- portant characters to continue with. Possibly, there are indeed comparatively distinct mor- phological characters which allow for the assumption of monophyly at least for the family Platycnemididae: i.e. the characteristic course of the MA and IR3 veins. However, these characters are somewhat more difficult to examine in taxa with comparatively short wings. In REHN's study (2003) Platycnemididae is rendered para- phyletic, and the old world Protoneuridae form a mono- phyletic group together with the subfamily Platycnemid- inae. Since REHN included only very few platycnemidid taxa, the significance of his findings has to remain open to further study. So, there have been indications for both the monophyly of Platycnemididae and Calicnemiinae, but also against it; apparently, only the monophyly of the subfamily Platycnemidinae has never been doubted. A detailed phylogenetic analysis of at least Coenagrion- oidea, if possible including Lestoidea (Megapodagrioni- dae) is urgently required. Although the present study was not designed to clarify the family status of dubiously assigned taxa, some notes might be appropriate. The genus Arrhenocnemis was in- Dirk GASSMANN: The Phylogeny of Southeast Asian and Indo-Pacific Calicnemiinae 65 cluded here because it shares the distinctly crenulate wing apex with all other Papuan Calicnemiinae (except Thaumatagrion, see below) and, in terms of wing vena- tion, mediates between Lieftinckia and Risiocnemis. It 1s not clear to the author why LIEFTINCK (1971; short note in type catalogue) placed this genus in the Megapo- dagrionidae, although he originally described it as a “platycnemine” (LIEFTINCK 1933). In 1965, he wrote in his account on the Odonata of Madagascar, referring to the genus Arrhenocnemis: “... Like Tatocnemis, this was originally placed in Platycnemididae, but it is quite evident that both should be removed therefrom.” An- other doubtful case is the New Guinean Thaumatagrion: possibly Lieftinck had already something different in mind when he named the genus by using the suffix -agrion instead of -cnemis. In any case, he placed it within the Platycnemididae (LIEFTINCK 1932), a deci- sion which is strongly questioned here. The general wing shape and colour, the size, but most of all, the largely zigzagged course of the MA and IR3 veins and the distinctly kinked arculus make the author consider it to be a potential coenagrionide; the 'very faintly denticu- late margin' of the wing apex (LIEFTINCK 1932) differs considerably from that of the other Papuan taxa. Beside that, the ligula structure 1s different from all other known calicnemiine taxa: the presence of spines at- tached to the ligula shaft (Figs. 37-38) suggests a closer relationship with the Papuan coenagrionide genus Oreagrion. BECHLY (1996) considered the absence of those spines to be an autapomorphy of the family Platy- cnemididae. And, finally, no internal lamina or filament seems to be present (Fig. 38). Based on these considera- tions, Thaumatagrion was excluded from the present analysis. Since REHN's study, also the status of Al- locnemis within the Platycnemididae is in question. In his analysis, this African genus turned out to be a sister group of a clade which contained the Calicnemiinae on one side and a clade consisting of Platycnemidinae and old world Protoneuridae on the other. As REHN points out himself, the sister group relationship between A/- locnemis and Platycnemididae/Protoneuridae is based only on a single homoplasious character, the extension of the premental cleft (REHN 2003). In the present analysis, Allocnemis, clusters quite 'logically' between the African platycnemidides. 4.2. Relationships between calicnemiine genera With regard to previous ideas on phylogenetic relation- ships within Calicnemiinae, both confirming and sur- prising results arose from the analysis. SCHMIDT sug- gested a close relationship between some Papuan genera (Thaumatagrion, Paramecocnemis) and the African Mesocnemis and Metacnemis (SCHMIDT 1951); these hypotheses are not at all corroborated by the present re- sults. However, FRASER's suggestion of a close relation- ship between the Oriental genera Calicnemia, Coeliccia and Indocnemis has been confirmed (FRASER 1932) While LIEFTINCK (1963), in the light of the present analysis, was very right to consider Lieftinckia and Ri- siocnemis Closely allied, he was apparently not with re- gard to Idiocnemis and Coeliccia. Some of these con- siderations were founded on larval morphology however, LIEFTINCK himself stated that those findings are not always reliable because of the difficulties of as- signing larval to adult stages in the field (LIEFTINCK 1984). Remarkably, the Madagascan genera Leptocnemis and Paracnemis are found at slightly displaced positions within the cladogram (Fig. 52): instead of splitting off from the first, Paracnemis alluaudi branches off from the Oriental taxa. It should be noted that for Paracne- mis, a total of 13 character states are either unknown to the author or - as those of the uniquely shaped male ap- pendages - inapplicable and therefore coded as missing values in the data matrix. Thus, it is possible that the position of Paracnemis in the cladogram represents an artefact; this will be taken into account for bio- geographic conclusions (VAN TOL & GASSMANN, in prep.). The ambigious position of Asthenocnemis stephanodera in the set of equally parsimonious trees (see above) can possibly be explained by taxon sampling. Preliminary results from work on the Coeliccia species of the Greater Sunda Islands and the Philippines have revealed that Asthenocnemis probably just represents an 'offshot' of one of two monophyletic radiations within the genus Coeliccia, one of them partly leading to the colonization of geological terranes which today constitute the west- ern and southern Philippines. In this context, 4. stephanodera apparently 1s the most 'apomorphic' spe- cies (DIJKSTRA & GASSMANN, in prep.). Thus, it is very likely that insufficient taxon sampling is the cause for the unresolved position of 4. stephanodera in the result- ing set of cladograms since the mediating morphologi- cal traits are lacking. Consequently, further phyloge- netic analyses of the Southeast Asian Calicnemiinae should include the highly interesting West Malesian ra- diations of Coeliccia. This would also contribute to the ongoing work on the historical biogeography of the group (GASSMANN, in prep.; VAN TOL & GASSMANN, submitted). 4.3. Perspectives for further studies of calicnemiine phylogeny The present study has resulted in a first estimate of the phylogeny of the damselfly subfamily Calicnemiinae. It is apparent that basal relationships are still unsatisfac- torily resolved (Fig. 51), and that the addition of taxa from the Oriental complex of genera (and the Palawan genus Asthenocnemis should be included in this context) is required. Firstly, this would allow for testing the un- 66 Bonner zoologische Beitráge 53 (2004) certain monophyly of genera as Calicnemia and Coelic- cia, which partly have been defined on the base of vari- able (polymorphic) wing venational traits of which the group-defining properties very much depend on our definition of the character states concerned. Secondly, it is necessary to newly define the generic borders be- tween Coeliccia and Asthenocnemis. Several new as Hainan Mindoro A] BS - 1 Moluccas Sulawesi Sumatra Zn Java Bali EEE Distribution range of Calicnemiinae well as described species should be transferred to the latter genus; distinct putative synapomorphies have been found (DIJKSTRA & GASSMANN, in prep.). Thirdly, by increasing the range of taxa and of both morphological (e.g. endosceletal) and molecular characters, the am- bigious phylogenetic position of Asthenocnemis could possibly be resolved. Bismarck Is 4: New Britain New Guinea Solomon Is a ~ Louisiade Is Australia Fig. 53: Map of Malesia, with distribution range of Calicnemiinae indicated. 4.4. Implications for biogeography As a group of insects occuring to the west and to the east of the Wallace line (Fig. 53), the Calicnemiinae are of special interest for biogeographical studies in the Indo-Australian transition zone. The phylogenetic hy- pothesis presented above allows for the postulation of active or passive dispersal events both into the East as well as, subsequently, back into the West (i.e. the Phil- ippines). Since a detailed scenario for the distributional history of the group has been developed elsewhere in the context of the historical biogeography of the fresh- water biota in Southeast Asia (VAN TOL & GASSMANN, submitted), the reader is referred to that forthcoming paper. Acknowledgments. It is an honour to me to dedicate the present paper to the memory of Professor Clas M. Naumann (Zoological Research Institute and Museum Alexander Koenig Bonn, and University of Bonn/ Germany), whom I owe a long lasting motivation and en- thusiasm to work in the field of organismic biology; this as well as his support during the developing PhD thesis will be gratefully remembered. The present project is conducted under the direction of Prof. Dr. E. Gittenberger and J. van Tol (Leiden University and Natural History Museum, Lei- den/Netherlands). I am grateful to Dr. H. Duffels (Univer- sity of Amsterdam) for comments on the manuscript. I thank K.-D. Dijkstra (Leiden), Dr. M. Hämäläinen (Hel- sinki), Dr. A. Martens (Braunschweig) and Dr. J. Kuhn (Seewiesen) for information of different kinds, as well as Dr. H. Turner and C. Riiffler (Leiden) for general discus- sions. Most of curators and collection management staff of various collections I have either visited or received loans from have been acknowledged in previous papers. For ad- ditional specimens included here, I thank K.-D. Dijkstra, V. J. Kalkman (Leiden), Dr. W. Schneider (Hessisches Landesmuseum, Darmstadt) and Dr. G. Vick (Basingstoke, UK). I am grateful to Mrs. G. Lamers and Dr. W. de Priester (Institute of Biology, Leiden) for their permission to let me do my SEM studies; a minor part of the micro- graphs, however, has been made at the Natural History Museum Leiden, with thanks to J. Goud for assistance. Dr. John Michalski and Prof. Dennis Paulson (USA), Roland A. Müller (St. Gallen, Switzerland) and J. van Tol provided field photographs. For scanning of negatives and slides, I thank M. Brittijn and J. Meijvogel (IBL, Leiden). Originally, the present research project was funded by the Dutch Life Science Foundation (ALW, formerly SLW), which is subsidized by the Netherlands Organisation for Scientific Research (NWO). Dirk GASSMANN: The Phylogeny of Southeast Asian and Indo-Pacific Calicnemiinae 67 REFERENCES ARCHIE, J. W. (1985): Methods for coding variable mor- phological features for numerical taxonomic analysis. Systematic Zoology 34(3): 326-345. BECHLY, G. H. P. (1996): Morphologische Untersuchungen am Flúgelgeáder der rezenten Libellen und deren Stammgruppenvertreter (Insecta; Pterygota; Odonata) unter besonderer Berücksichtigung der phyloge- netischen Systematik und des Grundplanes der Odo- nata. Petalura (Spec. Vol.) 1: 1-341. ERIKSSON, T. (1998): AutoDecay version 4.0 (program dis- tributed by the author). Bergius Foundation, Royal Swedish Academy of Sciences, Stockholm. FRASER, F. C. (1932): Indian Dragonflies. Journal of the Bombay Natural History Society 36(1): 141-151. FRASER, F. C. (1957): A reclassification of the Order Odo- nata. Royal Zoological Society of South Wales, Sydney. GASSMANN, D. (1999): Taxonomy and distribution of the inornata species-group of the Papuan genus /diocnemis Selys (Odonata: Zygoptera: Platycnemididae). Inver- tebrate Taxonomy 13: 977-1005. GASSMANN, D. (2000): A revision of the Papuan /diocnemis bidentata-group (Odonata: Platycnemididae). Zoolo- gische Mededelingen Leiden 74(23): 375-402. GASSMANN, D. & HAMALAINEN, M. (2002): A Revision of the Philippine subgenus Risiocnemis (lgneocnemis) Hämäläinen (Odonata: Platycnemididae). Tijdschrift voor Entomologie 145(2): 213-266. HAMALAINEN, M. (199la): The Philippine genus Ri- siocnemis Cowley (Zygoptera: Platycnemididae). I. Subgenus Risiocnemis. Odonatologica 20(2): 151-194. HAMALAINEN, M. (1991b): Preliminary descriptions of seven new Risiocnemis species of the subgenus /g- neocnemis Hämäläinen, 1991 (Zygoptera: Platycne- mididae). Odonatologica 20(2):195-201. HAMALAINEN, M. (2000): Risiocnemis seidenschwarzi spec. nov. - an endangered damselfly from Tabunan forest in Cebu, the Philippines (Odonata: Platycne- mididae). Entomologische Berichten, Amsterdam 60(3):46-49. HAMALAINEN, M. (2003): Platycnemis phasmovolans sp.nov. - an extraordinary damselfly from Laos with notes on its East Asian congeners (Odonata: Platy- cnemididae). Tombo 46(1/4):1-7. HENNIG, W. (1950): Grundzüge einer Theorie der phyloge- netischen Systematik. Deutscher Zentralverlag Berlin. HENNIG, W. (1966): Phylogenetic Systematics. University of Illinois Press, Urbana. KITCHING, I. J., FOREY, P. L., HUMPHRIES, C. J. & WIL- LIAMS, D. M. (1998): Cladistics. The theory and prac- tice of parsimony analysis. 2nd edition. Oxford Uni- versity Press. KORNET, D. J. & TURNER, H. (1999): Coding polymor- phism for phylogeny reconstruction. Systematic Biol- ogy 48(2): 365-379. LIEFTINCK, M. A. (1932): The dragonflies (Odonata) of New Guinea and neighbouring islands, Part I: Descrip- tions of new genera and species of the families Lesti- dae and Agrionidae. Nova Guinea 15(3): 485-602. LIEFTINCK, M. A. (1933): The dragonflies (Odonata) of New Guinea and neighbouring islands. Part II. De- scriptions of a new genus and species of Platycnemi- nae (Agrionidae) and of new Libellulidae. Nova Guinea 17(1): 1-66. LIEFTINCK, M. A. (1949): The dragonflies (Odonata) of New Guinea and neighbouring islands. Part VII. Results of the third Archbold Expedition 1938-1939 and of the Le Roux Expedition 1939 to Netherlands New Guinea. II. Zygoptera. Nova Guinea (N.S.) 5: |- 2: LIEFTINCK, M. A. (1956): Two new Platyenemididae (Odb- nata) from the Papuan region. Nova Guinea (N.S.) 7(2): 249-256, figs. 1-7. LIEFTINCK, M. A. (1958): A review of the genus /diocne- mis Selys in the Papuan Region, with notes on some larval forms of the Platyenemididae (Odonata). Nova Guinea (N.S.) 9: 253-292. LIEFTINCK, M. A. (1963): Contributions to the odonate fauna of the Solomon Islands, with notes on zygopter- ous larvae. Nova Guinea (Zoology) 21: 523-542. LIEFTINCK, M. A. (1965): Notes on Odonata of Madagas- car, with special reference to the Zygoptera and with comparative notes on other faunal regions. Verhand- lungen der Naturforschenden Gesellschaft Basel 76(2): 229-256, figs. 1-12. LIEFTINCK, M. A. (1971): A catalogue of the type- specimens of Odonata preserved in the Netherlands, with a supplementary list of the Odonata types de- scribed by Dutch scientists deposited in foreign institu- tional collections. Tijdschrift voor Entomologie 114(2): 65-139 + Pl. 1-7. LIEFTINCK, M. A. (1974): Dragonflies collected by the Noona Dan Expedition in the southwestern Philippine Islands (Insecta, Odonata). Steenstrupia 3(12): 11 1- 147. LIEFTINCK, M. A. (1984): Further notes on the specific characters of Calicnemia Strand, with a key to the males and remarks on some larval forms (Zygoptera: Platycnemididae). Odonatologica 13(3): 351-375. MADDISON, D. R. & MADDISON, W. P. (1992): MacClade. Analysis of phylogeny and character evolution. Ver- sion 3. Sinauer Associates, Sunderland, Massachusetts. MARTENS, A. (1996): Die Federlibellen Europas. Die Neue Brehm-Bücherei, Bd. 626. Magdeburg. REHN, A. C. (2003): Phylogenetic analysis of higher-level relationships of Odonata. Systematic Entomology 28(2): 181-239. SCHMIDT, E. (1951a): The Odonata of Madagascar. Zygop- tera. Mémoires de !'institut scientifique de Madagascar (Série A): 6(1): 115-283. SCHMIDT, E. (1951b): Uber neue und weniger bekannte af rikanische Platyenemididen (Odon.). Zygoptera. Me- moires de l'institut scientifique de Madagascar (Série A) 6(1): 115-283. SCHORR, M., LINDEBOOM, M. & PAULSON, D. (2004): List of Odonata of the world (Part 1, Zygoptera and Aniso- zygoptera). University of Puget Sound. [Internet web site at: http: //www.ups.edu/biology/museum/worldo- donates.html] SELYS-LONGCHAMPS, E. de (1863): Synopsis des Agrioni- nes, Quatriéme légion: Platycnemis. Bulletin de l'Aca- démie Belgique 2: 147-176. SELYS-LONGCHAMPS, E. de (1886): Révision du synopsis des Agrionines, I. Les Legions Pseudostigema, Poda- grion, Platycnemis et Protonevra. Mémoires Couron- nés et autre Mémoires. Académie Royale des Sciences des Lettres et des Beaux-Arts de Belgique 38(4): i-iv + 1-233. 68 Bonner zoologische Beitráge 53 (2004) SWOFFORD, D. L. & BEGLE, D. P. (1993): PAUP: Phyloge- netic analysis using parsimony, Version 3.1. User's manual. Smithonian Institution, Washington D.C. SWOFFORD, D. L. (2002): PAUP*: Phylogenetic Analysis Using Parsimony (* and other methods). Version 4.0b10. Sinauer Associates, Sunderland, Massachu- setts. TILLYARD, R. J. (1917): The biology of dragonflies (Odo- nata or Paraneuroptera). Cambridge University Press, Cambridge. TILLYARD, R. J. & FRASER, F. C. (1938-40): A reclassifica- tion of the order Odonata. Australian Zoologist 9: 125- 169, 195-221, 359-390. TOL, J. VAN (2000): The Odonata of Sulawesi and adjacent islands. Part 5. The genus Protosticta Selys (Platystic- tidae). Tijdschrift voor Entomologie 143(2): 221-266. WATERSTON, A. R. (1984): A new genus and species of platycnemidid dragonfly from the Arabian peninsula (Zygoptera). Odonatologica 13(1):139-146. WATSON, J. A. L. & O'FARRELL, A. F. (1991): Odonata (Dragonflies and Damselflies). Pp. 294-310 in: NAUMANN, I. D. (ed.) The Insects of Australia. 2nd ed., vol. 1., Melbourne. WIENS, J. J. (1999): Polymorphism in systematics and comparative biology. Annual Review of Ecology and Systematics 30: 327-362. WILSON, K. D. P. & WEN-BAO, Z., (2000): Sinocnemis yangbingi gen. nov. sp. nov. and Sinocnemis dumonti sp. nov., new platycnemidids from South-West China (Odonata: Platycnemididae). International Journal of Odonatology 3(2): 173-177. Y AKOBSON, G. G. & BIANKI, V. L. (1905): Orthoptera and Pseudo-Neuroptera of the Russian Empire and adjoin- ing countries. St. Petersburg (In Russian). Author’s address: Dirk GASSMANN, Institute of Biol- ogy, Leiden University, Evolutionary Biology Section, c/o National Museum of Natural History (Naturalis), P.O. Box 9517, NL-2300 RA Leiden, The Netherlands. E-mail: Gassmann@rulsfb.leidenuniv.nl Dirk GASSMANN: The Phylogeny of Southeast Asian and Indo-Pacific Calicnemiinae APPENDIX 1 Taxa included in the cladistic analysis and their distribution (m = male, f = female; [*] = only literature data available; [-] = not known). Allocnemis Selys, 1863 Arabicnemis Waterston, 1984 Asthenocnemis Lieftinck, 1949 Arrhenocnemis Lieftinck, 1933 Calicnemia Strand, 1928 Coeliccia Kirby, 1890 Copera Kirby, 1890 Cyanocnemis Lief- tinck, 1949 Idiocnemis Selys, 1878 bidentata species- group Allocnemis leucosticta Selys, 1863 Arabicnemis caerulea Waterston, 1984 Asthenocnemis stephanodera Lieftinck, 1949 Arrhenocnemis amphidactylis Lieftinck, 1949 Arrhenocnemis sinuatipennis Lieftinck, 1933 Calicnemia eximia (Selys, 1863) Coeliccia membranipes (Ram- bur, 1842) Copera marginipes (Rambur, 1842) Cyanocnemis aureofrons Lieftinck, 1949 Idiocnemis bidentata Selys, 1878 Idiocnemis dagnyae Lieftinck, 1958 Idiocnemis inaequidens Lieftinck, 1932 Idiocnemis mertoni Ris, 1913 m/f* m/f* m/f m/- m/- m/f m/f m/f m/f m/f m/f m/f m/f Southern Africa Arabian Peninsula: Oman, Yemen Philippines: Palawan Central northern New Guinea Northern New Guinea Mainland southern/southeastern Asia, Hainan (first record), Taiwan Greater Sunda Islands: Java, Sumatra Southern and southeastern Asia Central northern New Guinea Western New Guinea: Bird's Head Peninsula: Batanta; Salawati; Waigeo Western New Guinea: Bird's Head Peninsula and southwestern New Guinea: Misool Northeastern and eastern New Guinea Southern New Guinea: Aru Islands 70 [excluded, see GASSMANN (2000)] Indocnemis Laidlaw, 1917 Leptocnemis Selys, 1886 Lieftinckia Kimmins, 1957 inornata species- group Bonner zoologische Beitráge 53 (2004) Idiocnemis nigriventris Lieftinck, 1937 Idiocnemis obliterata Lieftinck, 1932 Idiocnemis polhemi Gassmann, 2000 Idiocnemis pruinescens Lieftinck, 1937 Idiocnemis adelbertensis Gassmann, 1999 Idiocnemis australis Gassmann, 1999 Idiocnemis chlo- ropleura Lieftinck, 1932 Idiocnemis fissidens Lieftinck, 1958 Idiocnemis huonensis Lieftinck, 1958 Idiocnemis inornata Selys, 1878 Idiocnemis kimminsi Lieftinck, 1958 Idiocnemis leonardi Lieftinck, 1958 Idiocnemis louisiadensis Lieftinck, 1958 Idiocnemis strumidens Lieftinck, 1958 Idiocnemis zebrina zebrina Lieftinck, 1958 Idiocnemis leonorae Lieftinck, 1949 Indocnemis orang (Forster in Laidlaw, 1907) Leptocnemis cyanops (Selys, 1869) Lieftinckia isabellae Lieftinck, 1987 Lieftinckia kimminsi Lieftinck, 1963 Lieftinckia lairdi m/f m/f m/f m/f m/f m/f m/f m/f m/f m/f m/f m/- m/f m/f m/f m/f m/f* m/f* m/- m/f m*/f* Northern New Guinea Northern and central northern New Guinea Southeastern central New Guinea Papuan Peninsula Northeastern New Guinea: Adelbert Range Southern central and southern New Guinea Northern New Guinea West Papuan Islands: Waigeo Northeastern New Guinea: Huon Peninsula Western New Guinea: Bird's Head Peninsula Bismarck Archipelago: New Britain, Mioko Louisiade Archipelago: Tagula (Sudest) Louis. Arch.: Misima, Rossel; Marshall Ben- nett Arch.: Woodlark Western New Guinea: Bird's Head Peninsula and southwestern New Guinea; Misool Northeastern and central Papuan Peninsula Northeastern and southern central New Guinea Southern and southeastern Asia Seychelles Solomon Islands: Santa Ysabel Solomon Islands: Bougainville Solomon Islands: Guadalcanal Lochmaeocnemis Lieftinck, 1949 Mesocnemis Karsch, 1891 Paracnemis Martin, 1903 Paramecocnemis Lieftinck, 1932 Platycnemis Burmeister, 1839 Rhyacocnemis Lieftinck, 1956 Risiocnemis Cowley, 1934 Dirk GASSMANN: The Phylogeny of Southeast Asian and Indo-Pacific Calicnemiinae Subgenus Risioc- nemis (Risiocne- mis) Cowley Lieftinck, 1963 Lieftinckia malaitae Lieftinck, 1987 Lieftinckia ramosa Lieftinck, 1987 Lieftinckia salomonis Kimmins, 1957 Lochmaeocnemis malacodora Lieftinck, 1949 Mesocnemis singularis Karsch, 1891 Paracnemis alluaudi Martin, 1903 Paramecocnemis erythrostigma Lieftinck, 1932 Paramecocnemis stilla-cruoris Lieftinck, 1956 Platycnemis pennipes (Pallas, 1771) Rhyacocnemis prothoracica Lieftinck, 1987 Rhyacocnemis sufficiens Lieftinck, 1956 Risiocnemis appendiculata (Brauer, 1868) Risiocnemis arator Hämäläinen, 1991 Risiocnemis asahinai Kitagawa, 1990 Risiocnemis confusa Hämäläinen, 1991 Risiocnemis elegans Kitagawa, 1990 m/f m/- m/f m/f m/f m/f m/f* m*/- m/f m*/- m/f m/f m/f m/f* m/f Solomon Islands: Malaita Solomon Islands: San Jorge; ? Bougainville Solomon Islands: Guadalcanal / Central northern New Guinea Western Africa Madagascar Northern New Guinea Northeastern New Guinea Western and central Europe Northeastern New Guinea D'Entrecasteaux Islands: Goodenough Philippines: Mindanao; Dinagat; Camiguin; Samar; Leyte; Panaon; Biliran; Bohol; Ho- monhon Philippines: northern/central Luzon Philippines: southern Luzon; Mindoro Philippines: southern Luzon: Catanduanes Philippines: Luzon Subgenus Risiocnemis (/gneocnemis ) Hämäläinen Bonner zoologische Beiträge 53 (2004) Risiocnemis erythrura (Brauer, 1868) Risiocnemis gracilis Hämäläinen, 199] Risiocnemis kiautai Hämäläinen, 1991 Risiocnemis laguna Hämäläinen, 199] Risiocnemis moroensis Hämäläinen, 1991 Risiocnemis praeusta Hämäläinen, 1991 Risiocnemis pulchra Hämäläinen, 1991 Risiocnemis rolandmuelleri Hämäläinen, 1991 Risiocnemis seidenschwarzi Hämäläinen, 2000 Risiocnemis serrata (Hagen ın Selys, 1863) Risiocnemis varians Hämäläinen, 1991 Risiocnemis antoniae Gassmann and Hämäläinen, 2002 Risiocnemis atripes (Needham and Gyger, 1941) Risiocnemis atropurpurea (Brauer, 1868) Risiocnemis calceata Hämäläinen, 1991 Risiocnemis flammea (Selys, 1882) Risiocnemis fuligifrons Hämäläinen, 1991 Risiocnemis haematopus (Selys, 1882) Risiocnemis ignea (Brauer, 1868) Risiocnemis incisa Kimmins, 1936 Risiocnemis kaiseri Gassmann and Hämäläinen, 2002 Risiocnemis melanops Hämäläinen, 1991 Risiocnemis nigra m/f m/f m/f* m/f m/f m/f m/f m/f m/f* m/f m/f m/f m/f m/f m/f m/f m/f m/f m/f m/f m/f m/f m/f Philippines: eastern Mindanao; Siargao Philippines: central Luzon Philippines: Sibuyan Philippines: southern Luzon Philippines: central Mindanao Philippines: Dinagat; Samar; Leyte; Panaon; Biliran Philippines: southern Luzon Philippines: Sibuyan; Panay; Negros; Mas- bate; Siquijor Philippines: Cebu Philippines: Luzon; Polillo; Marinduque; Ca- tanduanes Philippines: central northern Luzon. Philippines: northeastern/eastern Mindanao Philippines: central/eastern Mindanao Luzon; Marinduque Dinagat; Panaon Philippines: Mindanao; Dinagat; Biliran; Homonhon; Panaon; Leyte; Samar Philippines: western/northern Mindanao; Di- nagat; Basilan; Panaon; Leyte Philippines: southern Luzon; Catanduanes Philippines: northern/northeastern Luzon Philippines: northern and central Luzon Philippines: Samar Philippines: Samar Philippines: Samar Dirk GASSMANN; The Phylogeny of Southeast Asian and Indo-Pacific Calicnemiinae Gassmann and Hämäläinen, 2002 Risiocnemis odobeni m/f Philippines: southeastern Luzon; Catandua- Hämäläinen, 1991 nes Risiocnemis pistor m/- Philippines: southeastern Mindanao Gassmann and Hámáláinen, 2002 Risiocnemis plebeja m/f Philippines: Sibuyan; Panay Hámáláinen, 1991 Risiocnemis m/f Philippines: southeastern Luzon; Catandua- polilloensis nes; Polillo Hämäläinen, 1991 Risiocnemis m/- Philippines: eastern Mindanao rubricercus Gassmann and Hämäläinen, 2002 Risiocnemis rubripes m/f Philippines: eastern Mindanao: Dinagat (Needham and Gyger, 1939) Risiocnemis siniae m/f Philippines: Leyte; Samar Hämäläinen, 1991 Risiocnemis tendipes m/f Philippines: Mindanao (Needham and Gyger, 1941) Salomocnemis Lieftinck, 1987 Salomocnemis gerdae m/f Solomon Islands: Guadalcanal Lieftinck, 1987 Stenocnemis Karsch, 1899 Stenocnemis m/f Western Africa pachystigma (Selys, 1886) Torrenticnemis Lieftinck, 1949 Torrenticnemis m/f Central northern New Guinea filicornis Lieftinck, 1949 74 APPENDIX 2 Data matrix for the cladistic analysis of Calicnemiinae 1 1234567890 Idiocnemis adelbertensis __ Idiocnemis australis Idiocnemis chloropleura Idiocnemis fissidens Idiocnemis huonensis Idiocnemis inornata Idiocnemis kimminsi Idiocnemis leonardi Idiocnemis louisiadensis Idiocnemis strumidens Idiocnemis zebrina Idiocnemis bidentata Idiocnemis dagnyae Idiocnemis inaequidens Idiocnemis mertoni Idiocnemis nigriventris Idiocnemis obliterata Idiocnemis polhemi Idiocnemis pruinescens Risiocnemis appendiculata Risiocnemis arator Risiocnemis asahinai Risiocnemis confusa Risiocnemis elegans Risiocnemis erythrura Risiocnemis gracilis Risiocnemis kiautai Risiocnemis aguna Risiocnemis moroensis Risiocnemis praeusta Risiocnemis pulchra Risiocnemis rolandmuelleri Risiocnemis seidenschwarzi Risiocnemis serrata Risiocnemis varians Risiocnemis antoniae Risiocnemis atripes Risiocnemis atropurpurea Risiocnemis calceata Risiocnemis flammea Risiocnemis fuligifrons Risiocnemis haematopus Risiocnemis ignea Risiocnemis incisa Risiocnemis kaiseri 201 201 201 201 201 201 201 201 20 201 201 20 201 201 20 201 20 201 201 20 201 20 202 20 20 201 20 201 20 201 201 202 202 20 20 10 10 10 101 101 101 10 101 101 10 1400000 1400000 1300000 1200000 1400000 1300000 1400000 1300000 1300000 1300000 1300000 1400000 1300000 1300000 1300000 1300000 1300000 1300000 1300000 1400000 1400000 1400000 1400000 1400000 1400000 1400000 1400000 1400000 1400000 400000 1400000 400000 400000 1400000 400000 200000 200000 200000 1200000 200000 200000 1200000 200000 1200000 200000 1111111112 1234567890 2200001355 3200001355 2200001355 2200001355 2200001355 2200001355 2200001355 4200001355 3200001355 2200001355 3200001355 22000013A0 2200001397 2200001000 2200001334 2200001363 2200001334 2200001320 2200001040 4400011570 440001 10B0 340001 10C9 4400011010 440001 16C9 44000111B0 4400011123 440001 16C9 44000111C9 44000120B0 440001 10B0 440001 16C9 44000111B0 440001 10C0 440001 16C9 440001 10C9 4400001000 44000010B0 4400001000 4400001000 4400001574 4400001070 4400001000 4400001276 4400001242 4400001000 1234567890 3217720423 3217720423 3217720423 3217720423 3217720423 32177270423 3217720423 3217270423 3212720423 3201040400 32172270423 1201040400 3201040400 3201070403 3201040400 3201040401 3201040400 2217772423 0001017400 0217772400 0000117400 02127222400 1017772400 8001117408 1001117400 0217777400 0217772400 8001012400 0217777400 8000117400 0312772420 0312222400 2312772422 4317222422 0317777400 1234567890 3200100221 3200100221 3201101221 3201101221 3202100221 3201100221 3202100221 3201100221 3201100221 3201101221 3201100221 0223100021 3202}0100221 0223100021 0223100021 0203100021 0223 00021 1223100021 0223100021 2200 00221 0300110021 0300110021 0300 10021 0300111021 0300 10021 0200110021 0300110021 0200 10021 0200110021 0300110021 0200110021 8300110021 0300117071 0300111021 0300111021 0000100071 0000100001 0000100080 {02}000100051 5000100271 2000100271 0000100071 2000100271 2000100271 2000100041 Bonner zoologische Beiträge 53 (2004) 4444444445 5555555556 6666666667 1234567890 1234567890 1234567890 0052100000 200012710? 2010002155 0052100000 2000127107 2020002155 0072100000 200013210? 2010002155 0052101000 200013710? 2030002155 0050700000 2000127107 2010002255 0071200000 2000137107 2010002255 0070200000 2000127107 2010102255 0070201000 2000147107 2020002255 0070201000 2000127107 2020002255 0052100000 2000132102 2040002155 0070201000 2000237107 2020002155 0202300000 0100137117 2150003155 0101;72300000 010013711? 2030003155 0202300100 010014712? 2050003255 0200200000 01001472117 2050003155 0212300000 0100137137 2120003255 0212300000 010013211? 2150003155 0202300000 010013711? 2150003155 0202300100 0100237107 2050003255 0070202000 200022314? 2007000255 0270202000 200012114? 2002001455 0272012000 2000233187 2007001455 0270207001 200014113? 2002001455 0272012000 011022116? 2007001445 0270207001 200032114? 2002001455 0272022000 011033110? 2007001455 0270202000 200032114? 2007001455 0272012000 011032118? 2060001445 0270207001 200053314? 2007001455 0270207001 200032114? 2002001455 0272027000 0110321187 2060001445 0272002000 2000121137 2007001455 0272002000 200024117? 2072001455 0232012000 011034113? 2002001455 0272012000 011022118? 2060001445 0270202000 2000223100 1002001413 0270202000 2000141151 0002001433 0270702000 2000243162 0002001433 0270707001 2000233152 0007001434 0070702000 2000343100 1002001400 0070202000 2000345101 2007001411 0270202000 2000343101 1002001411 0070207000 2000255101 1007001433 0070202000 2000243433101 0002001433 0270702000 2000444101 1002001411 7777777778 1234567890 0025011101 0025001200 0125011101 0025011133 0025011101 0025001401 0025011101 0025001201 0025001201 0025011133 0025001300 0021101011 0025101011 0020101021 0025101011 0025101021 0025101011 0020101011 0021101011 0022001300 0020000701 0027000100 0022001300 1027000100 0022001300 0027000100 1027001300 0027000100 0023001300 1022001300 0027000100 0020000300 002000030? 0027000100 0027000100 2022001502 2021001502 2021001502 2022001502 2022001502 2021001502 2021001502 2022001202 2022001502 2021001502 88888888 12345678 00020218 00020219 00020214 00077219 00020218 0002021A 00020218 0002077? 0002021C 00070219 0011021C 000?021B 000%021B 00020210 000?021B 00020210 00070210 00020210 0007021D 0000020? 0000000? 0020000? 0000020? 0020000? 0000020? 0020000? 0000020? 0020000? 0000020? 0000020? 0020000? 0000020? 0000720? 0020000? 0020000? 00000213 0000021K 0000221H 0000321] 0000021E 0000021H 0000021H 0000021H 0000021) 0000021E APPENDIX 2 (CONTINUED) Risiocnemis melanops Risiocnemis nigra Risiocnemis odobeni Risiocnemis plebeja Risiocnemis pistor Risiocnemis polliloensis Risiocnemis rubripes Risiocnemis rubricercus Risiocnemis siniae 1011200000 Risiocnemis tendipes 1011200000 Paramecocnemis erythrostigma 2011200000 Paramecocnemis stilla-cruoris 2011200000 Cyanocnemis aureofrons 2001231110 Torrenticnemis filicornis 2001231000 Idiocnemis leonorae 2011230000 Lochmaeocnemis malacodora 2001232000 Arrhenocnemis amphidactylis 2001410000 Arrhenocnemis sinuatipennis 2001410000 Rhyacocnemis prothoracica 2011220000 Rhyacocnemis sufficiens 2011220000 Salomocnemis gerdae 2011200000 Lieftinckia isabellae 2031200000 Lieftinckia kimminsi 2031200000 Lieftinckia lairdi 2031400000 Lieftinckia malaitae 2031470000 Lieftinckia ramosa 2031400000 Lieftinckia salomonis 2031470000 Coeliccia membranipes 0011232020 Asthenocnemis stephanodera 0011222020 Calicnemia eximia 0001232010 Indocnemis orang 0001232221 Arabicnemis caerulea 0100030000 Leptocnemis cyanops 1011232020 Allocnemis leucosticta 0010220000 Mesocnemis singularis 0101032221 Paracnemis alluaudi 0000232120 Stenocnemis pachystigma 0001130000 Copera marginipes 0000400000 Platycnemis pennipes 0000200000 | 1234567890 1011200000 1011200000 1011200000 1011200000 1011200000 1011200000 1011200000 1011200000 1111111112 1234567890 4400001042 4400001000 4400001 102181061 4400001542 4400001000 4400001000 4400001245 4400001040 4400001000 4400001000 2210001720 2210021800 0001021028 0000031000 010000184B 0000031020 24000018E0 4400001660 1000021820 1000011820 02000018EB 2000011001 2300001001 22000712? 2200001271 2200011271 2200011276 11010010C0 22000016C0 0001001348 0101003020 00000007DA 0001011320 2201131820 0000001230 00010016B0 00000013A8 0000000450 000000145A 1234567890 0312222400 0217222400 (02331777241 {02} 3317772422 0317777400 0217772400 2312222422 0217772400 0312772 0317722420 7201070400 6201072400 0102057404 4201062400 6201032400 0217772400 600006 1300 8200017726 5200032400 5201032400 621227242? 0200007400 0200017400 2212229423 2217272423 22177724 {02}0 2712272493 8217722100 8200002006 4202052004 0219777100 7212992437 8200011000 4212722000 7217772000 7201022000 3333333334 1234567890 2000100271 0000100011 102/000100271 2000100271 0000100271 0000100041 2000100271 0000100001 0000100051 2000100271 0203101211 0203101011 0200100001 0203100001 0203100711 0300100001 0300100021 3203100721 0303100011 0300100011 2000101721 2000100721 0300100021 3300107121 3500100221 0100100221 2400100121 0300000021 6203102221 4300000061 0300000001 7200100281 0200100021 0500100011 0500100011 2200100081 4300100011 0100000030 0300001080 4444444445 1234567890 0070202000 0270702000 0070202001 0070707000 0270202000 0270702000 0070707000 0270702000 0270207001 0270707000 0222007000 0202607000 0272007000 0202402010 0062702000 0202707000 1202002000 0202007000 0222207000 0202207000 0272207000 0072202000 0227202000 0072202000 0077207000 0022202000, 0072207000 0202002000 0002002000 0202402000 0202502000 0000707000 0242207000 0202702000 0222707200 0202207000 0202702000 0202702000 0202502000 Dirk GASSMANN: The Phylogeny of Southeast Asian and Indo-Pacific Calicnemiinae 5555555556 6666666667 7777777778 888888! 1234567890 1234567890 1234567890 9000343101 1002001411 2021001502 0000021H 2000343100 1002001400 2021001502 0000021É y 2000245100 1002001400 2021001502 00000215 2000242102 0002001433 2024001502 00000217 2000343152 1002001433 2022001502 0000177? 2000343101 1002001410 2021001502 0000021 H 2000243152 1002001433 2022001502 0000121 H 2000143152 0007001422 2022001502 00001777 2000245152 0007001434 2022001502 0000321) 2000253152 0002001433 2022001502 00000217 010165702? 2082004255 3020001801 00020210 200164207? 2087004355 302020721 “777277? 010103?11? 2071002255 0020101201 00030210 010102607? 20072002255 0020021101 10020211 010106711? 2050001055 2720001801 11070212 010103710? 2040012255 2020001901 01020210 200113207? 2002001255 0024977277 27727777 200112207? 2032001355 0024001301 0002077 010116711? 2002002055 2020001701 2202077 010116711? 002002055 2720001101 110207? 200021411? 2002001455 2025001301 00110710 20001411A? 2011101255 0024707771 2077277? 200022119? 2012001355 0021001201 00010213 200071207? 2002001455 0025001300 00110713 200011207? 2002001455 0021901771 20772213 200022107? ?012001355 0024720171 2077777? 200011110? 2002001455 2024001301 00110213 011021119? 2070001455 1007000200 1030410? 010112607? 2082001355 3020001201 01007115 200011119? 2002001455 00150007?1 00000214 011011219? 2070001455 10000007?1 20000214 200023110? 2002001455 2117001 A01 0000010? 271024710? 2002002255 2120001701 0000020? 1110212192? 2002001255 0015??1B01 0007020? 211005110? 2002001455 20200007?1 2000420? 011012010? 2007007255 = 5220007701 00020210 111012410? 2007001455 2120001601 00007214 111040207? 2002002255 0016001201 00007116 111001102? 2002001455 4007000771 0007510? 76 Bonner zoologische Beitráge 53 (2004) APPENDIX 3 List of unambigious character state changes along branches of the preferred most parsimonious cladogram (ef. fig. 52). Branch 1: char. 18: 05; char. 19: B—7; char. 21: 0—2; char. 29: 02; char. 30: 03; char. 31: 02; - char. 36: 1—0; char. 38: 02; char. 42: 2—0; char. 67: 1—0; char. 68: 4—2. Branch 2: char. 17: 1—2; char. 50: 01; char. 56: 2—3; char. 74: 2—3. Branch 3: char. 11: 4-3; char. 55: 32; char. 56: 2—3; char. 57: 1—3. Branch 4: char. 59: 86; char. 71: 0-1; char. 88: 1—2. Branch 5: char. 63: 0—6. Branch 6: char. 21: 0—8; char. 43: 7—3; char. 56: 2—4; char. 59: 83; char. 88: 1—4. Branch 7: char. 19: C—2; char. 20: 9—3; char. 23: 01; char. 56: 23; char. 59: 80. Branch 8: none. Branch 9: char. 21: 1—8. Branch 10: char. 21: 08; char. 30: 0-8; char. 71: 0-1. Branch 11: char. 18: 0-1. Branch 12: char. 71: 0>1. Branch 13: char. 19: B—1; char. 50: 0-1. Branch 14: char. 18: 0>1; char. 21: 0>1; char. 31: 08. Branch 15: char. 19: B>C; char. 55: 1—2; char. 59: 3>7. Branch 16: char. 74: 2—0; char. 77: 1—0; char. 86: 2—0. Branch 17: char. 22: 23; char. 56: 4—2. Branch 18: char. 29: 0—2; char. 57: 35; char. 61: 1—2. Branch 19: char. 20: 0—2; char. 88: 1—2. Branch 20: char. 39: 7—4; char. 55: 3-4; char. 57: 3-4. Branch 21: none. Branch 22: char. 39: 7—4. Branch 23: char. 39: 7>1; char. 69: 1—0. Branch 24: char. 39: 7—8; char. 40: 1—0; char. 85: 02. Branch 25: char. 56: 4>3. Branch 26: char. 57: 35; char. 88: 4-43. Branch 27: char. 18: 25; char. 20: 6—4; char. 22: 3-2; char. 30: 2—5; char. 31: 2—5; char. 55: 23. Branch 28: char. 19: 7>8; char. 29: 2—1; char. 50: 0-1; char. 74: 2>1. Branch 29: char. 56: 4—5; char. 78: 6—2. Branch 30: char. 21: 2—4. Branch 31: char. 18: 25; char. 21: 23; char. 57: 32; char. 74: 2—4. Branch 32: char. 61: 0-1; char. 85: 0-1; char. 88: 4>1. Branch 33: char. 56: 4—5. Branch 34: char. 55: 2—3; char. 61: 0-1; char. 85: 0—1. Branch 35: char. 69: 3—2; char. 70: 3>2; char. 85: 0>1. Branch 36: char. 22: 2>3; char. 74: 2—1. Branch 37: char. 3: 3—1; char. 5: 4>2; char. 11: 2—0; char. 18: 2>8; char. 19: 7>E; char. 20: 1>B; char. 21: 26; char. 37: 0-1; char. 55: 1—2. Branch 38: char. 80: 1>0. Branch 39: char. 74: 5>1. Branch 40: char. 20: 1—6; char. 32: 0/3—4; char. 88: 4-3. Branch 41: char. 32: 0/3>1; char. 55: 1—2; char. 58: 1—0. Branch 42: char. 56: 224; char. 59: 0>A; char. 64: 231; char. 65: 0-1; char. 68: 3—2. Branch 43: char. 55: 1—2; char. 59: 09; char. 74: 4-41; char. 78: 3—2. Branch 44: char. 26: 1—6; char. 41: 0>1; char. 56: 2-3; char. 68: 3—2. Branch 45: char. 29: 0>2; char. 30: 0>6: char. 31: 03; char. 34: 0-3; char. 63: 0>3. Branch 46: none. Branch 47: none. Branch 48: char. 43: 5>7; char. 65: 0>1. Branch 49: char. 11: 2—3; char. 80: 1—0. Branch 50: char. 72: 0>1. Branch 51: char. 47: O>1. Branch 52: none. Branch 53: char. 44: 2>1; char. 45: 1>2; char. 68: 1—2; char. 88: 4>1. Branch 54: char. 11: 3—4. Branch 55: none. Branch 56: char. 55: 1—2; char. 80: 1—0; char. 83: 0>1; char. 84: 2—1. Branch 57: char. 63: 223. Branch 58: char. 59: 1—3; char. 68: 1>2; char. 79: 1—2. Branch 59: none. Branch 60: char. 44: 2—0; char. 56: 3—4; char. 62: 1—0. Branch 61: char. 5: 3>4. Branch 62: char. 30: 0>1; char. 31: 0-1. Branch 63: char. 21: 3>1; char. 56: 3—4; char. 79: 1—2. Branch 64: char. 55: 1>2; char. 74: 0>1. Branch 65: char. 21: 6>7; char. 38: 0—2; char. 43: 0>2. Branch 66: char. 19: 220; char. 45: 0>6; char. 52: 1—0; char. 68: 2>3. Branch 67: char. 12: 0>1; char. 19: 224; char. 20: O>B; char. 42: 2—0; char. 43: 0-6; char. 67: 2>1. Branch 68: char. 24: 1>0. Branch 69: char. 34: 3—0. Branch 70: char. 19: 2-40; char. 45: 0-4; char. 49: 0>1; char. 56: 3>2; char. 58: 1—0; char. 76: 0>2. Branch 71: char. 8: 0—1; char. 9: 0-1; char. 14: 0>1; char. 16: 0/3—2; char. 20: 0-8; char. 22: 251; char. 30: 0—4; char. 43: 0>7; char. 63: 0-7; char. 64: 0>1; char. 75: 0-1; char. 84: 2—3. Branch 72: char. 23: 0-1; char. 63: 0—4; char. 66: 0>1; char. 71: 0>2; char. 78: 2>A; char. 82: 0>1. Branch 73: char. 3: 0>1; char. 6: 3>2; char. 12: 0>2; char. 19: 2/B>C; char. 26: 1/5—0; char. 30: 06; char. 31: 0-6; char. 32: 3—2; char. 34: 0>3; char. 37: 0>2; char. 38: 0>2; char. 42: 220; char. 58: 1—0; char. 63: 0-8; char. 71: 0-3; char. 82: 0>1; char. 86: 2>1: char. 88: 4>3. Branch 74: char. 4: 1—0; char. 8: 0-1; char. 27: 2>1; char. 39: 28: char. 71: 05; char. 72: 0>2. Branch 75: char. 3: 0>1; char. 11: 0>1; char. 19: 2—C; char. 55: 1>2; char. 74: 0-7; char. 80: 1—0; char. 83: 0>3; char. 85: 0—4; char. 86: 2>1; char. 87: 1>0. Branch 76: char. 8: 0>2; char. 10: 0-1; char. 17: 123; char. 21: 8-0; char. 39: 2-0; char. 45: 0-5; char. 57: 1—2. Branch 77: char. 9: 21; char. 19: 2—4; char. 20: 08; char. 21: 8—4; char. 30: 0—4; char. 31: 0-4; char. 39: 26; char. 45: 0—4; char. 52: 1—0; char. 53: 1—0; char. 74: 0—5. Branch 78: char. 1: 0>1; char. 3: 0>1; char. 16: 0>1; char. 28: 0>2; char. 32: 3>2; char. 43: 04; char. 56: 1/2—4; char. 67: 1>2; char. 68: 4>2; char. 72: O>1; char. 78: 2—8. Branch 79: char. 2: 0-1; char. 5: 2-0; char. 8: 2—0; char. 10: O0>1; char. 19: 2—3; char. 32: 3>5; char. 48: 0>2; char. 55: 1/2>0; char. 56: 1/2—5; char. 77: 1—0; char. 85: 0>4. Branch 80: char. 5: 2>1; char. 19: 2—A; char. 20: 0-8; char. 31: 024; char. 57: 124; char. 72: 0>1; char. 78: 227; char. 87: 0>1. Branch 81: char. 3: 0>1; char. 6: 3>2; char. 11: 0>2; char. 12: 0-2; char. 14: 0>1; char. 15: 0>1; char. 16: 0>3; char. 23: 1—0; char. 27: 2-1; char. 32: 3>5; char. 59: 0-9; char. 68: 4—2; char. 78: 2—C. Branch 82: char. 2: 0>1; char. 5: 220; char. 29: 0>3; char. 30: 0-7; char. 31: 0>7; char. 32: 3>2; char. 38: 0>2; char. 42: 20; char. 44: 240; char. 52: 1—0; char. 53: 1>0; char. 56: 1>3; char. 72: 0-1; Dirk GASSMANN: The Phylogeny of Southeast Asian and Indo-Pacific Calicnemiinae char. 78: 2—B. Branch 83: char. 23: 1—0; char. 37: 0>1; char. 45: 7-5; char. 73: 1-0; char. 77: 1-0. Branch 84: none. Branch 85: none. Branch 86: none. Branch 87: char. 6: 0>3; char. 35: 0-1; char. 40: 0—1; char. 58: O>1. Branch 88: char. 39: 8>1; char. 86: 1>2. Branch 89: char. 4: 0-1; char. 73: 1>2. Branch 90: char. 7: 0>2; char. 9: 0>2. Branch 91: char. 21: 0>2; char. 30: 0>2; char. 42: 2—0. Branch 129: char. 59: 5—0. Branch 130: char. 60: 2>1. Branch 131: char. 19: 4-7; char. 20: 2>6; char. 61: 0>1. Branch 132: char. 60: 1—0; char. 69; 3-0; char. 70: 3-0. Branch 133: char. 20: 0-1; char. 63: 0>1: char. 84: 0>1. Branch 134: char. 16: 0>1; char. 42: 20, Branch 135: char. 5: 2—4; char. 18: 0>2; char. 19: char. 14: 0-1; char. 39: 1>2. Branch 92: char. 21: 1>8; char. 71: 2—0; char. 87: 0-1. Branch 93: char. 18: 3>6. Branch 94: char. 14: 1—0; char. 53: 1-0; char. 54: 0>1. Branch 95: char. 1: 0>2; char. 9: 2>0; char. 28: 0>4. Branch 96: char. 7: 2>0; char. 52: 1—0; char. 74: 0>4; char. 78: 2—3. Branch’ 97: char. 43: 0>7; char. 54: 1>0. Branch 98: char. 23: 0-1: char. 44: 2-0; char. 68: 3—4; char. 74: 4>2. Branch 99: char. 5: 2—4; char. 16: 0>1; char. 36: 0>1; char. 87: 1>0. Branch 100: char. 80: 1—0. Branch 101: none. Branch 102: char. 32: 3—2; char. 57: 1—3. Branch 103: none. Branch 104: char. 19: B>C; char. 20: 0—9; char. 22: 2-40; char. 23: 1—0; char. 74: 27: char. 88: 4-3. Branch 105: char. 44: 0>2; char. 46: 0>1; char. 59: 4-8; char. 77: 1>0; char. 78: 3-1; char. 83: 0-2; char. 86: 2—0. Branch 106: char. 52: 0—1; char. 53: 0-1; char. 88: 3>1. Branch 107: char. 24: 1-0; char. 37: 0>1. Branch 108: char. 55: 3—2; char. 69: 5—4. Branch 109: char. 21: 0-1; char. 32: 3—2. Branch 110: char. 63: 0>6; char. 69: 5—4. Branch 111: char. 50: 0>1. Branch 112: char. 3: 1—>2; char. 59: 4-43. Branch 113: char. 44: 0>2; char. 74: 2-0; char. 77: 0>1. Branch 114: char. 1: 2—1; char. 32: 3-50; char. 39: 2-0; char. 56: 2—4; char. 69: 5—3; char. 70: 5>3; char. 71: 0>2; char. 78: 3—6; char. 80: 1—2. Branch 115: char. 57: 1>3. Branch 116: char. 39: 0>7; char. 55: 1>2. Branch 117: char. 88: 4—2. Branch 118: char. 60: 2>0; char. 61: 0-1; char. 69: 3>1. Branch 119: char. 55: 2>3; char. 70: 3-0. Branch 120: char. 60: 0>1; char. 88: 2>1. Branch 121: char. 22: 2233; char. 70: 0-1. Branch 122: char. 31: 0—2. Branch 123: char. 38: 0>2; char. 42: 20. Branch 124: char. 22: 2—3. Branch 125: char. 39: 7>5; char. 50: 0>1; char. 70: 34; char. 85: 0>3. Branch 126: char. 38: 02. Branch 127: char. 29: 02; char. 31: 0>2. Branch 128: char. 18: 0>2; char. 19: 0-4; 0>7; char. 21: 0>2; char. 23: 0-1. Branch 136: char. 30: 0>3; char. 56: 2>1; char. 63: 1—0; char. 68: 3-4. Branch 137: char. 16: 1—0; char. 74: 4—5. Branch 138: char. 58: 1>0. Branch 139: char. 5: 2—4; char. 58: 1—0. Branch 140: char. 39: 220; char. 55: 1—0; char. 56: 2>3; char. 67: 1—2; char. 68: 3>2; char. 84: 02. Branch 141: char. 7: 2>1; char. 32: 3—2; char. 59: 0—1. Branch 142: char. 34: 0-3. Branch 143: char. 3: 0—1; char. 7: 1—0; char. 39: 0>1. Branch 144: char. 6: 3-50; char. 11: 0—2; char. 12: 0>2; char. 27: 20. Branch 145: char. 5: 2>3; char. 21: 6>3; char. 33: 0—2; char. 39: 1>2; char. 45: 0-3; char. 54: 1—0; char. 75: 0-1; char. 79: 0>1. Branch 146: char. 62: 0>1; char. 68: 2>1. Branch 147: none. Branch 148: char. 20: 024. Branch 149: char. 43: 0>1. Branch 150: char. 30: 0-3; char. 33: 2—0; char. 63: 5—2. Branch 151: char. 23: 0>1; char. 29: 02; char. 31: 03: char. 34: 3-0; char. 38: 02; char. 42: 220; char. 43: 1—7; char. 62: 1—0. Branch 152: char. 45: 3>1; char. 52: 10; char. 59: 1—0; char. 67: 3>2; char. 75: 1—0; char. 78: 0>1/2; char. 79: 1>0. Branch 153: char. 5: 3—4; char. 43: 7-5; char. 56: 3>2. Branch 154: char. 63: 2>1; char. 76: 0>1. Branch 155: char. 34: 0>2; char. 44: 2—0; char. 68: 1>2. Branch 156: char. 34: 0—1. Branch 157: char. 63: 2>1. Branch 158: char. 37: 0—1; char. 76: 0>1. Branch 159: char. 43: 7—5; char. 79: 03; char. 80: 1—3. Branch 160: char. 11: 23; char. 44: 2—0; char. 47: 0-1. Branch 161: char. 68: 1—2. Branch 162: char. 48: 0—1. Branch 163: char. 13: 0>1; char. 37: 0-1; char. 58: 1—0; char. 71: 0-3. Branch 164: char. 56: 3—6; char. 68: 2—0; char. 71: 0—2; char. 82: 01. Branch 165: char. 6: 3—2; char. 11: 0>1; char. 21: 6>5; char. 32: 2—3. Branch 166: char. 35: 1—0; char. 59: 0-9; char. 77: 1—0. Branch 167: char. 12: 0-1; char. 18: 3—0; char. 28: 0-1; char. 63: 0-7; char. 71: 0>1. 78 Bonner zoologische Beitráge 53 (2004) APPENDIX 4 For detailed information on specimens studied, the reader should refer to the 'material examined' sections in GASSMANN (1999), GASSMANN (2000) and GASSMANN & HAMALAINEN (2002). Additional taxa/specimens examined for the present study are listed in the follow- ing. Collections: BMNH= Natural History Museum: British Museum (Natural History), London. HLMD= Hessisches Landesmuseum, Darmstadt. ISNB = Institut Royal des Sciences Naturelles, Brus- sels. RMNH= National Museum of Natural History (Natu- ralis; formerly: Rijksmuseum van Natuurli- jke Historie), Leiden (including the former Roland Müller collection). SMFD = Senckenberg-Museum, Frankfurt. Allocnemis leucosticta: South Africa, Cape Province: 44, Natal, Durban, 8.111.1939, RMNH. 24, Klein- Swartberge, Seweweekspoort Pass, 5.1.1951, Swedish S. Africa Exped. 1950-1951, RMNH. Arabicnemis caerulea: Arabian Peninsula, Oman: 14, Wadi Al-Abyadh, appr. 21°32'N 56°18'E, 16.1x.1988, ex Oman Nat. Hist. Mus., HLMD-Od 226, HLMD. Arrhenocnemis amphidactylis: New Guinea, West Papua: 14 (paratype), central New Guinea, above Bernhard Camp, 700 m, 29.11.1939, RMNH. Asthenocnemis stephanodera: Philippines, Palawan Is- land: 14, Quezon, Lamakan, Magmuni str, 27.v.- 1.vi.1991, RMNH. 14, Quezon, Lamakan, Magmuni str, 1000 ft., 16-22.v.1991, RMNH. 1%, Quezon, Mag- muni str, 1000 ft., 16-22.v.1992, RMNH. Calicnemia eximia: China, Hainan: 14, China, Central Hainan Island, Tongshi City, Tongshi Holiday Resort, 400 m, 28.vi.1993, RMNH. India, Assam: 1, Assam, Shillong, 25.vi.1974, RMNH. India, West Bengal: 1°, Bom Busty, 1.vi.1976, RMNH. 13, 1%, Gumpha, 6.vi.1976, RMNH. 14, Kapkot, 26.ix.73, RMNH. 14, Loharkhet, 26.ix.73, RMNH. India, Sikkim: 1%, Kool- loo [Kooloo?], Carleton, ISNB. 14, ex Staudinger, acq. 1903, RMNH. 14, ex Mus. Ann Arbor (coll. Forster), acg. 1936, RMNH. Nepal: 14, Barabhise, Tibettan frontier, ca. 950 m, 4.vi.1973, RMNH. 24, Charnavati Khola/pine for., 1200 m, 15.1x.74, RMNH. 14, Kath- mandu valley, Waterfall west Chaunica, 1400 m, 24.v.1973, RMNH. 14 (juv.), 1%, Kodari, Tibettan fron- tier, ca. 1700 m, 4.vi.73, RMNH. 14, Rhedi, 1520 m, 14.1x.74, RMNH. Coeliccia membranipes: Indonesia, Java: 14, W. Java, Salak, 800 m, 31.x.60 [1961?, different labels], ex coll. A. Heymer, RMNH. Indonesia, Nias Island: 14, Cen- tral Nias Island, Lahago, 4.11.-10.111.1896, RMNH. In- donesia, Sumatra: 14, Bambar Baru, 950 m, 12.xi.50, RMNH. 14, Fanangtalu, Ophir Distrikten, Pad. Boven- land, v.1915 [1914 ?]. 13, Fort de Kok, x.1913. 19, Moesa Kambangan, 10.1x.1927. 24, N. Sumatra, Atjeh, Sangir, 1200 m, 19.11.1937. 14, N.E. Sumatra, Sibo- langiti, 400 m, 17.xi.1950. 24, S. Sumatra, S.W. Lam- poeng, Mt Tanggamoes, 3.1.35. 24, S. Sumatra, S.W. Lampoeng, Mt Tanggamoes, 24.x11.34, RMNH. DGS: Sumatra, Wai Teboe, 500 m, 24.vi.1934. 14, Sumatra Sungai Kumbang (Kurintji), ix.1915, RMNH. 19, Su- matra, Serapai (Kurintji), vii.1915. 14, Sumatra, Ben- gkulen, Suban Ajam (Redjang), vii.1916. 14, Suban Ajam (Redjang) Bengkulen, vii.1916. 24, Weg B. Ago- eng, grens Benkoelen, Ranaumeer, 2.1.1938, RMNH. Copera marginipes: Indonesia, eastern Java region: 14, Kangean Island, 21.ix.54, Petafan, RMNH. Indone- sia, Bali: 19, 19, ca. 2 km inland from Grokgak, 8°11'S 114°47'E", 150 m asl., 11.v.1991, RMNH. Cyanocnemis aureofrons: New Guinea, West Papua: 54, 29, paratypes, Araucariakamp, 700-800 m, 8.- 31.11.1939, RMNH. All paratypes; Neth. Ind.-Amer. New Guinea Exped., RMNH. Idiocnemis leonorae: Papua New Guinea: 33, 2%, NE New Guinea, E. Highlands, Yonkie Dome, 11 mi. NE Kainantu, 1500 m, 18-20.x.1972, ex coll. Donnelly, RMNH. Indocnemis orang: SE Asia, Malayan Peninsula: 22, Malaya, Selangor, Templer Park, K. L., 12-13 mi., 21.11.1963, RMNH. 14, Malaya, S. Kedah, Bading, Badenoch Estate, 21.11.1963, RMNH. Leptocnemis cyanops: Seychelles, Mahé island: 16, banks of the Grand Bois R., 6.11.1974, RMNH. 14, Bouin, 15.11.1974 RMNH. 14, St. Louis R., 8.11.1974, RMNH. Seychelles, Praslin island: 14, Vallée de Mai, 25.v11.1965, ex coll. Pinhey '66, RMNH. Lieftinckia isabellae: 24 (1 def.), Solomon Islands, Santa Ysabel island, Maring distr., Ta Matahi, 2.vii.1960, ex BISH, RMNH. Lieftinckia kimminsi: Paratypes: 14, Solomon Islands, southern Bougainville, west Kieta, Crown Prince Range, Kokorei (=Kokuze), 900 m, 12.vi.56, RMNH. ld, 12, west Kieta, Crown Prince Range, Kokorei (=Kokuze), 9.vi.56, RMNH. Lieftinckia malaitae: Paratypes: 2¢ (1 juv., incompl.), 1% (semi-adult), Solomon Islands, Malaita Island, Da- la, 50 m, 9-4.vi.1969, RMNH. 24 (1 semi-adult), Ngwaian, 1500 ft., 10.x.1967, RMNH. Dirk GASSMANN: The Phylogeny of Southeast Asian and Indo-Pacific Calicnemiinae 79 Lieftinckia ramosa: Paratype: 14, Solomon Islands, Bougainville island, RMNH. Lieftinckia salomonis: 25, 14, Solomon Islands, Gua- dalcanal island, Gold Ridge, 30.v1.1956, BMNH (no. BM 1956-712). 19, Sornolio R. [?], 30.vi.1956, BMNH (no. BM 1956-712). 12, Tenaligi R., 25.ix.65, "Dicht bos; juv., white", RMNH. Lochmaeocnemis malacodora: 2, New Guinea, West Papua, Araucaria Camp, 800 m, 111.1939. 24, Lower Mist Camp, 1400-1600 m, 1.1939. 6%, 29, Rattan Amer. New Guinea Exped., RMNH. Mesocnemis singularis: East Africa: 34, Rusinga Is- land, Lake Victoria, 111.1950, RMNH. 3%, Jinja, Uganda, vi. 1949, RMNH. 1%, Victoria Falls, 29.1x.1961, RMNH. 24, Ghana (Eastern Region), 31 km NW Koforidua: Birim R. at Bunso Waterworks. 6°16.0'N 0°28.3'W, 10.v.2000, RMNH. Paracnemis alluaudi: Madagascar: 14, Fort-Dauphin, 17.11.58, RMNH. 1%, Soanierana-Ivongo, 10.xi.57, RMNH. Paramecocnemis erythrostigma: New Guinea, West Papua: 4¢ paratypes, Hollandia, iv.-vi.1931 RMNH. 34, Hollandia, vii.1938, Dutch-Amer. New Guinea Ex- ped. 1938-1939, RMNH. Platycnemis pennipes: Europe, France: 14, Dampi- erre, bij Parijs, 19.v1.1966, RMNH. Europe, Turkey: 13, 19, Mugla, ca. 7 km SW of Kóycegiz: Lamnan Cayi, near bridge SW of Hamitkóy, 5 m asl, 1.v1.2000, RMNH. Rhyacocnemis sufficiens: Papua New Guinea, D'En- trecasteaux Islands: | paratype, Goodenough I., 1600 m — Camp (4th Archbold New Guinea Exped.), 12.x.1953, RMNH. 13, Normanby I., Camp 2 (Sth Archbold New Guinea Exped.), Mt Pabinama, 820 m, 8.v.1956, RMNH. Archbold Exped. Risiocnemis appendiculata: Philippines, Mindanao Is- land, Misamis Oriental Province: 34, 1%, Bal-ason, 2-3.1v.1960, RMNH. Philippines, Mindanao Island, South Cotabato Province: 24, Koronadal, Barrio 8, 4.11.1994, RMNH. 14, Mt Matutum, 500-700 m, 16- 19.1x.1993, RMNH. Philippines, Mindanao Island, Surigao del Sur Province: 14 (def.), Cotabato, Parang, 13.1.1954, RMNH. 34, Tandag, Hitaub Creek, 500-600 m, 16-19.iv.1995, RMNH. Philippines, Leyte Island: 14, Ormoc, 18.xi.1966, RMNH. Risiocnemis arator: Philippines, Luzon Island, Ifugao Province: 14, Jacmal Bunhian, 24 km E Mayoyao, 800-1000 m, 7-8.iv.1967, RMNH. Philippines, Luzon Island, Nueva Viscaya Province: 13, 19, Sta Fe, Dal- ton Pass, 800-1200 m, 25-30.v1.1989, RMNH. Risiocnemis asahinai: Philippines, Mindoro Island: Id, Barrio Luyang, Halcon Mts., 360-500 m, 14./17.v.1991, RMNH. 34, Mindoro Oriental Province, Calapan, Comonal, Mt Tarugin, ca. 350 m, 21./29.vi. & 17./26.vii.1990, RMNH. 23, 4%, Mt Halcon, 1000- 1500 m, 2-20.v.1994, RMNH. Risiocnemis confusa: Philippines, Catanduanes Is- land: 14, Gigmoto, San Pedro, Tongao Creeks, 400- 500 m, 20-30.v1.1996, RMNH. Risiocnemis elegans: Philippines, Luzon, Aurora Province: 73, 17, Dinalungan, Mt Anaguao, Alebit R. area, 600-900 m, 9-14.111.1997, RMNH. 24, Dinalun- gan, Mt Anaguao, Bolawan R., 800-1000 m, 11.11.1997, RMNH. Risiocnemis erythrura: Philippines, Mindanao Island, Surigao Province: 1, L. Mainit, 30.x1.1989, RMNH. Philippines, Mindanao Island, Surigao del Sur Prov- ince: 13, 12, Carmen, Upper Tandag R., km 9 Lanang Line, 500 m, 24.iv.1995, RMNH. 14, Tago, Meme R., 100-300 m, 12-18.v1.1996, RMNH. Risiocnemis gracilis: Philippines, Luzon Island, Nueva Ecija Province: 1, 1%, Caranglan, Batching R., 700-850 m, 1 1.vi.1991, RMNH. Philippines, Luzon Island, Nueva Viscaya Province: 14, 1%, Sta Fe, Dal- ton Pass area, 900 m, 8-17. vi11.1991, RMNH. Risiocnemis kiautai: Philippines, Sibuyan Island: 1 paratype, Magdiwang, Katingas, 29.111.1987, RMNH. 14, paratype, Magdiwang, Tampayan, Camp Ga-ong, 80-150 m, 1-12.1v.1987, RMNH. Risiocnemis laguna: Philippines, Luzon Island, Laguna Province: 14, 19, paratypes, Paete, 29.vi.1916, SMFD. Risiocnemis moroensis: Philippines, Mindanao Island, Bukidnon Province: 1, Philippines, Mindanao, Bukidnon, Mt Imbayo, 30.viii.1988, RMNH. 14, 2%, Mt Kalatungan, Talakag, Brgy Mebadiang, Magamana- son Creek, Sitio Olayan, 1000-1100 m, 5-18.v111.1995, RMNH. 1%, Philippines, Mindanao Id, Bukidnon, Ma- badiang, Olayan, Mt Kalatungan, Muntian Creek, 1200- 1400 m, 24./25.x1.1995, RMNH. 2%, Philippines, Min- danao Id, Bukidnon, Mebadiang, Dumatap, Mt Kalatun- gan, Mansabilan Creek, 1000-1300 m, 24.x1.1995, RMNH. 14, Mt Katanglad, Impasugong, Brgy Impalu- tao, Gantongan Creek, 800-900 m, 19-29.vili.1995, RMNH. Risiocnemis praeusta: Philippines, Dinagat Island: 14, Loreto, Balitbiton, Mt Canbinlio, vii.1989, RMNH. Philippines, Leyte Island: 17, (Mt Balocaue ?). 20.1.1989, RMNH. Philippines, Panaon Island: 1 San Francisco, Gabing Gamay, big river, x.1988, RMNH. 1d, 12, San Francisco, Batong Lapad. vill. 1988, RMNH. 80 Bonner zoologische Beitráge 53 (2004) Risiocnemis pulchra: Philippines, Luzon Island, Bataan Province:1Í, paratype, Luzon, Limay, 21.x.1913, SMED. Philippines, Luzon Island, Zam- bales Province: 6¢, 22, Masinloc, Mt Coto, Tal-tal, 400-700 m, 1-8.1v.1997, RMNH. Risiocnemis rolandmuelleri: Philippines, Negros Is- land: 94, 1%, Silay, Patag, Cuyong R., Dumalabdab Falls, 700-750 m, 22.v.1996, RMNH. Risiocnemis seidenschwarzi: Philippines, Cebu Island: 44, paratypes, Tabunan, 500 m, 19.xi.1998 & 9.11.1999, RMNH. Risiocnemis serrata: Philippines, Catanduanes Island: 34, 12, Gigmoto, San Pedro, Simohe Creeks, 300-500 m, 20-30.v1.1996, RMNH. Philippines, Luzon, Bataan Province: 23 (1 def.), Los Baños, 15.v.1977, RMNH. 22, Los Baños, 50 m, 28.v.1949, RMNH. Risiocnemis varians: Philippines, Luzon Island, Nueva Viscaya Province: 14, Dalton Pass, Sta Fe, Atbo, 500-900 m, 13.11.1989, RMNH. 14, Dalton Pass, Sta Fe, Torner, 500-900 m, 12.11.1989, paratype, RMNH. 2%, Dalton Pass, Sta Fe, Zigsag, 500-900 m, 16.11.1989, paratype, RMNH. 33, 39, Sta Fe, Barrio Lohong, Mounts east of Dalton Pass, 900-1000 m, 10- 23.1.1993, RMNH. 24, Sta Fe, Dalton Pass, 800-1200 m, 25./30.v1.1989 & 5./18.v1.1990, Paratype, RMNH. 19, Sta Fe, Atbo R., 550-800 m, 10.vi.1991, RMNH. ld, Sta Fe, Dalton Pass area, 900 m, 8./17.viii.1991, RMNH. Philippines, Luzon Island, Quirino Province: 19, 19, Maddela, Sulong R., 500-650 m, 26./27.iv.1991, RMNH. Salomocnemis gerdae: Solomon Islands, Guadalcanal: 13, Komugelea, forest, 1200 ft., 22.1x.1965, RMNH. Y, Komugelea, Guadalcanal, open brook, 1200 ft., 22.1x.1965, RMNH. Stenocnemis pachystigma: Western Africa, SW Cam- eroon: 14, Meme, Nyasoso, Mt Kupe, R. Nyesosoh, water catchment, 920 m, 30.11.1997, RMNH. 19, Tom- bel, Nyasoso, Mt Kupe, Shrike Trail, 920 m, 5.iv.1995, RMNH. Thaumatagrion funereum: Northern New Guinea: 64, paratypes, Hollandia, iii.1931, RMNH. 14, 19, Hollan- dia, 24.1.1933, RMNH. Torrenticnemis filicornis: Central northern New Guinea: 54, paratypes, Lower Mist Camp, 1400-1600 m, 1.1939, RMNH. 54, 19, paratypes, Sigi Camp, 1500 m, 11.1939. Neth. Ind. - Amer. New Guinea Exped., RMNH. Bonner zoologische Beitráge Band 53 (2004) | Heft 1/2 | Seiten 81-97 Neue Zygaena-Taxa aus Siidosteuropa, Vorder- und Zentralasien (Lepidoptera: Zygaenidae)' Fúr Clas, dem wir alle viel verdanken Axel HOFMANN, Breisach-Hochstetten Abstract. Several new taxa of the genus Zygaena Fabricius, 1775 are described from south-eastern Europe, Anatolia, Iran, Pakistan and Central Asia: Zygaena (Mesembrynus) rubricollis arachosica subsp. nov. from Quetta region (Paki- stan) with Z. manlia-like forewing spots. Zygaena (Mesembrynus) rubricollis shahkuhica subsp. nov. from Shah-Kuh (Iran) where it occurs syntopically with Z. manlia and Z. cacuminum. Zygaena (Mesembrynus) nocturna eberti subsp. nov. from the central Zagros range (Iran) typical with connected spots 5+6. Zygaena (Agrumenia) magiana pamirescens subsp. nov. from Kirghistan, a subspecies with yellow forewing spots thus reminiscent of Z. pamira. Zygaena (Agrume- nia) formosa molleti subsp. nov. from Bolkar Dagh (Turkey), the first high mountain population of this species, ecologi- cally and phenotypically reminiscent of Z. peschmerga. Zygaena (Agrumenia) tenhagenova sp. nov. from Iranian Kord- estan, a distinct biospecies closely related to Z. chirazica and Z. naumanni. Zygaena (Agrumenia) haberhaueri demangei subsp. nov. from north-western Iran, with red abdominal cingulum on | — 3 segments. Zygaena (Zygaena) viciae pelopis subsp. nov. from Peloponnesos peninsula (Greece) reminiscent of Z. filipendulae and Z. vicae laphria from the Taurus range in Turkey. Zygaena (Zygaena) filipendulae wiegelorum subsp. nov. from the province of Kars (Turkey) with red confluent forewing spots. Zygaena (Zygaena) lonicerae pseudangelicae subsp. nov. from Lake Van region (Turkey) phenotypically similar to Z. angelicae and Z. trifolii. Key words: Zygaena chirazica, Z. filipendulae wiegelorum subsp. nov., Z. formosa molleti subsp. nov., Z. haberhaueri demangei subsp. nov., Z. lonicerae pseudangelicae subsp. nov., Z. magiana pamirescens subsp. nov., Z. naumanni, Z. nocturna eberti subsp. nov., Z. rubricollis arachosica subsp. nov., Z. r. shahkuhica subsp. nov., Z. tenhagenova sp. nov., Bonn, Juni 2005 Z. viciae pelopis subsp. Nov. 1. EINLEITUNG Bei der Abfassung des ,,Systematic Catalogue of the Zygaeninae“ (HOFMANN & TREMEWAN 1996) zeigte sich bereits, dass einige sehr auffallige Populations- gruppen auch aus relativ gut bearbeiteten Regionen wie der Türkei und Griechenland bislang noch unbenannt geblieben sind. Die Beschreibung dieser Taxa wird hiermit nachgeholt. Des weiteren werden hier einige be- sonders markante Neuentdeckungen der vergangenen Jahre aus Zentralasien, dem Iran und aus Pakistan na- mentlich verfügbar gemacht. Eigene Aufsammlungen der Jahre 2001 bis 2004 im Iran (zusammen mit A. KALLIES, J. U. MEINEKE , W. G. TREMEWAN) sowie Beobachtungen von T. KEIL und G. TARMANN haben den Verdacht erhártet, dass unter Zy- gaena (Mesembrynus) rubricollis Hampson, 1900 bis- lang mehr als eine einzige Biospezies zusammengefasst wurde (HOFMANN & TREMEWAN 2003; KEIL 2003). Gleiches scheint für Zygaena (Mesembrynus) seitzi Reiss, 1938, Zygaena (Mesembrynus) manlia Lederer, 1870 und Zygaena (Agrumenia) rosinae Korb, 1903 zu- zutreffen. HOFMANN & TREMEWAN (2003) haben neu- erdings einen ersten vagen Versuch unternommen, die komplizierten taxonomischen Verháltnisse zu entwirren. 1 Clas Michael Naumann zu Königsbrück (26.06.1939 — 15.02.2004) zum Gedenken Als Ergebnis wurden Z. aisha, Z. nocturna und Z. fredi wieder in den Artstatus erhoben. Unklar bleiben aber nach wie vor zahlreiche Zuordnungen (kermanensis, gashgai, askarii, taftanica, escaleraiana, tenhageni etc.), und es bedarf noch intensiver Feldarbeit sowie umfassender biologischer und taxonomischer Untersu- chungen, bis die Verwandtschaftsverháltnisse insbeson- dere der irano-afghanischen Mesembrynus-Arten eini- germaßen befriedigend erklärt werden können. Auf die noch ungenügend begründete Taxonomie bei Z. afgha- na, Z. rosinae/sengana hat KEIL (2003) hingewiesen und Z. sengana wieder als bona species abgetrennt. Ein Artenkomplex könnte sich auch hinter Z. chirazica/- naumanni verbergen. Es ist bekanntermaßen bei allo- patrischen Taxa oft schwer zu entscheiden, ob es sich um eine einzige außergewöhnlich differenzierte, polyty- pische Art oder um mehrere engverwandte isolierte Bio- spezies handelt. Die geographische Isoliertheit und der bisweilen hohe Grad an Verschiedenheit z. B. zwischen Z. naumanni, Z. chirazica chirazica und Z. chirazica eckweileri, aber auch die Unterschiede zu anderen bis- lang noch nicht beschriebenen Populationen bzw.Taxa (s. Z. tenhagenova n. sp. , Populationen vom Zarde-Kuh und von Delijan) könnten bereits als Argumente für dis- tinkte Biospecies angeführt werden. Es muss jedoch be- tont werden, dass diese „Artverschiedenheit“ bisher eher schwach begründet wurde. Dies gilt für Z. naumanni aus der Region um Chonsar und Fereydun Shahr, aber auch 82 Bonner zoologische Beitráge 53 (2004) fiir die hier neu beschriebene Z. tenhagenova n. sp. aus iranisch Kurdistan. Eine Beschreibung als Unterart von Z. chirazica wäre nur möglich gewesen, wenn man Z. naumanni, die geographisch dazwischen siedelt, zuvor eingezogen hätte. Die nördlichste Population unweit Delijan zeigt bereits habituelle Annäherung an Z. ha- berhaueri, so dass evtl. auch diese Art in die Untersu- chung mit einbezogen werden müsste. Geeignete Feld- studien (z. B. Anlockversuche mit virginen Weibchen, Aufsammlungen in bislang unbesammelten Kontaktbe- reichen), Zucht- und Kreuzungsexperimente und die ge- zielte Suche nach weiteren taxonomischen Merkmalen (z. B. GU-Serienuntersuchungen) werden zukünftige Aufgabenbereiche darstellen, verbunden mit der Hoff- nung, weitere brauchbare Hinweisen zu erhalten, ob un- sere heutigen Anschauungen richtig waren oder korri- giert werden müssen. Wichtige Argumente zur Rekonstruktion stammesge- schichtlicher Verwandtschaftsverhältnisse wird uns die Analyse geeigneter DNA-Abschnitte liefern (O. NIE- HUIS, Bonn, in Vorbereitung). Entscheidungen auf Art- niveau (und tiefer), insbesondere die Bewertung allo- patrischer Taxa (Allo- oder Konspezifität), wird dem Taxonomen aber auch von diesem Wissenschaftszweig mit Sicherheit nicht abgenommen werden können. Es wäre vermessen zu erwarten, dass genetische Schwel- lenwerte gefunden würden oder definierbar wären, die eine solche Entscheidung fortan dem wertneutralen Computer überließen. 2. BESCHREIBUNG NEUER TAXA Die nachfolgenden Unterarten werden hier vorerst den derzeit noch gültigen Taxa zugeordnet. Zygaena (Mesembrynus) rubricollis arachosica subsp. nov. (Figs. 1 — 3) Zygaena (Mesembrynus) rubricollis Hampson, 1900, Journal of the Bombay natural History Society 13: 224, pl. B, fig. 9 Derivatio nominis: Nach der altpersischen Satrapie „Arachosien“. +: „Pakistan, Baluchistan, Quetta/Urak, 2400 — 2700 m, 24. — 26.V.83, leg. ECKWEILER“, coll. A. HOFMANN (später deponiert in coll. Staatliches Mu- seum für Naturkunde Karlsruhe, SMNK). Holotypus, : Paratypen: 1 ¢, 2 YY, Daten wie Holotypus, coll. A. HOFMANN; 3 44, 3 2%, Daten wie Holotypus, jedoch 23. — 24.V. 1979, coll. A. HOFMANN. Einzelne Belegtie- re (Daten wie Holotypus) in coll. C. M. NAUMANN und coll. T. WITT (via coll. G. REISS). Zum Vergleich liegen Topo- und Paratypen der aus Af- ghanistan beschriebenen Unterarten (Z. r. afghanica Reiss, 1940; Z. r. flavicola Naumann, 1969; Z. r. kabu- lica Naumann, 1974; Z. r. nasukmiri Naumann, 1974) sowie aktuelle Nachfänge der nominotypischen Unterart von Z. rubricollis aus Nord-Pakistan (Birmoghlash, Chitral: leg. W. ECKWEILER 1983, leg. J.-C. WEISS 1996, leg. M. NICOLLE 1997) vor. Neuerdings wurde die Art auch weiter östlich im Nanga Parbat-Gebiet gefun- den (DESSE 2003). Alle diese Populationen unterschei- den sich von Z. r. arachosica subsp. nov. durch ihre un- verkennbare Tendenz zur Reduktion von Fleck 6 (insbesondere im costalen Bereich) sowie ein wesentlich kleineres Fleckenpaar 3+4. Bei der Nominatunterart, bei Z. r. afghanica und Z. r. nasukmiri ist dieses nicht selten sogar durch eine schwarze Ader deutlich in zwei separa- te Flecken aufgelöst. Z. r. arachosica subsp. nov. besitzt stets einen vollentwickelten “Nierenfleck” (= Fleck 6), der sich mit seinem oberen und unteren Ende Fleck 5 annähert und dadurch eine geknickte Form erhält. Fleck 5 ist meist von dreieckiger Gestalt, ebenfalls groß wie auch die Basisflecken 1+2 und das Fleckenpaar 3+4, die jeweils zu einem einzigen Flecken zusammenfließen. Insgesamt sind alle Flecken größer als bei den zuvor genannten Taxa, so dass eine gewisse Ähnlichkeit mit Z. manlia Lederer, 1870, entsteht. Rote Halskrause und ge- schlossenes Abdominalcingulum sind auf einem Seg- ment vorhanden. Hinterflügelumrandung schmal, am beginnenden Analfeld nur schwach verstärkt. Hinterflü- gel wie bei allen afghanischen und pakistanischen Popu- lationen ohne hyalinen Wisch. Hierdurch unterscheiden sich die afghanisch-pakistanischen Z. rubricollis-Taxa auch habituell ziemlich deutlich von weiter westlich an- schließenden und ebenfalls heute (noch?) unter Z. rubri- collis subsumierten iranischen und armenisch-südost- anatolischen Populationen. Zygaena rubricollis shahkuhica subsp. nov. (Fig. 7) (Mesembrynus) Derivatio nominis: Nach der Typenlokalität am Shah- Kuh (Prov. Mazandaran/Golestan). Holotypus, ©: „Mazandaran, Shah Kuh (Est), versant nord, 2800 — 3000 m, 15./17. 7.1998, leg. B. MOLLET“, coll. A. HOFMANN (später deponiert in coll. SMNK). Dieses interessante Einzeltier wurde von B. MOLLET zusammen mit einer kleinen Serie Z. cacuminum Chris- toph, 1877, erbeutet. Gezielte Nachsuche seither blieb erfolglos. In den darauffolgenden Jahren konnten jedoch im selben Gebiet, unweit Hadjiabad topotypische Z. manlia manlia erstmals seit mehr als 130 Jahren wieder in wenigen Exemplaren nachgefangen werden (B. MOL- LET 2000; B. MOLLET & G. FLUTCH 2001; G. EBERT, R. TRUSCH, H. ALIPANAH, E. EBRAHIMI 2003). Somit kann nun eine Verwechslung mit einer dieser beiden nahe verwandten Arten ausgeschlossen. Ich zögere nicht, die- se Population anhand eines einzigen Exemplars mit ei- nem Namen zu belegen, zumal das hier neu beschriebe- Axel HOFMANN: Neue Zygaena-Taxa aus Súdosteuropa, Vorder- und Zentralasien 83 ne Taxon sicherlich nicht zu Z. manlia oder Z. cacumi- num gehört und Z. rubricollis aus dem gesamten El- bursgebirge bislang unbenannt geblieben ist (NAUMANN et al. 1984: 76 — als Z. araxis — HOFMANN 2000). Wei- ter westlich, aus dem zentralen Elburs, liegen zum Ver- gleich einige wenige Tiere von Z. rubricollis (s. HOF- MANN 2000: 227, leg. J. KLIR) vor. Sie sind jedoch abweichend gezeichnet, weshalb sıe nicht zur Typense- rie gezogen werden können. Insbesondere die Vorder- flügelflecken sind größer, die Nierenmakel ist geknickt und das Basisfleckenpaar 1+2 schließt fast senkrecht zum Innenrand hin ab. Durch Reduktion von Fleck 2a ist bei Z. r. shahkuhica subsp. nov. der Basisfleck am Innenrand deutlich eingebuchtet. Fleck 5 und 6 sind völ- lig freistehend, die Nierenmakel steht fast senkrecht und besitzt keine „birnenförmige“ Verdickung am unteren Ende. Der sichtbare Bereich der schwarz beschuppten Valven ist mit roten Schuppen durchsetzt. In Linneana Belgica XVIII: 229 (fig. 17) wurde das Tier bereits ab- gebildet. Die Unterschiede zu Z. manlia als auch zu Z. cacuminum und allen bekannten Z. rubricollis- Unterarten sind signifikant. Position und Größe der Fle- cken 5 und 6 erinnern am ehesten an die von NAUMANN & NAUMANN (1980) als Unterart von Z. manlia be- schriebene, inzwischen zur Art aufgewertete Z. aisha (HOFMANN & TREMEWAN 2003) aus Kerman. Hierbei scheint es sich jedoch um eine ökologisch und mor- phologisch wohl differenzierte endemische Biospezies aus den isolierten Gebirgen der Provinz Kerman zu han- deln. Als gesichert kann nun angesehen werden, daß mindes- tens drei verschiedene Arten aus der manlia-cuvieri- Gruppe im Elbursgebirge vorkommen. Erschwert wird die Bewertung der einzelnen Vorkommen durch überra- schend spärliche Nachweise. Während im Zagrosgebir- ge Z. manlia oder Z. fredi an manchen Standorten in Anzahl beobachtet werden können, sind aus dem Elburs immer nur Einzeltiere bekannt geworden. Zygaena (Mesembrynus) nocturna eberti subsp. nov. (Figs. 13 - 15) Zygaena (Mesembrynus) nocturna Ebert, 1974, Beiträge zur naturkundlichen Forschung in Südwestdeutschland 33: 164, figs. 2-6 Derivatio nominis: Nach dem Autor des nominellen Taxon „Zygaena nocturna” und Entdecker dieser Popu- lation am Osturan-Kuh, Günter EBERT. In Dankbarkeit für langjährige Freundschaft. Dag, 2600 m, e.p.: 7. 5. — 27. 5. 2000, A. HOFMANN cult., coll. A. HOFMANN“ (später deponiert in coll. SMNK). Paratypen: 9 ¿4,5 YY, Daten wie Holotypus, coll. A. HOFMANN; einzelne Belegtiere (Daten wie Holotypus) in coll. J.-M. DESSE, E. DROUET, T. KEIL, J. KLIR, C. M. Holotypus, ©: „Iran, e.o., Lorestan, Dorud, Golgolab NAUMANN, M. NICOLLE, W. G. TREMEWAN, T. W!IT1 (via G. REISS); 1 4, Prov. Lorestan, Dorud 22 km E. Dobastane 8 km E, 2300 — 2500 m, L. F., leg. A. Hol MANN & P. KAUTT, coll. A. HOFMANN; 2 , Ibidem, jedoch e.l., e.p.: 3.7.1997, coll. A. HOFMANN; 1 2, W- IRAN, Lorestan, Dorud 5 km SE, Saravand, "Kohyeh". 2300 m, 29. — 30. 7. 1975, L. F., leg. G. EBERT & H. FALKNER, coll. SMNK; 4 dd, W-IRAN, Lorestan, Do- rud, Paß S Darra-che-Gahar, "Partsche Kabud", 1. — 3. 8.1975, L. F., leg. G. EBERT & H. FALKNER, coll. SMNK. Patagia und geschlossenes Abdominalcingulum kráftig rot. Hinterflúgel zart rot, dúnn beschuppt, an der Wurzel glasig, schmal schwarz umrandet ohne nennenswerte Verdickung am Apex. Vorderfliigelflecken gelb, ver- gleichsweise groß, 1+2 und 3+4 flächig zu zwei Fle- ckenpaaren verbunden. Insbesondere durch die starke Annáherung von Fleck 5 an 6 unterscheiden sich die Vorkommen am Osturan-Kuh (Lorestan) recht auffallig von der nominotypischen Unterart im Kuh-e Dena Massiv (Boyer Ahmad va Kuhgiluye). Wahrend bei die- sen Populationen aus der Umgebung von Sisakht, von Ardekan, vom Gardaneh-ye Meymand oder von Chenar Mahmoudi die Flecken 5 und 6 stets deutlich durch eine breite schwarze Fláche getrennt sind und somit dem Zeichnungsmuster von Z. manlia entsprechen, tendieren bei Tieren aus der Umgebung von Dorud diese beiden Endmakeln zur Ausbildung eines einzigen zusammen- geflossenen Beilflecks (ca. 50 %). Nur weniger als 20 % haben oben und unten getrennte Flecken: aber auch bei diesen Tieren sind die Flecken dann stárker angenáhert als bei topotypischen Tieren. Gelegentlich sind die Fle- cken im unteren Bereich zusammenhángend, nach oben noch durch eine dünne „kommaförmige“ Einbuchtung getrennt. Diese Zeichnungsmerkmale treffen auch auf Tiere vom Kamaran-Tal (Fereydun Shahr S) zu, nicht jedoch die Farbung der Hinterfliigel dieser Population. Bei ca. einem Drittel der vorliegenden Tiere (n = 18) wird die rote Grundfarbe durch orange oder gar gelbe Schuppen ersetzt. Es handelt sich hierbei um eine phá- notypisch bereits abweichende, isolierte Population, die mehrheitlich ihre subspezifische Zugehörigkeit zu eberti subsp. nov. aber noch klar erkennen lässt. Charakteristisch für Z. nocturna (und Z. seitzi REISS, 1938) ist bekanntermaßen die konstant unterschiedliche Färbung von Vorder- (gelb) und Hinterflügeln (rot). Während nun am Osturan-Kuh in Höhenlagen zwischen 2200 und 3000 m Z. nocturna eberti subsp. nov. mono- morph in dieser Form auftritt, scheint in tieferen Lagen (1600 — 2000 m) des selben Massivs diese gelb/rot ge- färbte Form in polychromatischen Mischpopulationen mit gelb/rot, orange/rot oder rein rot gefärbte Individuen überzugehen. Zuchtergebnisse im Freiland aufgesam- melter L,, L> — Ráupchen (1999) sowie mehrerer Eige- lege (1997) vom Duschderaz-Paß bei Dorud (HOFMANN 84 Bonner zoologische Beitrage 53 (2004) 2000: plate III, figs. 22 - 27, 192 ff) lieferten folgende Resultate (1998 — 2000): [n = 35] manlia-artig cuvieri-artig gelb/rot 1/0 0/1 (nocturna-artig) orange/rot 0/2 rot/rot 6/3 14/8 Dabei gilt es zu bedenken, dass aus ein- und demselben Eigelege (1997) neben manlia- und cuvieri-artigen Indi- viduen auch ein Tier mit gelben Vorderflügeln hervor- ging (HOFMANN 2000: plate II, fig. 14). Entweder findet hier ım geographischen Kontaktbereich von man- lia/cuvieri mit Z. nocturna eine phänoptypisch-konver- gente Annäherung durch gleichgerichteten Selektions- druck statt oder es handelt sich hierbei um Vorkommen von Hybriden zwischen Z. nocturna und Z. man- lia/cuvieri. Hierfür könnte u.a. die sehr variable, von Z. nocturna abweichende, an Z. manlia/cuvieri aber bis- weilen stark angenäherte Raupenzeichnung sprechen (HOFMANN 2000a: 198, fig. 2; HOFMANN 2000b: 228, fig. 8b). Auch bestárken mehrere fertile Kreuzungsexpe- rimente (s. HOFMANN & TREMEWAN 2003: 10) diesen Verdacht. Neben den unterschiedlichen Raupenphána wird eme befriedigende taxonomische Interpretation auch durch verschiedene Kokonfarben (weiß bei Z. noc- turna, gelb bei durch den Zuchten vom Duschderaz- Paß) erschwert. Eine vergleichbar polymorphe ,,Popula- tion“ fand T. KEIL 2001 (pers. Mitt.) etwas weiter nord- östlich zwischen Dorud und Khoramabad (1800 m), die allerdings phánotypisch stärker an Z. cuvieri/Z. tamara angenáhert 1st. Zygaena (Agrumenia) magiana pamirescens subsp. nov. (Figs. 16 — 18) Zygaena (Agrumenia) magiana Staudinger, 1889, Stettiner en- tomologische Zeitung 50: 23 Derivatio nominis: Die Tiere zeigen starke habituelle Annáherung an Zygaena (Agrumenia) pamira Sheljuzh- ko, 1919. Holotypus, ©: „Kirghizstan, Alai mountains, Gaumich pass, H = 3.800 m, 26.07.1999, coll. A. HOFMANN“ (später deponiert in coll. SMNK). Paratypen: 31 94, 9 YY, Daten wie Holotypus, coll. A. HOFMANN. Abgesehen von einer meist nur angedeuteten rótlichen Patagia ist der Kórper zeichnungslos matt schwarz. Die Vorderfliigelflecken sind wohl entwickelt, Fleck 1 und 5 gelb umsáumt, rótlich gekernt, selten vóllig gelb, Fleck 2 — 4 und 6 gelb, meist ohne Spuren roter Schuppen. Hinterflügel rot von schwarzer, im Analfeld ausfransen- der Linie umgrenzt, am Apex verbreitert. Ein ( lässt magiana-typische Zeichnung mit roten Flecken 1+2, 3 und 5 erkennen, auch Fleck 6 und 4 sind bei diesem Tier leicht rot bestäubt, ein Y zeigt alle Flecken rot gekernt. Der Basisfleck (1+2) ist durch den schwarz beschuppten Stamm der Radialader getrennt. Fleck 2a ist nicht ent- wickelt. Von allen bekannten Z. magiana-Unterarten ist Z. m. pamirescens subsp. nov. durch ihre dominierende gelbe Vorderflügelfleckenfarbe leicht zu unterschieden. Sie kann weder mit den verdunkelten Populationen vom Kumbel-Paß (Z. m. kumbelica Tremewan & Naumann, 1989) noch mit den hyalinen nominotypischen Tieren oder mit den rotausgefärbten Unterarten (Z. m. alaudina Tremewan & Naumann, 1989, Z. m. alaica Holik & Sheljuzhko, 1956, Z. m. kohistana Grum-Grshimailo, 1893) verwechselt werden. Habituell erinnert die Mehr- heit der Tiere stärker an Z. pamira. Beide „Arten“ sind streng allopatrische Vertreter der Orealfauna Zentral- asıens. Zukünftige Aufsammlungen im tadschikischen Kontaktbereich könnten mehr Klarheit bringen, wie beide Taxa zu bewerten sind. Syntopes Auftreten zweier so eng verwandter und ökologisch nahestehender Taxa ist jedoch kaum zu erwarten. Sollte sich eine klinale phänotypische Annäherung ergeben, ist von einer ein- zigen Biospezies auszugehen. Zuchtexperimente (Kreu- zungsversuche, Anlockversuche etc.) stehen noch aus. Aus der selben Region (Kollektorny Range, Gaumich pass, 4.8.1998) liegt eine Serie SI, Y P von Z. cocan- dica Erschoff, 1874, vor. Auch diese Tiere sind durch kräftigere Farbanlagen, rote Ausfärbung der Basisfle- cken 1+2 zu einem einzigen großen Makel und stets vorhandenes rotes Abdominalcingulum mit Z. magiana pamirescens subsp. nov. nicht verwechselbar. W. ECK- WEILER fing 1993 in der östlichen Turkestanski-Kette (Kirgistan, Osh oblast, Ljailjak — Aksu valley, 3100 — 3500 m) einige Einzeltiere, die vermutlich zu Z. magia- na gehören und zu pamirescens subsp. nov. gezogen werden könnten. Im Gegensatz zu den Tieren vom Gaumich-Pass besitzen diese 2 44, 1 Y jedoch ein Ab- dominalcingulum sowie einen zusammengeflossenen Basisfleck (1+2), wobei die Rotfärbung von Fleck 1 auch auf Fleck 2 übergreift. Fleck 2a ist angedeutet. Zygaena (Agrumenia) formosa molleti subsp. nov. (Figs. 25 — 27) Zygaena (Agrumenia) formosa Herrich-Scháffer, 1852, Syste- matische Bearbeitung der Schmetterlinge von Europa 6: 45; 1851, ibidem 2: pl. 14, fig. 99 [non-binominal] Derivatio nominis: Nach Bernard MOLLET (Gometz-le- Chátel). „Merci“ für wertvolles Belegmaterial und zahl- reiche Zuchtmitbringsel. Holotypus, : „Turkiye, prov. Nigde, Ulukisla, 2500 — 3000 m, 4.8.1995, leg. B. MOLLET“, coll. A. HOFMANN | Axel HOFMANN: Neue Zygaena-Taxa aus Siidosteuropa, Vorder- und Zentralasien 8. (spáter deponiert in coll. Staatliches Museum fiir Natur- kunde Karlsruhe). Paratypen: 5 44,4 YY, Daten wie Holotypus, coll. A. HOFMANN; | 9, „Bolkar daglari N, 2000 — 3000 m, (Nigde) — Turkquie, 19-20 juillet 1998, Frédéric CAR- BONELL leg.“, coll. A. HOFMANN; 50 $4, 30 9 9, ,,Sou- thern Turkey, Prov. Konya, Bolkar dagları, Aydos dagi, NE side, 37°20’ N, 34°24’ E, 2700 m, 23.V11.1998, C. u. A. NAUMANN leg.”, coll. C. M. NAUMANN; 1 3, 2 29, „Turkey, Nigde, 3100 m, Bolkar daglari, N side, SW Maden, 28.VII.1994, H. v. OORSCHOT, H. v.d. BRINK, H. v.d. POORTEN, W. DE PRINS [leg.]”, coll. C. M. NAUMANN; ibidem, 1 Y, 30.V11.1997, W. DE PRINS, A. OLIVIER, H. v.d. POORTEN, [leg.]”, coll. C. M. NAU- MANN. Habituell und ökologisch scheint Z. f. molleti subsp. nov. zwischen Z. formosa Herrich-Schäffer, 1852 und Z. peschmerga Eckweiler & Görgner, 1981 zu stehen. Insbesondere das tristere Rot und die verkleinerten Vor- derfliigelflecken sowie das Fehlen von Fleck 2a erin- nern stark an Z. peschmerga. Der Nierenfleck tendiert wesentlich stárker zur Loslósung von Fleck 5 als dies bei anderen Z. formosa-Populationen festzustellen ist. Die Flecken sind klein, insgesamt jedoch größer als bei Z. peschmerga, der Abstand zwischen dem Basisfleck 1+2 und Fleck 3 ist flachiger, 3+4 sind fast immer durch die Weißumrandung verbunden, ebenso kontaktiert Fleck 5 mit dem Fleckenpaar 3+4. Die Basisflecken 1+2 sind zu einer einzigen Makel verschmolzen, die im Ge- gensatz zu Z. peschmerga auch nicht andeutungsweise durch schwarze Schuppen auf der Radialader getrennt ist. Die Patagia ist nur bei den Y Y rötlich, bei den 44 weißlich-grau oder bräunlich. 2 Y Y zeigen oberseits ein schwach ausgeprägtes einfaches Abdominalcingulum, bei den anderen beiden Y Y ist der Gürtel nur unterseits in Spuren zu erkennen. Die 4 sind uncinguliert. Das Vorkommen einer gut differenzierten hochmontan- alpinen formosa-Population in der Dornpolsterstufe am Bolkar Dag darf durchaus als überraschend bezeichnet werden. Im selben Gebiet (Ulukisla 11 km E) fliegt nämlich zur gleichen Zeit auf 1100 — 1200 m Höhe eine völlig normal aussehende Z. formosa-Population mit großen leuchtend roten Flecken, roter Patagia, zwei- bis dreifachem Abdominalcingulum und wohl entwickeltem Fleck 2a. Auch das Auffinden einer weiteren — pháno- typisch fast identischen — Hochgebirgspopulation mehr als 500 km (Luftlinie) weiter nordóstlich in der Prov. Erzincan ( „Turquie, Erzincan, Mercan Daglari, N 39555. -E 39°29". 3100 .m, 27.VII.. 1999, Dominique Dumont leg.”, 1 $, 1 Y, weiteres Belegmaterial in coll. D. DUMONT) könnte bei besserer Kenntnis der 6kolo- gisch-biologischen Eigentiimlichkeiten Anlass sein, die Eigenständigkeit bzw. Zugehörigkeit von Z. f. molleti subsp. nov. neu zu diskutieren. Klärungsbedarf besteht weiter hinsichtlich Z. formosa kotzschi Reiss, 1935 Diese Unterart wurde aus der Geröllschuttzone nahe der armenischen Grenze oberhalb 3000 m beschrieben. Sie zeigt Fleckenreduktion, lässt aber noch wesentlich kla- rer (rotes Cingulum, rote Patagia etc.) ihre Zugehörig- keit zu Z. formosa erkennen. / Z. formosa molleti subsp. nov. (2600 — 3000 m) ist am Bolkar Dag etwas höher eingenischt als Zygaena (Zy- gaena) loti senilis Burgeff, 1914 (TREMEWAN & TAR- MANN 2003) und die zeitgleich fliegende Adscita capi- talis (Staudinger, 1879), die 2600 m Höhe nicht wesentlich zu überschreiten scheint (B. MOLLET, pers. Mitt.). Die Falter fliegen auffallend niedrig über dem Grund, eine Anpassung an die häufigen, teilweise recht heftigen Winde. Als Raupennahrungspflanze konnte ei- ne rotblühende, flachliegende bis kriechende, nicht- stachelige Fabaceae ausgemacht werden. Eine Eiablage in die Mittelrispe der auffällig stark behaarten Fieder- blätter konnte von B. MOLLET beobachtet werden. Zygaena (Agrumenia) tenhagenova sp. nov. (Figs. 31, 32) Derivatio nominis: Dem Entdecker Dr. Wolfgang TEN HAGEN in Anerkennung seiner verdienstvollen Feldar- beit (1996 — 2003) fiir die Erforschung der iranischen Zygaenenfauna gewidmet. Das Art-Epitheton ist ein Substantiv in Apposition, da der Name fenhageni bereits fiir Zygaena rubricollis tenhageni Hofmann & Treme- wan, 2003 vergeben ist. Holotypus, ©: „Iran, Kordestan, Paß NE Baneh, (Ost- seite); 1900 — 2100 m, 16. VI. 2001, leg. [W.] TEN HA- GEN “, coll. A. HOFMANN (spáter deponiert in coll. SMNK). Paratypus: 1 ©, Daten wie Holotypus. coll. A. HOF- MANN. Y noch unbekannt. Die habituelle Ubereinstimmung beider Tiere zeigt, dass hier eine eigenständige, zweifelsfrei beschreibenswerte Population vorliegt. Einige Merkmale und Merkmals- kombinationen sind als taxonomische Besonderheiten hervorzuheben: Gelbe Halskrause (Patagia) bei rotem Abdominalcingulum (auf zwei Segmenten), hell-ocker- gelbe Beine, Vorderfliigelflecken 2, 3, 4 und 6 gelb, Fleck 1 entlang der Costa rot, Fleck 5 rot gekernt, Fleck 2a rot bepudert; Fleck 4 nahezu quadratisch. Hinterflii- gelumrandung schmal, am Apex verstarkt, mit auffalli- ger Verdickung („Zahn“) am beginnenden Analfeld: ein fast unbeschupptes, somit durchsichtiges Feldchen durchzieht die roten Hinterflügel von der Flügelwurzel fast bis zur schwarzen Analfeldverdickung reichend. Die beiden einzigen bislang bekannten Tiere erlauben keine endgültige Entscheidung, ob es sich um eine wohldifferenzierte Biospezies oder „nur“ um eine au- Bergewóhnliche Subspezies von Z. chirazica Reiss, 86 Bonner zoologische Beitráge 53 (2004) 1938, oder Z. naumanni Hille & Keil, 2000, handelt. Eine Beschreibung als Unterart von Z. chirazica waren jedoch nur móglich gewesen, wenn gleichzeitig die geographisch dazwischen siedelnde Z. naumanni syn- onymisiert worden wáre. Dieser Schritt sollte jedoch erst vollzogen werden, wenn weitere gut begriindete Argumente (Larvalbiologie, Pheromone, DNA, Kreu- zungs- und Anlockexperimente) vorliegen. Habituelle Unterschiede zu diesen beiden Taxa, die den úblichen Rahmen einer Subspezies übertreffen, sind nicht zu tibersehen. Insbesondere der ausgepragte hyali- ne Wisch auf den Hinterfliigeln von der Fliigelwurzel bis zum „Zahn“ der Umsäumung stellt eine eigenartige Besonderheit dar, die in dieser Form bei keiner anderen Z. chirazica-Unterart einschließlich Z. naumanni auf- tritt. Vielmehr erinnert diese Merkmalskombination („schwarzer Zahn — hyaliner Wisch**) eher an Z. escale- rai Poujade, 1900, (etwa aus Dorud oder Sibak) zu der auch weitere Ahnlichkeiten bestehen, so z. B. die unter- schiedliche Vorder- (gelb) und Hinterflúgelfárbung (rot). Die Mehrzahl signifikanter Merkmale (zweifaches Abdominalcingulum, Ausgestaltung und Anhángung der beiden Vorderflügelflecken 5+6 etc.) lässt jedoch eine nähere Verwandtschaft mit Z. chirazica/naumanni vermuten. Die nächsten bekannten Vorkommen von Z. naumanni sind ca. 500 km weiter südöstlich gelegen, dazwischen sind bislang keine Populationen bekannt geworden. Z. chirazica hat ihre nächsten Fundorte bei Semirom und Meymand (ca. 730 km Luftlinie entfernt). Am Locus typicus zwischen Baneh und Saqez unweit der irakischen Grenze fliegt die neue Art zusammen mit Z. tamara Christoph, 1889, und Z. cuvieri Boisduval, [1828]. Die habituelle Ähnlichkeit mit der syntopen Z. tamara (anderes Subgenus!) ist nicht zu übersehen (vel. Abbildungen). Sie tritt hier in der gleichen zweifarbigen Form („mahabadicoid“) mit gelber Patagia, gelben Vor- der- und roten Hinterflügeln auf. Zygaena (Agrumenia) haberhaueri demangei subsp. nov. (Figs. 34 — 39) Zygaena (Agrumenia) haberhaueri Lederer, 1870, Annales de la Societé entomologique de Belge 13 (1869-70): 29, 45 Derivatio nominis: Fiir Jacques DEMANGE (Denis du Val), dem wir die Entdeckung der am stárksten abwei- chenden Population dieser Unterart verdanken. Holotypus, ©: „Iran prov. Azerbaijan E, Kandovan 2350m, 37°N47,063 46%E 15,689, 18-20 VII 2000, J&M DEM[ANGE leg.]*; coll. A. HOFMANN (später deponiert in coll. SMNK). Paratypen: 7 Í9, 3 Y Y, Daten wie Holotypus, coll. A. HOFMANN; weitere Paratypen (Daten wie Holotypus) coll. J. DEMANGE. Von der nominotypischen Unterart und allen nachfol- gend beschriebenen hierzu gehórigen Taxa unterschei- det sich Z. h. demangei subsp. nov. durch auffallig gro- Be, kräftig weiß umsáumte Vorderfliigelflecken und ein gut entwickeltes rotes Abdominalcingulum (meistens úber 2-3 Segmente). Eine rote Halskrause ist stets vor- handen, 7% besitzen zumindest andeutungsweise rote Tegulae. Diese Schönheitsmerkmale erinnern an Z. oli- vieri Boisduval, [1828], von der sich Z. h. demangei subsp. nov. jedoch deutlich durch ihren haberhaueri- typischen, kälteren Rotton abhebt. Die Vorderflügelfle- cken 1+2 und 5+6 sind paarweise im Rot verbunden, Fleck 3+4 werden nicht selten durch weiße Schuppen getrennt. Entlang der Costa der Vorderflügel zieht sich ein Wisch weißer Schuppen. Am stärksten ist diese Ausprägung bei Tieren von der Typenlokalität zu er- kennen. Individuen aus Hamadan sind zwar ebenfalls noch durchwegs cinguliert, häufig jedoch nur auf einem oder zwei Segmenten. Die roten Tegulae bei den YY sind auf wenige Schuppen reduziert. Unweit Zanjan tre- ten dann die ersten uncingulierten Individuen hinzu (ca. 5 %). Die Flecken werden kleiner, die prominente Weißumrandung schmäler. Z. h. haberhaueri und Z. h. elbursica TREMEWAN, 1975, sind uncinguliert (ca. 90 %), durch ihre kleineren Vorderflügelflecken und schwächer ausgeprägte Weißumsäumung nicht mit Z. h. demangei subsp. nov. verwechselbar. Z. haberhaueri optima Reiss, 1939 , ist ebenfalls uncinguliert und be- sitzt darüber hinaus ein weißes Collar. Z. haberhaueri demangei subsp. nov. besiedelt ein ver- gleichsweise großes Areal in Nordwestiran. Eindeutig hierzu gehöriges Vergleichsmaterial liegt aus den Pro- vinzen Azarbaygan-e Sharqi (Kuh-e Sahand, Talish, Ka- laybar), Zanjan und Hamadan (Gardaneh-ye Avaj) vor. Auch Tiere weiter nordwestlich anschlieBender Regio- nen (Buzgov/Nachitschevan, Talysh/Republik Azar- beidjian) besitzen — wenngleich etwas abgeschwácht — eindeutig hierher verweisende Merkmalsziige (Cingu- lum, weiße Fleckenumrandung). Zygaena (Zygaena) viciae pelopis subsp. nov. (Figs. 46 — 48) Zygaena (Zygaena) viciae [Denis & Schiffermiiller], 1775, Ankiindung eines systematischen Werkes von den Schmetter- lingen der Wienergegend: 45 (Sphinx) Derivatio nominis: Nach Pelops, Gestalt der griechi- schen Mythologie, máchtigster Kónig der Insel, die von ihm den Namen erhält. „Peloponnesos“: Insel des Pe- lops. Holotypus, : „HELLAS/Peloponnes, Aroania-Oros, Chelmos, Kalavrita 1600 m, 5.-21.VH.1987, H. Axel HOFMANN: Neue Zygaena-Taxa aus Súdosteuropa, Vorder- und Zentralasien 8 ARHEILGER [leg.]“; coll. A. HOFMANN (später deponiert in coll. SMNK). Paratypen: | Y, Datum wie Holotypus, coll A. HOF- MANN; ibidem, 5 44, 1 9, coll. H. ARHEILGER; ibidem, 8 88, 4 99, 1300 m, 15. - 22. VI. 1987, coll. H. ARHEILGER; ibidem 1 $, „T. ARHEILGER [leg.]“, coll. H. ARHEILGER; ibidem 1 6, „1750 m, 5. — 21. VII. 1987, H. ARHEILGER“ [leg.]; 1 7, „ Griechenland, Pelo- ponnissos, Aroanıa Ori, Umgebung Kalavryta, 20.6.1976“, leg. J. WOLF; ibidem 1 Í, 28.6.1976; ibi- dem 1 Y, 24.6.1986; 1 @, „ Griechenland, Peloponnis- sos, Erymanthos Oros, 1500 m, 29.6.1990, leg. J. WOLF: 1 $, „Graecia mer., Peloponnes sept., 7.6. 1976, Trikala, 1400 m, leg. K. BERNHAUER“, ex coll. G. REISS, coll. A. HOFMANN; 1 ©, „Greece, Prov. Ahaia, versant nord du mont Chelmos, 1450/1650 m, 3/VII/1996, Dominque DUMONT leg.“. Z. viciae pelopis subsp. nov. zeichnet sich durch schma- le, spitze Vorderfliigel und habituelle Annáherung an Z. filipendulae aus. Die Tiere sind groß, im Durchschnitt sogar größer als syntope Z. filipendulae. Ein Abdomi- nalcingulum fehlt stets. Die Vorderflügelflecken 1+2 und 5+6 sind paarweise verbunden, 3+4 werden von schwarzen Setae getrennt. Fleck 5 hängt abgewinkelt an Fleck 6 (ähnlich Z. graslini Lederer, 1855). Fleck 6 ist verkleinert; nur bei einem einzigen von 25 vorliegenden Tieren fehlt diese Endmakel. Die Rotfärbung ist ver- gleichsweise trist, keinesfalls leuchtend oder orangefar- ben. Die Z. viciae-typische Hinterflügelumsäumung ist am Apex und im Analfeld erweitert. Dieses Merkmal in Kombination mit dem winzigen Fleck 3 erlaubt eine si- chere habituelle Unterscheidung von Z. filipendulae. Im Vergleich mit anderen Z. viciae-Unterarten zeigt Z. v. pelopis subsp. nov. gewisse Ähnlichkeiten mit Z. v. amanica Reiss, 1935 vom Nurudag Gegidi unweit der türkisch-syrischen Grenze. Die geographisch nächstste- henden Unterarten Z. viciae silbernageli Reiss, 1943 und Z. v. bosniensis Reiss, 1922 sind fast ausnahmslos 5-fleckig. Sie sind dichter beschuppt mit wesentlich wärmerem Rotton und breiteren Vorderflügeln. Dies trifft auch für mazedonische (Ochird- u. Prespan-See vic.) und nordgriechische Populationen (Kalambaka, Katara, Pilion etc.) zu. Abdominalberingung tritt hier gelegentlich auf. Zygaena (Zygaena) filipendulae wiegelorum subsp. nov. (Figs. 52 — 54) Zygaena (Zygaena) filipendulae (Linnaeus, 1758), Systema Naturae, ed. X, 1: 494 (Sphinx) Derivatio nominis: Dem Ehepaar Dr. Karl-Heinz und Charlotte WIEGEL in Anerkennung seiner Verdienste um die Erforschung der Biologie mehrerer Zygaena- Arten gewidmet. Als Autoren dieses Taxon zeichnen A. HOFMANN und W. G. TREMEWAN verantwortlich Holotypus “y, „Turkey: Erzurum, Askale (7 km SW.), 2000 m. 20.v11.1999“ leg. et coll. W. G. TREMEWAN (spáter deponiert in coll. Staatliches Museum fiir Natur- kunde Stuttgart) j Paratypen: 8 43,8 29, „Turkey: Erzurum, Askale (7 km SW.), 2000 m. ab ovo, 27.111.-8.1v.2000. W. G. & S. M. TREMEWANS, coll W.G. TREMEWAN; 1 Ú, „Türkei, Erzurum, 20 km nórdl. Askale, Kop Gec., 1700 m, 19.7.1992, leg. G. BAISCH”, coll. A. HOFMANN; | Ú, i- bidem, 2300 m, 29.7.1992; 2 44, „Türkei, n. Askale, Kopdag, 2000 m, Tasagil, Weideland/Máhwiesen, 7.7.1997, J. U. MEINEKE leg.“, coll. A. HOFMANN; 2 74, „Türkei, Bayburt, Nórdl. Kop. Gec., 2100 m, 8.7.1996, leg. W. TEN HAGEN“, coll. A. HOFMANN; 4 SC, „Turkey, Erzurum, Kop. gec., 2370 m, 40.01.40N, 40.31 E, 17.7.1996, K. SPATENKA lgt.*, coll. A. HOF- MANN; ibidem, 4 5d, 1 Y, coll. C.M. NAUMANN; 2 dd „Kop-gecidi, 2350 m, 29.7.77, loc. 50, leg. S. WAGE- NER, coll. C.M. NAUMANN. Zu Z. f. wiegelorum subsp. nov. gehören ferner zahlrei- che weitere Vorkommen — so vom Palandöken Dag (Er- zurum SE), den Munzur und Mercan Dagları, aus der Umgebung von Ovacik (Erzurum) und Yusufeli sowie vom Buglan Gecidi (Bingöl — Mus) — die wir jedoch nicht als Paratypen werten. Z. filipendulae wiegelorum subsp. nov. ist die am stärks- ten rotkonfluente Unterart. Sie stellt insofern das extre- me Gegenstück zu den melanistischen Unterarten Z. f gigantea Rocci, 1913 und Z. f. himmighofeni Burgeff, 1926 dar. Die 6 Vorderflügelflecken sind paarweise so miteinander verbunden, dass sie nur noch als 3 einzelne Großmakel auszumachen sind. Bei den vorliegenden Tieren vom Locus typicus geht diese Tendenz noch wei- ter: Ein durchgehender roter Striemen von Fleck 2 nach 4 und weiter nach 5+6 verbindet alle Flecken zu einem geradezu neuen Phänotypus. Bei einigen Männchen sınd die Makel flächig auch entlang des Vorderflügelaußen- randes verbunden; solche Tiere erinnern stärker an kräf- tig beschuppte Z. purpuralis als an Z. filipendulae. Der Hinterflügelsaum ist sehr schmal. Z. filipendulae wiegelorum subsp. nov. besiedelt ein re- lativ großes Areal vom Buglan Gecidi im Süden (Prov. Bingöl) bis Erzurum und Bayburt (Prov. Gümüshane) im Norden. Auch ein einzelnes vorliegendes Weibchen von Yusufeli (Prov. Coruh/Artvin) sowie Tiere aus den Rize dagları und aus der Umgebung von Gümüshane scheinen hierher zu gehören. Nach Nordwesten und Os- ten schließen sich hiervon völlig abweichende, normal aussehende Z. filipendulae — Populationen an (Z. f ti- rabzonica Koch, 1942 und Z. f. kulpiensis Reiss, 1935). Nur die im Südosten kontaktierenden Z. filipendulae 88 Bonner zoologische Beitráge 53 (2004) schuberti Reiss & Reiss, 1973 vom Van-See-Gebiet tendieren bereits zur Fleckenverschmelzung. Hier sind die Vorderfliigelflecken oft paarweise angenáhert oder verschmolzen, meistens jedoch noch deutlich als Ein- zelmakel zu erkennen. Größerflächige Konfluenz hin zu Z. purpuralis-artigen Tieren tritt bei Z. f. schuberti nicht auf. - Erythristische Populationen/Unterarten sind aus der Ost- türkei auch von Z. brizae, Z. purpuralis, Z. olivieri (NAUMANN & HOFMANN, im Vorbereitung) und Z. loti (latifa Naumann & Naumann, 1978) bekannt. Dabei fallt auf, dass mehrere Arten gerade in der Provinz Er- zurum zur Verstárkung der roten Zeichnungsanlagen tendieren (Z. purpuralis, Z. olivieri, Z. carniolica etc.) und mindestens 2 Arten (Z. loti und Z. filipendulae) hier ihre extremsten roten Unterarten hervorgebracht haben. Zygaena (Zygaena) lonicerae pseudangelicae subsp. nov. (Figs. 55 — 57) Zygaena (Zygaena) lonicerae (Scheven, 1777), Naturforscher, Halle 10: 97 (Sphinx) Derivatio nominis: Ostanatolische Populationen von Z. lonicerae (Scheven, 1777) zwischen Buglan Gegidi und Van-See zeigen starke habituelle Anklánge an Z. ange- licae Ochsenheimer, 1808. Holotypus, ©: „Türkei or., Prov. Van, Kuzgunkiran Gecidi, 1900 m, 19. — 23. 7. 1981, GROß, HERBST, R. & A. HFM.“, coll. A. HOFMANN (später deponiert in coll. Staatliches Museum für Naturkunde Karlsruhe). Paratypen: 1 Y, Datum wie Holotypus; ibidem, 4 44, 14.7.1982, P. STRAUB & A. HOFMANN leg., coll. A. HOFMANN; 2 44, „15 km östl. Tatvan, 1800 — 1900 m, 16.7.1986, leg. K. SCHURIAN, Coll. Nr. 215”, coll. A. HOFMANN; 6 Cd, 9 2&,“Asia min., Kurdistan, Van- See-Gebiet, Bitlis Vill., 20 km óstl. Tatvan, 1750 m, 10. — 12. 7. 68, feuchte Wiesen, leg. MITTENDORF“, coll. C. M. NAUMANN; ibidem, 3 33, 2 Y Y, coll. A. HOFMANN via coll. C.M. NAUMANN. Zu Z. I. pseudangelicae subsp. nov. gehören ferner die Aufsammlungen vom Buglan Gecidi (die ich jedoch nicht als Paratypen werte): 1 Y, „Prov. Mus, Buglan Gecidi, 1600 m, 28.6. — 4.7. 1977, leg. HOLZSCHUH & RESSL“, coll. A. HOFMANN; ibidem, 1 4, „Bingöl/Mus, Buglan Gecidi, 1650 — 1750 m, 03.07.1986, A. HOF- MANN & G. & S. REISS“, coll. T. WITT (via coll. G. REISS); 1 Y, “Türkei, Prov. Bingól, Buglan gec., 1600 m, 26.V1.82, leg. W. THOMAS”, coll. C. M. NAUMANN. Neben der geringen Größe unterscheiden sich die zierli- chen Tiere recht augenfällig von allen anderen Z. loni- cerae-Unterarten durch leuchtend orangerote Färbung, vergleichsweise dünne Fühler und kleine, fast runde Vorderflügelflecken 3, 4 und 5. Die schwarze Hinter- flügelumsäumung ist im Analbereich sehr schmal, vom Cupido bis zum Apex mäßig verstärkt. Insgesamt erin- nern die Tiere stark an Z. angelicae. Die eurosibirisch verbreitete Art, Z. lonicerae, gehört in Zentralanatolien zu den großen Seltenheiten. Während das Areal der nordanatolischen und armenischen Z. lo- nicerae-Populationen (subsp. abbastumana Reiss, 1922) im pontischen Waldgürtel liegt, wo die Art auch flä- chenhaft verbreitet ist, besiedelt Z. /. pseudangelicae subsp. nov. nur sehr kleinräumige, heute größtenteils waldfreie Lebensräume des östlichen Zentralanatolien. Die Biotope am Buglan und Kuzgunkiran Gecidi sind azonale, mesophile Kleinbiotope in der subalpinen Grassteppe, stellenweise Mähwiesen (Mahd ab Anfang, Mitte Juli), in ihren Randbereichen oft von lichtem Ei- chenbuschwald durchsetzt. In diesem ökologisch beson- ders reich strukturierten Raum überlagern sich rezent die Areale zahlreicher arborealer und eremialer Arten und bedingen so eine besonders hohe Artendiversität. Z. lonicerae tritt hier syntop mit Z. cuvieri Boisduval, [1828], Z. cambysea Lederer, 1870, Z. purpuralis (Brünnich, 1763), Z. minos (|Denis & Schiffermiiller], 1775), Z. olivieri Boisduval, [1828], Z. sedi Fabricius, 1787, Z. carniolica (Scopoli, 1763), Z. loti ([Denis & Schiffermüller], 1775), Z. viciae ([Denis & Schiffermiil- ler], 1775), Z. doryenii Ochsenheimer, 1808 und Z. fili- pendulae (Linnaeus, 1758) auf. In der náheren Umge- bung sind weitere Arten (Z. manlia Lederer, 1870, Z. rubricollis Hampson, 1900, Z. tamara Christoph, 1889, Z. punctum Ochsenheimer, 1808, Z. brizae (Esper, 1800), Z. osterodensis Reiss, 1921, Z. peschmerga Eck- weiler & Górgner, 1981) anzutreffen, so dass in tiirkisch Kurdistan zwischen Buglan Gegidi und Van-See-Gebiet mit 19 Arten ein rezentes Mannigfaltigkeitszentrum der Gattung Zygaena erreicht sein dürfte. Für Z. lonicerae bilden diese Nachweise zugleich ihre südöstliche Areal- grenze. Danksagung. Beiträge zum Gelingen dieser Arbeit haben beigesteuert: H. Arheilger, T. Arheilger, G. Baisch, F. Car- bonell, J.-M. Desse, J. Demange, D. Dumont, G. Ebert, Dr. W. Eckweiler, G. Flutsch, J. J. de Freina, Dr. A. Kallies, H. u. P. Kautt, T. Keil, R. Leestmans, Dr. J.-U. Meineke, B. Mollet, M. Nicolle, Prof. Dr. C. M. Naumann (7), Dr. G. Reiss (+), Dr. K. Schurian, Dr. G. Tarmann, Dr. K. Spatenka, M. Ströhle, Dr. W. ten Hagen, Dr. W. G. Tre- mewan und J.-C. Weiss. Ihnen allen bin ich einmal mehr sehr zu Dank verpflichtet. LITERATUR DESSE, J.-M. (2003): An extension of the known distributi- on of Zygaena rubricollis Hamspon, 1900 (Lepidopte- ra: Zygaenidae, Zygaeninae) in EFETOV, K. A., TRE- MEWAN, W. G. & TARMANN, G. M. (eds.) Proceedings of the 7th International Symposium on Zygaenidae (Insecta, Lepidoptera), 10. Innsbrucker Lepidopteren- Axel HOFMANN: Neue Zygaena-Taxa aus Südosteuropa, Vorder- und Zentralasien 89 gesprach, Innsbruck 4 - 8 September 2000 Crimean State Medical University Press, Simferopol: 15, 16. EFETOV, K. A., TREMEWAN, W. G. & G. M. TARMANN (2003): Proceedings of the 7th International Symposi- um on Zygaenidae (Insecta, Lepidoptera), 10. Innsbru- cker lepidopterengespräch, Innsbruck 4 - 8 September 2000 Crimean State Medical University Press, Simfe- ropol. HILLE, A. & KEIL, T. (1999): Eine neue Zygaena aus dem Iran - Zygaena naumanni n. sp. (Lep., Zygaenidae). Entomologische Nachrichten und Berichte Dresden 43: 249-253. HOFMANN, A. (2003): Problems concerning the systematic position of Zygaena formosa molleti n. subsp. Abs- tracts of the VIII International Symposium on the Zy- gaenidae, Dresden, 10-14 September 2003: 23. HOFMANN, A. (2000): Contribution to the knowledge of the genus Zygaena Fabricius, 1775 in Iran (Lepidoptera, Zygaenidae. Part I: Introduction, systematic part: Zy- gaena (Mesembrynus) seitzi, Z. ?seitzi nocturna, Z. manlia. Linneana Belgica 17(5): 171-196. HOFMANN, A. (2000): Contribution to the knowledge of the genus Zygaena Fabricius, 1775 in Iran (Lepidoptera, Zygaenidae. Part Il: Zygaena rubricollis. Linneana Belgica 17(6): 227-232. HOFMANN, A. (2000): Contribution to the knowledge of the genus Zygaena Fabricius, 1775 in Iran (Lepidoptera, Zygaenidae. Part IV: Zygaena haematina (addendum), Z. cacuminum. Linneana Belgica 17(8): 339-347. HOFMANN, A. & TREMEWAN, W. G. (1996): A Systematic Catalogue of the Zygaeninae (Lepidoptera: Zygaeni- dae). 251 pp., Harley Books, Colchester, Essex , Eng- land. HOFMANN, A. & W. G. TREMEWAN (2003): Contribution to the knowledge of the genus Zygaena FABRICIUS, 1775 in Iran (Lepidoptera, Zygaenidae. Part VI: Zygaena nocturna, Z. aisha, Z. rubricollis and Z. fredi (addenda based on results of fieldwork in 2001 and 2002). Lin- neana Belgica 19(1): 9-20. KEIL, T. (2003): New data on the biology of Zygaena (Agrumenia) rosinae KORB, 1903 and Z. (4.) sengana HOLIK & SHELJUZHKO, 1956 stat. rev. in South Iran (Lepidoptera: Zygaenidae, Zygaeninae). Abstracts VI- I International Symposium on the Zygaenidae, Dres- den, 10-14 September 2003: 29-30. KEIL, T. (2003): Attend for a new classification of the taxa around Zygaena rubricollis Hampson, 1900 (Lepidop- tera: Zygaenidae, Zygaeninae). Abstracts VIII Interna- tional Symposium on the Zygaenidae, Dresden, 10-14 September 2003: 31-32. NAUMANN, C. M. (1969): Zur Kenntnis der Mesembrynus- Arten Afghanistans (Lep., Zygaenidae). Bonner zg0- logische Beitráge 20: 266-278. NAUMANN, C. M. (1974): Neue Zygaena-Unterarten aus Afghanistan (Lep., Zygaenidae). Entomologische Zeit- schrift Frankfurt am Main 84: 29-36. NAUMANN, C. M. (1977): Zygaena (Mesembrynus) halima n. sp. und einige Bemerkungen zur stammesgeschicht- lichen Gliederung der Gattung Zygaena F. (Lepidopte- ra: Zygaenidae). Zeitschrift der Arbeitsgemeinschaft österreichischer Entomologen 29: 35-40. NAUMANN, C. M. & TREMEWAN, W. G. (1984): Das Bi- ospecies-Konzept in seiner Anwendung auf die Gat- tung Zygaena Fabricius, 1775 (Insecta, Lepidoptera, Zygaenidae). Spixiana 7: 161-193. NAUMANN, C. M., FEIST, R., RICHTER, G. & WEBER, U. (1984): Verbreitungsatlas der Gattung Zygaena Fabri- cius, 1775 (Lepidoptera, Zygaenidae). Theses zoologi- cae 5: 1-45, text-fig., maps 1-97. NAUMANN, S. & NAUMANN, C. M. (1978): Eine neue ost- anatolische Unterart der Zygaena (Zygaena) loti (De- nis und Schiffermúller, 1775) (Lep., Zygaenidae). En- tomologische Zeitschrift Frankfurt am Main 88: 37-40. NAUMANN, S. & NAUMANN, C. M. (1980): Ein Beitrag zur Kenntnis der Zygaenen-Fauna Nord- und Ost- Anatoliens (Lep., Zygaenidae). Entomofauna 1: 302- 353. TARMANN, G. M. & TREMEWAN, W. G. (2003): The status of Zygaena (Zygaena) loti senilis Burgeff, 1914 (Lepi- doptera: Zygaenidae, Zygaeninae). Entomologist's Gazette 54: 183-186. TREMEWAN, W. G. & NAUMANN, C. M. (1989): A revision of Zygaena (Agrumenia) magiana Staudinger, 1889 (Lepidoptera: Zygaenidae). Entomologist’s Gazette 40: 114-122. Anschrift des Autors: Axel Hofmann, Verenenweg 4, D-79102 Breisach-Hochstetten, Germany; axel.hofmann(@debitel.net 90 Bonner zoologische Beitráge 53 (2004) 1-3: Zygaena (Mesembrynus) rubricollis arachosica n. subsp.; 1: Holotypus, 4, 31 mm: „Pakistan, Baluchistan, Quetta/Urak, 2400 — 2700 m, 24. — 26.V.83, leg. ECKWEILER“; 2: Paratypus, 3, 36 mm: Daten wie Holotypus je- doch 23. — 24.V. 1979; 3: Paratypus, Y, 34 mm: ibidem. 4-6: Zygaena (Mesembrynus) rubricollis rubricollis Hamspon, 1900; 4: Topotypus, 4, 34 mm: "Pakistan, Chitral, Birmoglasht, 2700 - 2900 m, 6. VIL83, leg. ECKWEILER“; 5: Topotypus, Y, 32 mm: „Chitral, 2800 - 3000 m, Bir- moglasht, Chagh Bini, Pakistan, 30-VI-97, [ leg.] MARC NICOLLE”; 6: Topotypus, Y, 33 mm: Daten wie fig. 4 je- doch 29. V1.83. 7: Zygaena (Mesembrynus) rubricollis shahkuhica n. subsp.; Holotypus, ©, 30 mm: „Iran, prov. Mazandaran, Shah Kuh (Est), versant nord, 2800 — 3000 m, N36°32-E54°26, 15/17-VH-1998, [B.] MOLLET leg.“. 8: Zygaena (Mesembrynus) manlia manlia Lederer, 1870: 4, 30 mm: „Iran, prov. Mazandaran, Shah Kuh (Est), versant nord, 2800 — 3000 m, N36°32-E54°26, 17/19-V1-2000, G. FLUTSCH & B. MOLLET leg.*. Sharud S, Shah Kuh-e Pa’in vic., 2700 — 2900 m, 19. — 21. 7. 1999, A. HOFMANN, B. MOLLET & J.U. MEINEKE leg.”. 9: Zygaena (Mesembrynus) cacuminum Christoph, 1877; Topotypus, ©, 31mm: Iran, Mazandaran, Shah Kuh, 10-12: Zygaena (Mesembrynus) nocturna nocturna Ebert, 1974; 10: Paratypus, ©, 33 mm: “[Iran], Fars, 50 km NW Ardekan, Tange Surkh, 2250 m, 16.6.1972, EBERT/PAZOUKI leg. “; 11: Í, 34 mm: „Iran, L. F., Prov. Boyer Ahmad- va-Kohgiluyeh, Gardaneh Meymand, 2450 — 2800 m, 15. — 17.6.2001, J.U. MEINEKE & A. HOFMANN leg.“; 12: 3, 31 mm: „Iran, e.o., Chaharmahal-va-B., Borujen S, Gardaneh-ye Chenar Mahmoudi S, e.p.: 22.4. — 8.6.2002, A. HOFMANN cult.*. 13-15: Zygaena (Mesembrynus) nocturna eberti n. subsp.; 13: Holotypus, ©, 32 mm: Iran, e.o., Lorestan, Dorud, Golgolab Dag, 2600 m, e.p.: 7.5. — 27.5. 2000, A. HOFMANN cult.“; 14: Paratypus, ©, 32 mm: Daten wie Holotypus; 15: Í, 33 mm: „Iran, e.p., Prov. Esfahan, Fereydun Shahr S, Kamaran vic., 2700 — 2800 m, e.p.: 29.6.2001, A. HOFMANN & J.U. MEINEKE leg.“. 16-18: Zygaena (Agrumenia) magiana pamirescens n. subsp.; 16: Holotypus, ¢, 26 mm: “Kirghizstan, Alai mountains, Gaumish pass, H = 3.600m, 26.07. 1999, coll. A. HOFMANN“; 17: Paratypus, ©, 25 mm, Daten wie Holotypus; 18: Paratypus, Y, 27 mm, ibidem. Axel HOFMANN: Neue Zygaena-Taxa aus Súdosteuropa, Vorder- und Zentralasien 9] 92 Bonner zoologische Beitráge 53 (2004) 19: Zygaena (Agrumenia) magiana magiana Staudinger, 1889: Í, 26 mm: “Turkestan” 20: Zygaena (Agrumenia) magiana alaica Holik & Sheljuzhko, 1956: Í, 27 mm: “UdSSR, Kirghisistan, Alai-Geb., Kaindy b. Ferghana, 3000 — 3500 m, 10.-20.7. 1986, Coll. Dr. A. SCHULTE, coll. A. HOFMANN”. 21: Zygaena (Agrumenia) magiana alaudina Tremewan & Naumann, 1989: Topotypus, Í, 24 mm: “N. Tajikistan, Zerevshan, Fanskiye Gory, Alaudin Lakes, 3.600 m, 29. VII. 1997, leg. G. LUTSENKO, ex coll. C.M. NAUMANN, coll. A. HOFMANN“. 22-24: Zygaena (Agrumenia) cocandica ad cocandica Erschoff, 1874: “Kirgiztan, Alai, Kollectorniy Range, Gau- mich Pass, 4.8.1998, coll. A. HOFMANN.“; 22: Í, 24 mm; 23: Í, 24 mm; 24: 9, 26 mm. 25-27: Zygaena (Agrumenia) formosa molleti n. subsp.; 25: Holotypus, 4, 24 mm: “Turkiye, prov. Nigde, Ulukisla, 2500 — 3000 m, 4-VIII-1995, MOLLET B. leg.”; 26: Paratypus, ©, 25 mm, Daten wie Holotypus; 27: Paratypus, Y, 26 mm, ibidem. 28-30: Zygaena (Agrumenia) formosa malatiana Rebel, 1901; 28: 3, 23 mm: “Türkei centr., Sivas 10 — 14 km S, 1350 m, 15./16.7.1986, A. HOFM., G. & ST. REISS”; 29: Í, 23 mm: “Türkei mer., Nigde S, Ulukisla 11 km E, Ha- sangazi vic., 1160 m, 14.7.1986, A. HOFM., G. & ST. REISS”; 30: Y, 26 mm: Daten wie fig. 29. 31, 32: Zygaena (Agrumenia) tenhagenova n. spec.; 31: Holotypus, Í, 26 mm: “Iran, Kordestan, Paß NE Baneh (O- stseite), 1900 — 2100 m, 16.VI. 2001, leg. TEN HAGEN”; 32: Paratypus, Í, 26 mm, Daten wie Holotypus. 33: Zygaena (Mesembrynus) tamara mahabadica Reiss, 1978; ©, 27 mm, Daten wie fig. 31. 34-39: Zygaena (Agrumenia) haberhaueri demangei n. subsp.; 34: Holotypus, ©, 27 mm: “Iran, prov. Azerbaijan E, Kandovan, 2350 m, 37°N47,063 46°E15,689, 18-20 VII. 2000, J & M DEM[ANGE]“; 35: Paratypus, Í, 27 mm, Da- ten wie Holotypus; 36: Paratypus, Y, 30 mm, Daten wie Holotypus; 37: ¢, 28 mm: „Iran, Prov. Hamadan, Hamadan NNE, Razan N, Gardanye Avaj, 2300 — 2400 m, 19. 06. 1998, A. HOFMANN/J.-U. MEINEKE [leg.]“; 38: Y, 33 mm: „Iran, Zanjan, 25 km SW Zanjan (Pass), 1900 — 2000 m, 4. VII. 2003, leg. ECKWEILER + 890%; 39: : Q, 30 mm: ,,I- ran, Prov. Zanjan, Zanjan — Gilvan, Gargovol Dag, 1. Paß, ca. 1 km N. des Passes, 2400 — 2500 m, 26.6. 2001, A. HOFMANN & J.U. MEINEKE leg. ,,. Axel HOFMANN: Neue Zygaena-Taxa aus Súdosteuropa, Vorder- und Zentralasien 93 94 Bonner zoologische Beitráge 53 (2004) 40, 41: Zygaena (Agrumenia) chirazica eckweileri Naumann & Naumann, 1980; 40: Paratypus, ©, 22 mm: “S-Iran, Kerman, Bam/ Deh Bakri, 2000 — 2500 m, 23. — 25.V. 78, leg. ECKWEILER“, ex coll. C.M. NAUMANN; 41: Topoty- pus, Y, 24 mm: „Iran, Prov. Kerman, Bam W, Dehbakri 1 km S, 2200 m, 1.6. 1997, (26/97), A. HOFMANN & P. KAUTT [leg.]“. 42, 43: Zygaena (Agrumenia) chirazica chirazica Reiss, 1938; 42: 3, 24 mm: “Siidiran, Dasht-e-Arjan, 2000 — 2200 m, 60 — 75 km westl. Shiraz, Prov. Fars, 15. — 26. 5. 1978, leg. K. ROSE“; 43: Y, 28 mm: “Iran, Fars [sic], Kuh-e Dinar, ca. 15 km E Sisakht (PaB), 3200 m, 8. VII. 1997, leg. ECKWEILER”. 44, 45: Zygaena (Agrumenia) naumanni Hille & Keil, 2000; 44: Topotypus, 4, 25 mm: “Iran, Esfahan, 20 km NW Damaneh, Kuh-Derre Bid, Godar Chonsar, 2660 m,14.6.2000, leg. KEIL“; 45: Paratypus, Y, 27 mm: ,, “Iran, Esfa- han, 20 km NW Damaneh, Kuh-Derre Bid, Godar Chonsar, 2660 m,7.6.99, leg. HILLE & KEIL“, ex coll. C. M. NAUMANN. 46-48: Zygaena (Zygaena) viciae pelopis n. subsp.; 46: Holotypus, ©, 30 mm: „Hellas/Peloponnes, Aroania-Oros, Chelmos, Kalavrita, 1600 m, 5.-21. VII. 1987, H. ARHEILGER [leg.]“; 47: Paratypus, Í, 31 mm: ,,Griechen- land/Peloponnes, Prov. Achaia, Kalavrita, Aroania Oros, Mt. Chelmos, 5.-21.7.1987, 1750 m, HORST ARHEILGER leg.”; 48: Paratypus, Í, 32 mm: ,,Umg. Kalaryta, 1300 m, Greece, Prov. Achaia, Peloponnes, 15.-22.6.1982, leg. T. ARHEILGER™. 49: Zygaena (Zygaena) filipendulae noacki Reiss, 1962; 32 mm: ,,Kalavrita, Prov. Achaia, Peloponnes, Griechen- land, 28.5.1981, 800 m, H. ARHEILGER leg. Oberursel 1. T.“. 50, 51: Zygaena (Zygaena) viciae bosniensis Reiss, 1922; 50: Cotype, 4, 28 mm: „24/6 Bosnia 04, Koriéna, G. LE- ONHARD [leg.]*, ex coll. G. REISS; 51: ¢, 31 mm: „Griechenland, Thessalien, Halbinsel Pilion, Chania vic., 800 m, 26.6.1987, H. & A. HOFMANN leg.“ 52-54: Zygaena (Zygaena) filipendulae wiegelorum n. subsp.; 52: Holotypus, Y, 32 mm: „Turkey: Erzurum, Askale (7 km SW), 20. vii. 1999, W. G. TREMEWAN [leg.]“, coll. W. G. TREMEWAN; 53: Paratypus, 3, 34 mm: „Türkei, Er- zurum, 20 km nórdl. Askale, Kop Gec., 1700 m, 19.7. 1992, G. BAISCH [leg.]*; 54: Paratypus, ©, 31 mm: „Türkei, n. Askale, Kopdag, 2000 m, Tasagil, Weideland/Mähwiesen, 7.7. 1997, J.U. MEINEKE [leg.]“. (Fig. 52: Holotypus, Y, Zygaena (Zygaena) filipendulae wiegelorum n. subsp. in coll. W. G. TREMEWAN; alle ande- ren abgebildeten Tiere befinden sich in coll. A. HOFMANN/Breisach-Hochstetten; die Holotypen coll. A. HOFMANN werden später im Staatlichen Museum für Naturkunde/Karlsruhe deponiert werden). Axel HOFMANN: Neue Zygaena-Taxa aus Siidosteuropa, Vorder- und Zentralasien 95 96 Bonner zoologische Beitrage 53 (2004) 55-57: Zygaena (Zygaena) lonicerae pseudangelicae n. subsp.; 55: Holotypus, 4, 29 mm: „Türkei or., Prov. Van, Kuzgunkiran Gecidi, 1900 m, 19. -23. 7. 1981, GROß, HERBST, R. & A. HFM. [leg.]“; 56: Paratypus, Í, 29 mm: „Türkei or., Prov. Van, Kuzgunkiran Gecidi E, Karayollari vic., 1900 — 2000 m, 14. 7. 1982, STRAUB & A. HOF- MANN leg.“; 57: Paratypus, Y, 32 mm: ,,Anatolien, Prov. Mus, Buglan gecidi, 28.6. — 4.7. 1977, leg. HOLZSCHUH & RESSL“. 58-60: Zygaena (Zygaena) lonicerae abbastumana Reiss, 1922; 58: $, 35 mm: “Türkei sept. or., Prov. Kars, Posof 2 — 5 km E, 1750 m, 24.7. 1981, GROß, HERBST, R. & A. HFM. [leg.] 59: Í, 35 mm: ibidem; 60: 2, 37 mm: ibi- dem. 61: Lebensraum von Zygaena (Agrumenia) formosa molleti n. subsp. im Bereich der Dornpolsterstufe am Bolkar Dag-Massif unweit Ulukisla, 2600 — 3000 m, wo die Art zusammen mit Pyrgus bolkariensis und Polyommatus mol- leti vorkommt. 62: Raupennahrungspflanze (vermutl. Onobrychis spec.) von Zygaena (Agrumenia) formosa molleti n. subsp. 63: Eigelege von Zygaena (Agrumenia) formosa molleti n. subsp. (Figs. 61-63: 4-VIII-1995, B. MOLLET phot.) Axel HOFMANN: Neue Zygaena-Taxa aus Súdosteuropa, Vorder- und Zentralasien Sa Kal. Bonner zoologische Beitráge | Band 53 (2004) | Heft 1/2 | = — — 4 Seiten 99-107 3onn, Juni 2005 Leptopholcus (Araneae: Pholcidae) in Continental America: Rare Relicts in Low Precipitation Areas' | Bernhard A. HUBER”, Abel PÉREZ G.” & Renner L. C. BAPTISTA”? 2) Zoological Research Institute and Museum Alexander Koenig, Bonn, Germany Museu Nacional, Universidade do Brasil, Rio de Janeiro, Brazil Abstract. The genus Leptopholcus has a cireumtropical distribution, but the only New World records previously known were for the Greater Antilles. On these islands, Leptopholcus is quite common, probably due to the absence of Merago- nia from the Antilles. On the other hand, Leptopholcus was previously thought to be absent from the mainland where Metagonia is ubiquitous and species-rich, occupying the preferred habitat of Leptopholcus (the underside of leaves). The present paper describes three new species of Leptopholcus from South America: L. brazlandia from Distrito Federal, Brazil, L. pataxo from Bahia, Brazil, and L. evaluna from Sucre, Venezuela. All these species were collected in low numbers at low precipitation areas, and no further specimens could be found in large collections of South American pholcids, indicating a relict status of the genus in South America. The known distribution of the genus is documented, and taxonomic problems regarding its relationship with Pholcus and other genera are briefly discussed. Key words. New species, taxonomy, relict, South America, competitive exclusion, Metagonia 1. INTRODUCTION Representatives of Leptopholcus Simon, 1893 are rela- tively rare in collections, which is probably mainly due to their cryptic habits. Almost all known species live on the underside of leaves (PETRUNKEVITCH 1929; BRIGNOLI 1980; IRIE 1999), where they spend most of the day pressed against the surface (HUBER € PEREZ 1998). Their greenish coloration when alive and their extremely thin legs make them highly cryptic, both for predators and collectors. SIMON (1893) created the group Leptopholceae to in- clude pholcids with a flattened prosoma lacking depres- sions and sutures, and with a sternum that is longer than wide. Two genera were included: Leptopholcus (eight eyes) and Micromerys Bradley, 1877 (six eyes). Even though SIMON noted the similarities in the male genita- lia between Pholcus Walckenaer, 1805 and Leptophol- cus, he put Pholcus in a different group, the Pholceae. BRIGNOLI (1980) and DEELEMAN-REINHOLD (1986) suggested that the female external genitalia provide fur- ther means to separate Pholcus from Leptopholcus: they are covered by a chitinous plate in Pholcus, by unscle- rotized cuticle in Leptopholcus. In this regard, Lep- topholcus is similar to Micromerys, Micropholcus Deeleman-Reinhold & Prinsen, 1987 and Calapnita Simon, 1892. Substantial homoplasy is obviously pre- sent, and neither the relationships among genera nor the monophyly of the genera involved has ever been tested by cladistic analysis or supported by convincing argu- ments. Even worse, neither Pholcus nor Leptopholcus 1 In commemoration of Clas Michael Naumann zu Königsbrück (26.06.1939 — 15.02.2004) have ever been revised, and together they include some 120 nominal species. Pending a large scale revision, we consider BRIGNOLI’S (1980) statement still valid: “il conviendrait de se montrer prudent avant de réunir les deux groupes”. Considering this taxonomic situation, the assignment of the species below to Leptopholcus needs justification. First, we chose to follow the traditional, non- phylogenetic concept of the genus, i.e. pale pholcids with long abdomen, long and thin legs, rather flat cara- pace, widely spaced lateral eye triads, unsclerotized ex- ternal female genitalia, and without the synapomorphies or distinctive characters of similar genera (Panjange Deeleman-Reinhold & Deeleman, 1983 and Calapnita, see DEELEMAN-REINHOLD & DEELEMAN 1986: Micro- merys, see HUBER 2001; Pehrforsskalia Deeleman- Reinhold & van Harten, 2001, see DEELEMAN-REIN- HOLD & VAN HARTEN 2001; Micropholcus, see DEELE- MAN-REINHOLD & PRINSEN 1987). Second, the main point of this paper is barely affected by these taxonomic problems. Neither Pholcus nor Leptopholcus (nor any of the other genera mentioned) have previously been re- corded from South America [except for the synan- thropic P. phalangioides (Fuesslin, 1775) and some taxa that were misplaced and have been transferred to New World genera; see HUBER 2000]. The only previously known autochthonous representatives of the Pholcus- group sensu HUBER (1995) in the New World are some Leptopholcus species on the Antilles and some Pholcus species in the eastern United States (HUBER 2000). Third, the new species seem related to the species on the Antilles (both groups have modified hairs on the tips of the male paipal trochanter apophyses). and these are presently considered as Leptopholcus. 100 Bonner zoologische Beitráge 53 (2004) According to the concept above, Leptopholcus has a cir- cumtropical distribution, with all known records be- tween 29°S and 26°N. Figure | shows all published re- cords (circles) as well as unpublished records (squares) of material deposited in various institutions. DEELEMAN-REINHOLD & VAN HARTEN (2001) suggested that Leptopholcus dioscoridis Deeleman-Reinhold & van Harten, 2001 on Socotra Island might represent a relict. Pholcids in general, and Leptopholcus in particu- lar (PETRUNKEVITCH 1929; MILLOT 1946), seem to pre- fer humid habitats, but Socotra has a mean annual pre- cipitation of no more than about 170-190 mm (DEELEMAN & VAN HARTEN 2001). The three species a “Ge tu e SES @P-@. »:- ha j 9 40 go. described below might also be relicts for two reasons: first, all were collected in low precipitation areas: L. evaluna along a river in an area of xeric shrublands near the northern coast of Venezuela, L. brazlandia in *cer- rado” (i.e., tropical grasslands, savannas and shrub- lands), and L. pataxo in a transitional zone between “cerrado” and mixed palm-sand vegetation. Second, the preferred microhabitat of Leptopholcus (underside of leaves) is in the New World occupied by the very spe- cies-rich genus Metagonia Simon, 1893. Intriguingly, Metagonia is absent from the Antilles (except one cav- ernicole species: PEREZ & HUBER 1999), and it is only on the Antilles that Leptopholcus is fairly common and species-rich (HUBER 2000; see also Fig. 1). Fig. 1: Known distribution of Leptopholcus. Circles represent published records, squares represent unpublished material from the following institutions: National Museum of Natural History, Washington, D.C.; Zoological Research Institute and Museum Alexander Koenig, Bonn; collection Suresh Benjamin; National Museum, Bloemfontein; National Collection, Pretoria; California Academy of Sciences, San Francisco; Musée royal de l’Afrique Centrale, Tervuren. 2. TAXONOMY Style of descriptions is as in HUBER (2000). Measure- ments are in mm unless indicated otherwise. Material is deposited in the Museu Nacional, Rio de Janeiro (MNRJ) and in the Museo de Historia Natural de la Fundacion La Salle, Caracas (MFLS). 2.1. Leptopholcus brazlandia Huber, Pérez & Baptista, new species (Figures 2, 3, 8, 9, 14-18) Types. Male holotype and two female paratypes from Brazlándia (15°41’S, 48°12’W), Nucleo Rural, Chacara 33, Distrito Federal, Brazil; Dec. 19, 2003 (A. Chagas, B. Segal), in MNRJ (04149-50). Etymology. The species name is a noun in apposition, taken from the type locality. Diagnosis. Easily distinguished from other New World species by the spotted abdomen (Figs. 2, 3); also by the shapes of procursus, bulbal apophyses, and the internal female genital sclerites (Figs. 14, 15, 17, 18). Male (holotype). Total length 2.8 (3.0 with clypeus), carapace width 0.97. Leg 1 missing, tibia 2: 3.8, tibia 3: 2.3, tibia 4 missing. Habitus as in Figures 2 and 3. Pro- soma ochre-grey with blackish mark dorsally, clypeus barely darkened, posterior half of sternum darker, with light spots near coxae 3 and 4; legs ochre-yellow, patel- lae and tibia-metatarsus joints blackish; abdomen ochre- grey with many dark marks except ventrally. Ocular area slightly elevated (Fig. 3), apparently with brush of hairs between triads (most hairs missing), thoracic fur- row absent; distance PME-PME 285 um; diameter PME 90 um; distance PME-ALE 20 um; AME totally absent. Clypeus unmodified. Sternum wider than long (0.6/0.5), unmodified. Chelicerae as in Fig. 16, with distal apophyses carrying two modified hairs each, with some macrosetae on each side near the apophyses, proximally with slightly sclerotized projections. Palps as in Figures 14 and 15; coxa unmodified, trochanter with unscle- rotized retrolateral hump and long retrolatero-ventral apophysis with modified hair distally (small and cone- shaped rather than cylindrical as in fig. 105 in Huber Bernhard A. HUBER, Abel PÉREZ & Renner L. C. BAPTISTA: Leptopholcus in Continental America 101 Figs. 2-13: Habitus and female external genitalia photographs. 2-3. L. brazlandia. 4-5. L. pataxo. 6-7. L. evaluna. 8-9. L. brazlandia, female abdomen, lateral view, and external genitalia, ventral view. 10-11. L. pataxo, female, lateral view, and exter- nal genitalia, ventral view. 12-13. L. evaluna, female abdomen, lateral view, and external genitalia, ventral view. 102 Bonner zoologische Beitráge 53 (2004) 17 Figs. 14-18: L. brazlandia, left male palp in prolateral (14) and retrolateral (15) views, male chelicerae (16), and cleared female genitalia in ventral (17) and dorsal (18) views. a: appendix, b: bulb, e: embolus, p: procursus, t: trochanter-apophysis, asterisk: proximal sclerotized element of bulb. Scale lines: 0.5 mm (14, 15), 0.2 mm (16), 0.3 mm (17, 18). 2000); femur with indistinct ventral protrusion and two distinct retrolateral apophyses; procursus relatively sim- ple except distally, with capsulate tarsal organ; bulb with proximal sclerotized element, apparently without uncus, with complex appendix(?) and slightly scle- rotized embolus. Retrolateral trichobothrium of tibia 2 at 15%; legs apparently without spines, curved hairs, and vertical hairs (most hairs missing); tarsus 2 with about 10 fairly distinct pseudosegments. Abdomen pos- teriorly pointed dorsally (Fig. 3). Female (paratype). In general similar to male. Tibia 1 missing in both specimens. Female external genitalia protruding but simple (Figs. 8, 9), consisting of frontal Bernhard A. HUBER, Abel PEREZ & Renner L. C. BAPTISTA: Leptopholcus in Continental America 103 plate and smaller median plate that carries a small indis- tinct knob-like structure. Internal genitalia with complex system of distinctive sclerites and folds (Figs. 17, 18). Distribution. Known only from type locality. Remarks. The collecting area is located near the bor- ders of the Cupim river, at the northeast of the Federal District. The Cupim river flows down the Chapada da Contagem flanks, forming a series of rapids known lo- cally as Mumunhas. The specimens were collected in gallery forest at the borders of one of the rapids. That spot is much more humid than the xeric ‘cerrado’ areas around it. The type locality is part of an area preserved by Brazilian Government, the Area de Protecáo Ambi- ental de Cafuringa. 2.2. Leptopholcus pataxo Huber, Pérez & Baptista, new species (Figures 4-5, 10-11, 19-23) Types. Male holotype, seven female paratypes and 2 ju- veniles from Gentio do Ouro, Toca do Encantado (11°25’S, 42°30’W), Bahia, Brazil; Nov. 10, 2002 (R.L.C. Baptista, A.P.L. Giupponi), in MNRJ (04147- 48). Etymology. The species name honours the Pataxó- Hahahai Indians, formerly one of the largest Indian tribes in Bahia state. Only a few isolated communities remain, mainly due to extermination campaigns to clear land for cacao plantations. The epithet is a noun in ap- position. Diagnosis. Distinguished from other New World spe- cies by the shapes of procursus, bulbal apophyses, and the internal female genital sclerites (Figs. 19, 20, 22, 23). Male (holotype). Total length 3.2 (3.4 with clypeus), carapace width 1.13. Leg 1: 8.7 + 0.5 + 8.3 + 15.3 (tar- sus missing), leg 2 missing, tibia 3: 2.9, tibia 4: 4.1; tibia | L/d: 85. Habitus as in Figures 4 and 5. Prosoma pale ochre-grey with pair of brown marks dorsally and brown mark on clypeus, sternum light brown except frontally; legs ochre-yellow, patellae and tibia-metatar- sus joints brown; abdomen monochromous ochre-grey, ventrally with pair of brown marks in booklung-area. Ocular area slightly elevated (Fig. 5), with brush of hairs between triads, thoracic furrow absent; distance PME-PME 375 um; diameter PME 105 um; distance PME-ALE 25 um; AME-AME 15 um, AME diameter 20 um. Clypeus unmodified. Sternum wider than long (0.75/0.55), unmodified. Chelicerae as in Fig. 21, with distal apophyses carrying four modified hairs each, with some macrosetae on each side near the apophyses, proximally with slightly sclerotized projections. Palps as in Figures 19 and 20; coxa unmodified, trochanter with unsclerotized retrolateral hump and long retrola- tero-ventral apophysis with characteristic modified hair distally (cf. fig. 105 in Huber 2000); femur with large ventral protrusion and two distinct retrolateral apophy- ses; procursus relatively simple except distally, with capsulate tarsal organ; bulb with proximal sclerotized element connected to small uncus(?), with complex ap- pendix and slightly sclerotized long embolus (Fig. 19). Retrolateral trichobothrium of tibia 1 at 7%; tibia | without prolateral trichobothrium (present in all others): legs without spines, curved hairs, and vertical hairs; tar- sus 3 with about 7 fairly distinct pseudosegments. Ab- domen posteriorly not pointed dorsally (Fig. 5). Female (paratype). In general similar to male, but ab- domen thicker, clypeus barely darker. Tibia 1 in 4 fe- males: 5.5-6.8. External genitalia protruding but very simple externally (Figs. 10, 11), apparently without knob-like structure. Internal genitalia with complex sys- tem of distinctive sclerites and folds (Figs. 22, 23). Distribution. Known only from type locality. Remarks: The collecting area is located at the border of a small stream, the Encantado river, at less than 15 km from the Sao Francisco river. It is a small rocky gorge with the stream flowing through it and forming isolated ponds. Most of the area between the Sao Francisco river and Toca do Encantado is covered by a mixed vegeta- tion of palms, trees and shrubs over a sandy and damp soil. The higher areas after Toca do Encantado are cov- ered by a transitional xeric vegetation of ‘cerrado’- ‘caatinga’, over a rocky soil. All the Leptopholcus specimens were collected from webs placed against the underside of big boulders at the stream borders. The spiders were closely pressed against the rock, in a man- ner similar to other species of Leptopholcus that live on the underside of leaves. A search for Leptopholcus on the underside of leaves was made in the area, but no specimen was found. However, only a few bushes, palms and trees with small leaves cover the stream bor- ders, which would not offer an adequate substrate for Leptopholcus. 2.3. Leptopholcus evaluna Huber, Pérez & Baptista, new species (Figures 6-7, 12-13, 24-28) Types. Male holotype and one female paratype from underside of leaves along riverbed, Marigiiitar (10°26.5’N, 63%54.5"W), ~30 m a.s.l., Sucre, Venezuela: Nov. 29, 2002 (B. A. Huber), in MFLS. Etymology. Named for Isabel Allende’s Venezuela- born protagonist and fantastic story-teller. The epithet is a noun in apposition. Diagnosis. Distinguished from other New World spe- cies by the very long male palpal trochanter apophysis. and by the shapes of procursus, bulbal apophyses, and internal female genital sclerites (Figs. 24, 25, 27, 28). 104 Bonner zoologische Beitráge 53 (2004) Figs. 19-23: L. pataxo, left male palp in prolateral (19) and retrolateral (20) views, male chelicerae (21), and cleared female geni- talia in ventral (22) and dorsal (23) views. a: appendix, b: bulb, e: embolus, p: procursus, t: trochanter-apophysis, u: uncus, asterisk: proximal sclerotized element of bulb. Scale lines: 0.5 mm (19, 20, 22, 23), 0.3 mm (21). 105 Bernhard A. HUBER, Abel PÉREZ & Renner L. C. BAPTISTA: Leptopholcus in Continental America Figs. 24-28: L. evaluna, left male palp in prolateral (24) and retrolateral (25) views, male chelicerae (26), and cleared female genitalia in ventral (27) and dorsal (28) views. a: appendix, b: bulb, e: embolus, p: procursus, t: trochanter-apophysis, u: uncus, asterisk: proximal sclerotized element of bulb, arrow points to transparent projection of bulb. Scale lines: 0.3 mm (24, 25, 27, 28), 0.2 mm (26). 106 Bonner zoologische Beitráge 53 (2004) Male (holotype). Total length 2.95 (3.05 with clypeus), carapace width 0.77. Leg 1: 26.8 (6.5 + 0.4 + 6.4 + 12.4 + 1.1), tibia 2: 4.1, tibia 3: 2.5, tibia 4: 3.6; tibia 1 L/d: 96. Habitus as in Figures 6 and 7. Entire animal very pale whitish (in live rather greenish), with pair of dark dots on carapace, patellae and tibia-metatarsus joints slightly darkened, abdomen dorsally with two pairs of indistinct dark marks. Ocular area barely elevated (Fig. 7), only triads on low elevations, thoracic furrow absent; distance PME-PME 195 um; diameter PME 70 um; dis- tance PME-ALE 20 um; AME-AME 20 um, AME di- ameter 30 um. Clypeus unmodified. Sternum wider than long (0.55/0.45), unmodified. Chelicerae as in Fig. 26, with distal apophyses carrying two (or three?) modified hairs each, with two macrosetae on each side near the apophyses, proximally with unsclerotized projections. Palps as in Figures 24 and 25; coxa unmodified, tro- chanter with short retrolateral apophysis and very long retrolatero-ventral apophysis with characteristic modi- fied hair distally (cf. fig. 105 in Huber 2000), femur with distinct retrolateral apophysis, procursus relatively simple, with capsulate tarsal organ, distally more com- plex with various membranous structures, bulb with proximal sclerotized element, sclerotized uncus and ap- pendix and membranous embolus with distal transparent fringes, with additional slender transparent projection (arrow in Fig. 24). Retrolateral trichobothrium of tibia | at 8%; tibia 1 without prolateral trichobothrium (present on other legs); legs without spines, curved hairs, and vertical hairs; tarsus | apparently with >20 pseudoseg- ments, but very indistinct and difficult to see, only dis- tally about 5 fairly distinct. Abdomen _ posteriorly slightly pointed dorsally (Fig. 7). Female. In general similar to male, but without dark spots on carapace. Tibia 1: 5.1. External genitalia very simple, slightly protruding (Figs. 12, 13), with distinct knob at posterior rim (Fig. 27), with small oval pore plates (Fig. 28). Distribution. Known only from type locality. Acknowledgements. We thank Osvaldo Villarreal, Celsa Señaris and Daniel Lew for their support in obtaining a collecting permit in Venezuela and for the loan of speci- mens, the Dirección General de Fauna y Oficina Nacional de Diversidad Biológica in Caracas for issuing permit N° 01-11-0966, and the Deutsche Forschungsgemeinschaft for financing a collecting trip to Venezuela (DFG grant HU 980/1-1). Venezuelan results are part of the project "Ex- pedición Aracnológica a Venezuela", between the Zoologi- cal Research Institute and Museum Alexander Koenig, the arachnological laboratory of the National Museum of the Federal University of Rio the Janeiro and the Natural His- tory Museum of the "La Salle" Foundation, Caracas. Thanks are also due to Pedro Gnaspini and FAPESP for funding the "Expedicáo Amblypygi" to Bahia State, Brazil, to Alessandro Giupponi, Amazonas Chagas Jr. and Barbara Segal for collecting specimens, to Frederico Araujo Ramos for providing information about the type locality of L. brazlandia and to Adriano B. Kury for the loan of the Bra- zilian specimens. REFERENCES BRIGNOLI, P. M. (1980): Sur le genre Leptopholcus Simon, 1893 (Araneae, Pholcidae). Revue de zoologie afri- caine 94(3): 649-655. DEELEMAN-REINHOLD, C. L. (1986): Studies on tropical Pholcidae II. Redescription of Micromerys gracilis Bradley and Calapnita vermiformis Simon (Araneae, Pholcidae) and description of some related new spe- cies. Memoirs of the Queensland Museum 22(2): 205- 224. DEELEMAN-REINHOLD, C. L. & DEELEMAN, P. R. (1983): Studies on tropical Pholcidae I. Panjange, a new genus of Indo-Australian leaf- and rock-dwelling pholcid spiders (Araneae). Zoologische Mededelingen 57(14): 121-130. DEELEMAN-REINHOLD, C. L. & PRINSEN, J. D. (1987): Mi- cropholcus fauroti (Simon) n. comb., a pantropical, synanthropic spider (Araneae: Pholcidae). Entomolo- gische berichten, Amsterdam 47(5): 73-77. DEELEMAN-REINHOLD, C. L. & VAN HARTEN, A. (2001): Description of some interesting, new or little known Pholcidae (Araneae) from Yemen. Pp. 193-207 in: ISHWAR PRAKASH (ed.) Ecology of Desert Environ- ments. Scientific Publishers, India, Jodhpur. HUBER, B. A. (1995): Copulatory mechanism in Holocne- mus pluchei and Pholcus opilionoides, with notes on male cheliceral apophyses and stridulatory organs in Pholcidae (Araneae). Acta Zoologica (Stockholm) 76(4): 291-300. HUBER, B. A. (2000): New World pholcid spiders (Ara- neae: Phoicidae): a revision at generic level. Bulletin of the American Museum of Natural History 254: 1- 348. HUBER, B. A. (2001): The pholcids of Australia (Araneae; Pholcidae): Taxonomy, biogeography, and relation- ships. Bulletin of the American Museum of Natural History 260: 1-144. HUBER, B. A. & PÉREZ G., A. (1998): Leptopholcus delica- tulus (Araneae, Pholcidae) is a valid name. Journal of Arachnology 26: 251-256. IRIE, T. (1999): A new species of the genus Leptopholcus (Araneae: Pholcidae) from the Yaeyama Islands, Ja- pan. Acta arachnologica 48(1): 37-39. MILLOT, J. (1946): Les pholcides de Madagascar (Aranéi- des). Mémoirs du Musée National d’ Histoire Naturelle (NS)22(3): 127-158. PEREZ G., A. & HUBER, B. A. (1999): Metagonia debrasi n. sp., the first species of the genus Metagonia Simon in Cuba (Pholcidae, Araneae). Revue arachnologique 13(4): 69-72. PETRUNKEVITCH, A. (1929): The spiders of Porto Rico. Transactions of the Connecticut Academy of Arts and Sciences 30: 1-158. SIMON, E. (1893): Histoire Naturelle des Araignées. 2° édi- tion, vol. 1(2). Paris. Bernhard A. HUBER, Abel PÉREZ & Renner L. C. BAPTISTA: Leptopholcus in Continental America 107 Authors” addresses: Bernhard A. HUBER (correspon- ding author), Zoological Research Institute and Museum Alexander Koenig, Adenauerallee 160, 53113 Bonn, Germany, b.huber.zfmk@uni-bonn.de; Abel PÉREZ Gon- zález & Renner L. C. BAPTISTA: Laboratorio de Aracno- logía, Departamento de Invertebrados, Museu Nacional, Universidade do Brasil, Quinta da Boa Vista, Rio de Ja- neiro, Brazil, CEP 20940-040, baptistr@mn.ufrj.br abelpg«mn.ufrj.br. i tn 7 u *. o | L . » 7 . 7 I 7 ma 4 i ’ 7 A x wiv = ha 1 , ; a . y . ' + u \ . o ñ ! ‘ % . + ay 1 „ 1 ony i 4 Bonner zoologische Beiträge Band 53 (2004) | Heft 1/2 | Seiten 109-110 Bonn, Juni 2005 First Record of Hyles dahlii (Geyer, 1827) (Lepidoptera: Sphingidae) from the African Mainland ' Anna K. HUNDSDÖRFER, Dresden The hawkmoth species Hyles dahlii (Geyer, 1827) is considered to be restricted to Corsica, Sardinia and the Balearic Islands (REBEL 1934), although there are occa- sional reports from Sicily (PITTAWAY 2004). In addi- tion, it has also been found on the Catalan coast, in northeastern Spain, in 1975 (MASO PLANAS et al. 1979) and on the island of Pianosa, off the Tuscan coast (Italy) in 1998 (DAPPORTO et al. 1999). Here I report the discovery of about 15-20 first instar (L1) larvae of H. dahlii feeding on Euphorbia paralias in northern Tunisia in October 2004. The exact locality is the beach at Residence Shiri, a small hotel and camp- ing site on the coast north of Sejane between Cap Serrat and Bizerte. I collected and reared about a dozen indi- viduals. After two moults it was possible to determine them as A. dahlii (a caterpillar is illustrated in Fig. 1 and an adult in Fig. 2). This species has to date not been reported from Africa. Voucher specimens will be depos- ited at the Zoologisches Forschungsinstitut und Museum Alexander Koenig (Bonn) and the Natural History Mu- seum (London). I doubt that HA. dahlii is already established on the Afri- can mainland. I observed very few habitats for spurge- feeding hawkmoths in Tunisia, as I found little food plant in the parts of the country I visited. During my one-week tour (from the northern coast due south near the Algerian border, then back north along the eastern coast), I found herbaceous Euphorbia only along one inland stretch of road; otherwise there was only £. paralias along the coast. In the north, there are few sandy coastlines and these are often short and very iso- lated, with stretches of rocky coast and cultivated pine forest surrounding them. In parts of the eastern coast, the air is so humid that water condenses on the plants and the sand becomes baked onto them, so that they do not appear to be easily eatable by caterpillars. Also, the beaches are often partly or wholly developed for the tourist industry and thus it is possible that the popula- tions of E. paralias have become relictual or destroyed due to human activities. 1 found no other Hy/es caterpil- lars at the beach of Residence Shiri, or any indications 1 Clas Michael Naumann zu Kónigsbrúck (26.06.1939 — 15.02.2004) zum Gedenken of prior occurrence, such as droppings or nibbled plant sprouts. In contrast, on other beaches in Tunisia, and one especially on the north coast, as well as at one local- ity inland, I found numerous caterpillars of H. tithymali, but I did not find any other H. dahlii caterpillars. There- fore, I consider this first record of H. dahlii from the Af- rican mainland is probably the result of a rare dispersion event, possibly of only a single fertilised female from Sardinia, which lies geographically nearest to the Tuni- sian locality. Fig. 1: Fifth instar larva of Hyles dahlii found on Euphorbia paralias in Tunisia in October 2004 (feeding in captivity on £. myrsinites). 110 Bonner zoologische Beitráge 53 (2004) a cs Fig. 2: An adult moth of Hyles dahlii, emerged from the pupa in February 2005. The individual was collected as first instar larva in Tunisia in October 2004 and reared in captivity. Acknowledgements. I would like to thank Michael Korn (Konstanz) for this successful joint collecting trip to Tuni- sia and his specific help in looking out for Euphorbia plants and Hyles caterpillars. Special thanks also go to Paolo Mazzei (Rome), who first determined these larvae as H. dahlii on the basis of the single specimen I gave to him during my return journey. I am indebted to Ian J. Kitching (London) for his correction of my language and for very helpful suggestions to this note. I would also like to ex- press gratitude to the late Clas Naumann for his long-term interest and participation in the collection of Hyles samples for our joint studies with Ian J. Kitching and Michael Wink (Heidelberg). REFERENCES DAPPORTO, L., CECCHI, B., LO CASCIO, P. & SFORZI, A. (1999): Contributi alla conoscenza dell'artropodofauna dell'Isola di Pianosa (Arcipelago Toscana). II. Prima nota sui macrolepidotteri (Insecta Lepidoptera). Bol- lettino della Societa Entomologica Italiana 131: 245- 232. MASO PLANAS, A., PEREZ DE-GREGORIO, J. & SIMO CA- STELS, L. L. (1979): Primera cita d'Hyles dahlii (Lep. Sphingidae) al continent Europeu. Treballs de la So- ciétat Catalana de Lepidopterologia 2: 11-17. PITTAWAY, A. R. (2004): Sphingidae of the Western Palae- arctic: Hyles dahlii (Geyer, 1827). http://tpittaway.tri- pod.com/sphinx/h_dah.htm REBEL, H. (1934): Lepidopteren von den Balearen und Pi- tyusen. Deutsche Entomologische Zeitschrift Iris 48: 122-138. Author’s present address: Anna K. HUNDSDORFER, Staatliches Naturhistorische Sammlungen, Museum fiir Tierkunde — Zoological Museum, DNA — Laboratory, Königsbrücker Landstr. 159, D-01109 Dresden, Germa- ny, anna.hundsdoerfer@snsd.smwk.sachsen.de Bonner zoologische Beitrage Band 53 (2004) | Heft 1/2 Seiten 111-114 Bonn, Juni 2005 A New Genus and Species of Scorpion from Afghanistan (Scorpiones, Buthidae)' Wilson R. LOURENCO Département de Systématique et Evolution, Section Arthropodes (Arachnologie), Museum national d’Histoire naturelle, Paris, France Abstract. A new genus and species of a psammophilic buthid scorpion, Afghanobuthus naumanni gen. n., sp. n., are de- scribed on the basis of one female specimen collected by Professor Clas Naumann on October 1971 in Afghanistan. The specimen was collected in an extremely arid zone (400 m), at Vic Shiberghan, Dasht-e-Leili on the North range of the country. 1. INTRODUCTION In a recent publication, FET et al. (2001) call attention to the important and interesting diversity of psammophilic scorpions, from the Palaearctic deserts of central and southern Asia. In this contribution, the authors de- scribed a new genus and species of a buthid scorpion from the Baluchistan Province of Iran. They also in- cluded a very complete table of characters for six buthid psammophilic genera, namely, Anomalobuthus Kraepe- lin, 1900, Liobuthus Birula, 1898, Plesiobuthus Pocock, 1990, Psammobuthus Birula, 1911, Pectinibuthus Fet, 1984 and Polisius Fet, Capes & Sissom, 2001. During his field trips to Afghanistan on the early 1970s Professor Clas Naumann collected a few scorpions, which were subsequently sent to Prof. Max Vachon in Paris. After examination, Vachon determined two specimens as Compsobuthus sp. A recent study of these two specimens revealed that the one collected in the eastern region of the country, S of Kunduz (Qonduz), NE of Pul-I-Khumri (Poi-e-Khomri), on April 7, 1972 was indeed a species of Compsobuthus. This specimen was described as a new species, Compsobuthus tofti Lourenco, 2001 (LOURENCO 2001a). Careful examination of the second specimen revealed that it was not a Compsobuthus, but rather a psammophilic element. From the table of characters proposed by FET et al. (2001), it appears that the specimen from Afghani- stan shares some common characters with the listed genera. It shows, however, a combination of different characters, and this leads me to describe here a new ge- nus and species. This new scorpion was collected by Professor Clas Naumann in October 1971, on the North range of the country, at Vic Shiberghan, Dasht-e-Leili. 1 In commemoration of Clas Michael Naumann zu Königsbrück (26.06.1939 — 15.02.2004) 2. TAXONOMIC TREATMENT Afghanobuthus gen. n. Diagnosis. Small sized scorpions, 27.3 mm in total length. Colouration of a generally pale yellow without any spots or pigmented zones on the body and its ap- pendages. Dentate margins on movable and fixed fin- gers of pedipalp chela composed of 11/12 almost linear rows of granules. Absence of inner and outer denticles on both fingers. Pectinal tooth count 24-25. Chelicerae with one basal denticle on the fixed finger flattened in its apex; basal denticles of the movable finger absent. Anal arc with 3 lateral lobes. Trichobothrial pattern A-f (Beta). Derivatio nominis: After Afghanistan, its country of origin. Relationships: From its general morphology, the new genus Afghanobuthus gen. n. appears to have some relation- ships with other psammophilic genera present in the Palaearctic deserts, but also in North African deserts. It shares with other genera such as Polisius, Anomalobuthus or Plesiobuthus an orthobothriotaxy pattern for trichobothria. It also shares with Pectinibuthus, Anomalobuthus and the Somalian genus Sabinebuthus Lourenco, 2001 (LOURENCO 2001b) a very slender chela with rows of granules essentially straight. The new genus can, how- ever, be readily distinguished from other psammophilic genera, by a combination of characters: (1) Basal denti- cles of chelicera movable finger absent. (11) Absence of inner and outer accessory denticles on pedipalp chela fingers. (111) Sternum pentagonal. (iv) Small size. Type species 4f2hanobuthus naumanni sp. n. Description: Afghanobuthus naumanni sp. n. (Figs. 1-8) Diagnosis: as for the new genus. Type material: | female holotype. Afghanistan, North range, Vic Shiberghan, Dasht-e-Leili, 400 m, X/1971 Bonner zoologische Beitráge 53 (2004) . ... 99900070 „nooe® e ven... 0 e... yo e. eo 9900 Beecveee ce .. e o o o o e e o o o o o e Figs. 1-8: Afghanobuthus naumanni gen. n., Sp. N. Female holotype. 1. Chelicer ct, s most linear rows of granules. 3. Metasomal segment V and telson, lateral aspe dal spurs. 5. Sternum and genital operculum. 6-7. Trichobothriotaxy. 6. Chela, Femur, dorsal aspect. a, 2. Movable finger of pedipalp chela with al- howing anal arc. 4. Leg IV with tibial and pe- dorso-external aspect. 7. Patella, dorsal aspect. 8. Wilson R. LOURENGO: A New Genus and Species of Scorpion from Afghanistan 113 Fig. 9: Map of Afghanistan showing the type locality of the new species. (C. Naumann leg.). Deposited in the collection of the ‘Muséum national d’ Histoire naturelle’, Paris. Patronym: In honour of the late Professor Clas M. Naumann, Zoologisches Forschungsinstitut und Mu- seum Alexander Koenig (ZFMK), Bonn (Germany), who collected the specimen and contributed signifi- cantly to the scientific infrastructure of Afghanistan. Description based on female holotype (Morphometric measurements in Table 1). Table I: Morphometric values (in mm) of the female holotype of Afghanobuthus naumanni gen. n., sp. n. Total length 27.3 Carapace: - length 35 - anterior width 2.4 - posterior width 3:1 Metasomal segment I: - length 2a - width 1.9 Metasomal segment V: - length 3.9 - width le - depth 1.6 Vesicle: - width 1.2 - depth 2 Pedipalp: - Femur length 3.1 - Femur width 0.9 - Patella length 4.1 - Patella width 1:3 - Chela length 6.4 - Chela width 1.1 - Chela depth 1.1 Movable finger: - length 4.6 Colouration. Generally pale yellow without any spots or pigmented zones on body and appendages. Prosoma: carapace yellowish; only eyes surrounded by black pigment. Mesosoma: yellowish. Metasoma: all seg- ments yellowish. Vesicle yellowish; aculeus yellowish at base and light reddish at extremity. Venter pale yel- low. Chelicerae yellowish; teeth light reddish. Pedi- palps: yellowish overall; rows of granules on dentate margins of fingers very slightly darker. Legs yellowish. Morphology. Prosoma: anterior margin of carapace weakly emarginate. Carapace carinae weakly to moder- ately developed; anterior median carinae moderate to 114 Bonner zoologische Beitráge 53 (2004) weak; posterior median carinae moderate; central lateral moderate to weak; central median vestigial; posterior median carinae not terminating distally in spinoid proc- ess. Intercarinal spaces somewhat moderately granular laterally; almost smooth centrally. Median ocular tuber- cle slightly anterior to centre of carapace; median eyes separated by 2.5 ocular diameters. Three pairs of lateral eyes. Mesosoma: Tergites I-VI tricarinate. Lateral cari- nae on I-VI moderate to weak, granular; each carina terminating distally in very small spinoid process that extends very slightly beyond posterior margin of tergite. Median carinae on I weak; on II-VI moderate, to weak, terminating slightly distally on each segment with spi- noid process that extends very slightly beyond posterior margin of tergite. Tergite VII pentacarinate, with lateral pairs of carinae moderate to strong; median carinae pre- sent on proximal half, moderate. Intercarinal spaces weakly granular, almost smooth. Sternites: absent from sternites IH-VI; Sternite VII with 4 carinae moderate to weak. Pectines moderately long; pectinal tooth count 24-25. Metasoma: Segments I-II with 10 carinae, crenu- late; III-IV with 8 carinae. Dorsolateral carinae moder- ate, without spinoid denticles. Ventral submedian cari- nae moderate on segments I-IV. Segment V with 5 carinae; ventromedian carinae moderate to strong. Dor- sal furrows of all segments moderately to weakly devel- oped, smooth; intercarinal weakly granular almost smooth. Telson weakly granular, almost smooth, with only some granules ventrally; aculeus weakly curved, and slightly longer than vesicle. Subaculear tubercle ab- sent. Chelicerae: Basal denticles of movable finger ab- sent; basal denticles of fixed finger flattened at apex (Vachon, 1963). Pedipalps: Trichobothrial pattern or- thobothriotaxic, type A (Vachon, 1974); dorsal trichobothria of femur in f (beta) configuration (Va- chon, 1975). Femur pentacarinate; all carinae moder- ately crenulate. Patella with 8 carinae; all carinae weak; dorsointernal carinae with one slightly spinoid granule. Chelae slender, with elongated fingers; all carinae weak to vestigial. Dentate margins on movable and fixed fin- gers composed of 11/12 almost linear rows of granules. Legs: Ventral aspect of tarsi with one row of thin setae. Tibial spurs present on legs III-IV but reduced; pedal spurs present on all legs, moderate to strong. Acknowledgements. | am very grateful to J. Köhler (ZFMK, Bonn) for inviting me to participate in this publi- cation and to B. Sinclair (ZFMK, Bonn) for reviewing the English style of preliminary drafts of the manuscript. REFERENCES FET, V., CAPES, E.M. & Sissom, W. D. (2001): A new ge- nus and species of Psammophilic scorpion from east- ern Iran (Scorpiones: Buthidae). Pp. 183-189 in: FET, V. & SELDEN, P. A. (eds.) Scorpions 2001, In Memor- iam Gary A. Polis. British Arachnological Society, London. LOURENCO, W. R. (2001a): A new species of Compsobut- hus Vachon, 1949 from Afghanistan (Scorpiones, Buthidae). Entomologische Mitteilungen aus dem zo- ologischen Museum Hamburg 13(164): 315-319. LOURENGO, W. R. (2001b): Un nouveau genre de Buthidae, probable vicariant géographique d’Anomalobuthus Kraepelin (Chelicerata, Scorpiones). Biogevgraphica 77(1): 15-20. VACHON, M. (1963): De l'utilité, en systématique, d'une nomenclature des dents des chéliceres chez les Scor- pions. Bulletin du Muséum National d’Histoire Natu- relle Paris 2e sér. 35(2): 161-166. VACHON, M. (1974): Etude des caracteres utilisés pour classer les familles et les genres de Scorpions (Arach- nides). 1. La trichobothriotaxie en arachnologie. Sigles trichobothriaux et types de trichobothriotaxie chez les Scorpions. Bulletin du Muséum National d’Histoire Naturelle Paris 3e sér., n° 140, Zool. 104: 857-958. VACHON, M. (1975): Sur l'utilisation de la trichobothrio- taxie du bras des pédipalpes des Scorpions (Arachni- des) dans le classement des genres de la famille des Buthidae Simon. Compte Rendu hebdomadaire des Séances de l’Academie des Sciences Paris, ser. D 281: 1597-1599. Author’s address: Wilson R. LOURENCO, Departement de Systematique et Evolution, USM 0602, Section Arthropodes (Arachnologie), Museum national d’His- toire naturelle, CP 053, 61 rue Buffon 75005 Paris, France, e-mail: arachne@mnhn. fr Bonner zoologische Beitráge Band 53 (2004) | Heft 1/2 | Seiten 115-119 3onn. Juni 2005 Two New Species of Eoophyla Swinhoe, 1900 from Continental South East Asia (Lepidoptera: Crambidae, Acentropinae)' Wolfram MEY’ & Wolfgang SPEIDEL” ; ’Museum für Naturkunde, Humboldt Universität, Berlin, Germany > Zoologisches Forschungsmuseum Alexander Koenig, Bonn, Germany Abstract. Eoophyla clasnaumanni Speidel & Mey sp. nov. from North Thailand and Eoophyla myanmarica Mey & Speidel sp. nov. from North-west Myanmar (Burma) are described. These new species are closely related to each other and to Eoo- phyla aureolalis (Snellen, 1876) from Sundaland. Differential characters and figures of all three species are provided. 1. INTRODUCTION The species of Eoophyla Swinhoe, 1900 are currently placed in a number of species groups. SPEIDEL & MEY (1999) recognised six groups including an assemblage of nine isolated species which could not be assigned then to one of the defined groups. The study of the Phil- ippine species of the genus resulted in the establishment of three further groups (SPEIDEL 2003). The definition of the groups are based on external characters. Espe- cially the number and arrangement of the eye-spots on the hindwings are used as a simple and discernable trait. It has proved to be a practical way in sorting species and to cope with the increasing species numbers. The spe- cies treated in this paper belong to the crassicornalis group of species (SPEIDEL & MEY 1999). Since the phy- logenetic value of this and other characters is unclear yet, the ordering group system of the genus should be regarded as provisional. The Acentropinae collection in the ZMHB and the col- lection of W. Speidel contain a number of undescribed species. In the crassicornalis — group Eoophyla aureo- lalis (Snellen, 1876) is the only species hitherto known with a dark spot in the anal field of the forewings. The species is distributed in Sundaland. The examination of specimens with the same wing pattern from Thailand and Myanmar showed that they are related to E. aureo- lalis but represent separate species. They are described in the present paper. Abbreviation: ZMHB = Museum fiir Naturkunde, Humboldt Universitat, Berlin 1 In commemoration of Clas Michael Naumann zu Königsbrück (26.06.1939 — 15.02.2004) 2. DESCRIPTION OF SPECIES Eoophyla clasnaumanni Speidel & Mey, sp. nov. Material: Female holotype (genitalia slide: Mey 10/04) and 1 male paratype (genitalia slide: Mey 11/04), North Thailand, Nan Province, Pua, Doi Phu Kha, 1600m, km 35, 16. — 20. 2. 1993, lux, leg. W. Speidel, deposited in ZMHB (holotype) and coll. Speidel (paratype). Description (Fig. | male, fig. 2 female): Length of forewing 12 mm (male) — 14 mm (female). Head and thorax yellow-white, scape of male with curved proces- sus and enlarged antennal flagellomeres. Tegulae with brown spots. Legs yellow, tarsal segments of foreleg brown at the articulations; femora and tibia of forelegs dark-brown on inner side; male hindfemora with scale- brush. Forewing predominantly yellow except the slightly curved, antemarginal white band which is darkly lined on its margins; frontside of costa with dark line; cell between Cu 1+2 and A 1+2 filled with white in the centre, but not lined; disco-cellular bar reduced to a brown, rounded spot at the base of M2 and M3: a simi- lar brown spot in the anal field between anal vein and wing margin. Cu 2 and A 3 present with their basal stems. Male with tuft of broad scales on R before bifur- cation with R 5. Hindwing with subapical incision, strongly developed in male, weaker in female: antemar- ginal white band short, just reaching the marginal spots: short yellow band at the base of the hindwing present: two small marginal, oval, white spots, lined with black scales. Male genitalia (Fig. 9): General structure as usual in the genus; valvae long, with 3 apical, caudally directed setae. Gnathos slightly shorter than uncus, with 5 subapical, dorsal teeth. Aedeagus without cornuti. Female genitalia (Fig. 6): Ninth segment shorter than apophyses posteriores; ductus bursae long, beginning with a wide antrum; colliculum laterally sclerotised. 116 Bonner zoologische Beitráge 53 (2004) Fig. 1-5: Adult specimens Fig. 1: Eoophyla clasnaumanni Speidel & Mey sp. nov. Male, paratype. Forewing length 12 mm Fig. 2: Eoophyla clasnaumanni Speidel & Mey sp. nov. Female, holotype. Forewing length 14 mm Fig. 3: Eoophyla myanmarica Mey & Speidel sp. nov. Female, holotype. Forewing length 13 mm Fig. 4, 5: Eoophyla aureolalis (Snellen, 1876) (Indonesia, Sumatra, Sumatera Utara, Prapat, 2°46’ N 98°59’ E, log- ging trail 2, 1050 m, 1.-30.12.1994, lux, leg. E.W. Diehl Sumatra), fig. 4 male (Forewing length 9 mm), fig. 5 fe- male (Forewing length 12 mm) Wolfram Mey & Wolfgang SPEIDEL: Two New Species of Eoophyla Swinhoe Fig. 6-9: Genitalia Fig. 6: Eoophyla clasnaumanni Speidel & Mey sp. nov. Female holotype. Fig. 7: Eoophyla myanmarica Mey & Speidel sp. nov. Female Paratype. Fig. 8: Eoophyla aureolalis (Snellen, 1876). Female. Fig. 9: Eoophyla clasnaumanni Speidel & Mey sp. nov. Male paratype. , 1900 from Continental South East Asia 117 118 Bonner zoologische Beitráge 53 (2004) Corpus bursae elongate, club-shaped. Signum consists of small teeths, arranged in one short and one long pair of elongate bands, which are situated on the ridge of the somewhat flattened corpus bursae. Etymology: The species is named in memory of the late Clas M. Naumann, our distinguished lepidopterist col- league and adviser, who always supported and promoted our studies on Lepidoptera and who passed away far too early. Relationship: The wing pattern of E. clasnaumanni sp. nov. is similar to that of E. myanmarica sp. nov. The shape of the corpus bursae and the structure of the sig- num in the female genitalia are also very similar. They differ, however, in the length of the ninth segment and in the structure of the antrum, being short and wide in E. clasnaumanni sp. nov. and long and narrow in E. myanmarica sp. nov. Both species probably form a spe- cies pair. According to female genitalic structures a re- lated species is E. simplicialis sensu Yoshiyasu, 1987, recorded from Thailand. Eoophyla myanmarica Mey & Speidel, sp. nov. Material: Female holotype (genitalia slide: Mey 12/04) Myanmar, Chin, Tiddim, Thaing-gnin, 2310 m, 25.5.2001, lux, leg. S. Naumann, deposited in ZMHB. Paratypes: 3 females, same locality, 2200 m, 17.— 2.5.2001, lux, leg. S. Naumann, deposited in ZMHB and coll. Speidel. Description (Fig. 3): Length of forewing 12 mm- 13 mm (females). Head and thorax yellow-white, Tegulae with brown spots. Patagia white. Legs yellow, tarsal segments of foreleg brown at the articulations; femora and tibia of forelegs dark-brown on inner side. Fore- wing predominantly yellow except the slightly curved, antemarginal white band which is darkly lined on its margins; frontside of costa with dark line; cell between Cu 1+2 and A 1+2 filled with white in the centre, but not lined; disco-cellular bar light brown, but its apex dark brown forming a rounded spot at the base of M2 and M3; a similar brown, slightly elongate spot in the anal field between anal vein and wing margin. Cu 2 and A 3 present with their basal stems. Hindwing with subapical incision, antemarginal white band short, just reaching the marginal spots; short yellow band at the base of the hindwing present; two small marginal, oval white spots, lined with black scales. Female genitalia (Fig. 7): Nineth segment as long as the apophyses posteriores; ductus bursae long, begin- ning with a narrow and long antrum; colliculum later- ally sclerotised. First part of corpus bursae beyond col- liculum a long ductus, widening to a somewhat elongate, trapeze-like form, pleated at the widening. Signum consists of small teeths, arranged in one short and one long pair of elongate bands, which are situated on the ridge of the somewhat flattened corpus bursae; short pair darker than long pair. Etymology: The specific epithet is derived from Myanmar, today the name of the former Burma. Relationship: See £. clasnaumanni sp. nov. Eoophyla aureolalis (Snellen, 1876) Oligostigma aureolalis Snellen, 1876: 193 (key), 200, pl. 8, fig. 5 a-c. Type locality: Java. Material: 2 44, 2 Y Y (genitalia slide: Mey 13/04), In- donesia, Sumatra, Sumatera Utara, Prapat, 2°46’ N 98259” E, logging trail 2, 1050 m, 1.-30.12.1994, lux, leg. E.W. Diehl (coll. Speidel, ZMHB). Several other specimens, all from Sumatra, in coll. Speidel. The lectotype, a female, labeled ‘Java, Heckmeijer” was selected by Munroe et al. (1958) and is in the Leiden Museum. The lectotype and the only male paralectotype still in Leiden both lack the abdomen (Munroe et al., 1958). The coloured figure in the original description is good and fits very well to our Sumatran specimens. Description (Fig. 4 male, Fig. 5 female). Length of forewing 8 - 9 mm (males), 10 - 12 mm (females). In comparison with E. clasnaumanni sp. nov. and E. myanmarica sp. nov. the female of this species has a third, dark brown spot on the forewings. It is situated in the cell before bifurcation of Cu la and Cu 1b. The males have a long costal fold in the forewings and the cape is without basal process. The femur and tarsus of the forelegs bear another separating character. In £. aureolalis the dark brown, inner and front sides are in- terrupted by a long, yellow line. The tarsal segments are not dark at the articulations. The yellow line is lacking in the other two species. In the genitalia of the female (Fig. 8), the bursa copulatrix 1s a large, oval sac, enlarg- ing shortly behind the colliculum. Only a single, elon- gate, paired signum is present on its ridge. The antrum is wide and the ductus bursae a broad, folded tube. Relationship: The species of the crassicornalis-group exhibit a distinct sexual dimorphism. In general, the fe- males are more uniformly patterned as the males who have developed remarkable, secondary sexual charac- ters. The variety of these traits are not accompanied by appropriate differences in the genitalia. It is therefore difficult to acertain the relationships among species. The wing pattern of E. clasnaumanni sp. nov., E. myan- marica sp. nov. and E. aureolalis show a similar ar- rangement of spots. It possibly points to a closer rela- tionship of these three species. Wolfram Mey & Wolfgang SPEIDEL: Two New Species of Eoophyla Swinhoe, 1900 from Continental South East Asia 119 REFERENCES MUNROE, E., DIAKONOFF, A. & MARTIN, E. L. (1958): Catalogue of Snellen’s types of Pyralidae, with selec- tions of lectotypes. Tijdschrift voor Entomologie 101: 65-88. SNELLEN, P. C. T. (1876): Over Oligostigma Guenée, een genus der Pyraliden. Tijdschrift voor Entomologie 19: 186-209, pl. 8-9. SPEIDEL, W. & MEY, W. (1999): Catalogue of the Oriental Acentropinae (Lepidoptera, Crambidae). Tijdschrift voor Entomologie 142: 125-142. SPEIDEL, W. (2003): New species of aquatic moths from the Philippines (Lepidoptera, Crambidae). Insecta Ko- reana 20: 7-49. YOSHIYASU, Y. (1987): The Nymphulinae (Lepidoptera: Pyralidae) from Thailand, with descriptions of a new genus and six new species. Microlepidoptera of Thai- land 1: 133-184. / Authors’ addresses: Wolfram MEY, Museum fiir Naturkunde, Humboldt Universität, Invalidenstr. 43, D 10115 Berlin, Germany, e-mail wolfram.mey@mu- seum.hu-berlin.de; Wolfgang SPEIDEL (corresponding author), Zoologisches Forschungsinstitut und Museum Alexander Koenig, Adenauerallee 160, D-53113 Bonn, Germany, e-mail: w.speidel.zfmk@uni-bonn.de Seiten 121-133 _— | Bonner zoologische Beiträge Band 53 (2004) Heft 1/2 Bonn, Juni 2005 Of Phenotypes and Genotypes: Two Sides of one Coin in Taxonomy?' Bernhard MISOF, Cornelya F. C. KLUTSCH, Oliver NIEHUIS & Alexandra PAT1 Zoologisches Forschungsmuseum Alexander Koenig, Bonn, Germany Abstract. We discuss the recent proposals of DNA taxonomy and review some of the basic principles necessary to as- sess the importance of promoted ideas. We try to show that distinctness and independence of species taxa do not neces- sarily correlate, at least in newly formed species, which limits the applicability of any applied pattern analysis method. We do not intend to dismiss DNA based taxonomy, but would like to emphasize that molecular characters pose com- pletely new problems to taxonomy often not appreciated enough in the discipline. DNA taxonomy is currently promoted because of its potential for automation. We show that species identification and circumscription can not be entirely automated since the result of a species description in taxonomy is equivalent to the formulation of a locally valid hy- pothesis. Consequently, every delimitation of a species taxon is open for empirical falsification only on a local scale. It is impossible to extend locally derived criteria, a species delimitation based on criteria derived from single sister species, to a global scale without sacrifying relevant information. We show from a pragmatic perspective that specimen assign- ment based on molecular characters can be automated under restricted circumstances employing character-based ap- proaches. 1. INTRODUCTION It is certainly true that taxonomy still faces a major task. It has been estimated that only one tenth of assumed ex- isting species has been recognized and described so far. Considering the long tradition in taxonomy, which has been conducted over centuries, this is not really encour- aging. Taxonomists estimate that hot spots of biodiver- sity, tropical rain forests, and also deep sea biotas are still only fragmentarily known. The biodiversity of tropical rain forests is threatened by massive destruction and consequently eradication of largely unknown biodi- versity. In many densely populated areas, like Europe or Africa, massive anthropogenic destruction has left de- serted areas with presumably significant loss of species and biodiversity. It is comprehensible that taxonomy has to search for ways of speeding up the process of identi- fying and describing species in the face of these chal- lenges. Here surfaces the second conundrum. Taxono- mists generally feel underfunded and it appears that the profession is a vanishing one among biologists. It is hard to find students interested in taxonomic activities, | Several recently published articles advocate the incorpo- | ration of molecular techniques into taxonomic protocols (for example HEBERT et al. 2001, 2004; TAUTZ et al. 2002, 2003; STOECKLE 2003, to name just a few). For | many practicing taxonomists the incorporation of mo- lecular data seems highly desirable. Without any doubt, the incorporation of as many different character sets as possible into a species taxon description will improve the fit between species taxa and real evolutionary units. What is new in recent articles advocating DNA taxon- omy is the emphasis on molecular techniques over mor- phological approaches, which, as the authors claim, would be the only way to meet the challenges in taxon- | omy of the 21” century. This is indeed slightly contrary to the perception of taxonomic practice of many tax- onomists and aroused obvious opposition. | “Let's face it, the morphological approach has had 250 | years to advance the task, and we're only 10% of the way towards the goalposts” said HEBERT in a report on BioMedNet News. His solution, to which many taxono- mists are strongly opposed, is to distinguish between species on the basis of similarities and differences in their DNA. If taxonomists fail to embrace molecular technology, HEBERT is clear about the consequences: “There is no more likely death of a discipline than the failure to innovate.” (HEBERT, BioMedNet News, DNA: the barcode of life? 8 January 2003) | In commemoration of Clas Michael Naumann zu Königsbrück (26.06.1939 — 15.02.2004) and for several invertebrate groups specialists are al- ready absent. Consequently, training in taxonomy is not being provided, further promoting the loss of knowledge and momentum. Traditional taxonomy thrives from the life-long experience of specialists with trained intuition. The delimitation of species taxa by dedicated specialists and their intuition is often inaccessible for the outsider, making the process of species description quite mysteri- ous. There is no single character with which species taxa can be recognized across phyla. The taxonomic challenge is the discovery of the relevant individual cri- teria given a population of individuals. Consequently. taxonomists are sometimes seen outside of the realm of 122 22; Bonner zoologische Beiträge 53 (2004) natural sciences presenting themselves as book keepers of mental projections. This seemingly subjective and highly artificial approach to recognizing taxa often in- timidates the novice student. Another immanent problem is the publication process in taxonomy. Currently, species descriptions are most of- ten not placed in peer reviewed journals and do not en- joy a high standing among many biologists. It would be ill advised for students hoping for a respectable career to enter the field of taxonomy. As can easily be envis- aged, a massive brain drain away from taxonomy is the result. Among others, molecular taxonomy has been proposed to cushion this development. But what could be the advantages of using molecular techniques in taxonomy? The advocates of DNA taxonomy, as it is also called, claim that the genotype provides a universal code with which species could be automatically identified and classified. This indeed seems to draw taxonomic inves- tigation out of the realm of mystery and subjectivity. Automatic detection of species taxa based on genotypic differences would draw taxonomy out of its classroom corner and would transfer taxonomy to a technical en- terprise solely executable by machines (TAUTZ et al. 2003; GASTON & O'NEILL 2004). It is indeed true that the composition of DNA presents a universal resource of variation by the combination of only four character states. Including the phenomenon of indels (a composite word derived from insertion and de- letion), we have five clearly distinct characters states with which we can discriminate between DNA se- quences. Consequently, genotypes offer the potential of automatic detection of differences between DNA se- quences. Different to the plethora of morphological variation, this seems indeed a big advantage as it might alleviate the problem of educating taxonomic special- ists. Computers are inexhaustible co-workers doing the job while we are sleeping. Proponents of DNA taxonomy claim that the simplicity of molecular characters help to solve the problems with which current taxonomy is confronted. Specialists are not needed anymore and identification of new sequence groups can be computerized, greatly speeding up the process of describing the nine-tenths of the unknown biodiversity. It has been already put forward that the de- scription of new species must include a mandatory analysis of selected target sequences thus giving a bar- code (HEBERT et al. 2001) of every newly described taxon. To be fair, we have to note that most proponents of DNA taxonomy do not call for a complete disregard of phenotypes but instead argue for a priority of the genotype in species descriptions. Once a species taxon has been recognized based on genotypes, the corre- sponding phenotypes should be recorded afterwards completing species descriptions (HEBERT et al. 2001; TAUTZ et al. 2003). In other words, the shift from phe- notypes to genotypes could provide the answer to the current taxonomic challenges. In this essay, we will try to sort potentials and pitfalls of the molecular approach in taxonomy. We are particu- larly interested in theoretical implications, which are surprisingly often omitted in the actual discussion. We will not discuss political implications which have been put forward (LIPSCOMB et al. 2003; MALLET & WILL- MOTT 2003) and we will also refrain from discussing the developments in internet technologies which are often promoted concurrently with matters of DNA taxonomy (GODFRAY 2002; SEBERG et al. 2003). Our starting point is a taxonomy which seeks the description of evo- lutionary units, characterized by reproductive isolation. In this sense, taxonomists try to delimit species taxa by matching phenotypic and possibly genotypic classes with these evolutionary units. This prescription differen- tiates the taxonomic activity from a mere classification of biological variation for our convenience and text books. DNA taxonomists claim that the genotype is bet- ter suited for the delimitation of species taxa matching evolutionary units than any other biological properties of organisms. This proclamation is the focus of our analysis. Consequently, we will first review the princi- ples of genetic variation and variability. Second, we will spend time to review the role of species in evolution. We will not review all of the ideas once promoted but instead give a biased view on what species and speci- ation probably are and is. Taxonomy as a discipline is often surprisingly ignorant of theoretical issues behind species definitions and the process of speciation. Third, we will discuss the inherently assumed parallelism of character distinctness and population independence, which is at the theoretical foundation of taxonomy in general. But we will also try to show where DNA tax- onomy can indeed help to recover lost ground in de- scribing and (re-)identifying specimens. Finally, we will develop ideas of how to incorporate molecular and mor- phological methods in an automatic way. A final word: we do not claim to present radically new ideas about the incorporation of molecular techniques in taxonomy, but instead try to draw the reader’s attention to the limits of DNA based approaches. This is by no means intended to damn DNA taxonomy or glorify the proposals. We feel that the ongoing discussion on DNA taxonomy and tax- onomy in general is becoming unbalanced with a strong impetus towards political points of view instead of sci- entific reasoning. Bernhard Misor, Cornelya F. C. KLUTSCH, Oliver NIEHUIS & Alexandra PATT: Of Phenotypes and Genotypes 2. INTRA- VS INTERSPECIFIC VARIABILITY/VARIATION Considering genetic consequences of speciation, one has to ask how many, what kind of, and in which rate genomic organization of loci change during the process of speciation. Observed genetic differences between species may accumulate not only during or after the speciation process when populations have become inde- pendent, but also within populations in terms of intra- specific variation. In the following, we will use the term ‘variability’ to designate the differences actually present between individuals within a species. The term 'varia- tion', which is often used in a synonymous way, is used to describe differences of individuals of different spe- cies. 2.1. Intraspecific variability As DNA-Taxonomy is often conducted on mtDNA se- quences, we will concentrate on the evolutionary pat- terns of this DNA type, which is nevertheless quite similar to genomic DNA with the exception of its usu- ally maternal inheritance and its haploid occurrence. Genetic differences, as mentioned before, result from four different nucleotides, conventionally symbolized A, T, G, and C. These four character states (or rather five by counting indels) are potentially assumed at each position given enough time for substitution to occur. Genetic differentiation occurs not only between subspe- cies or species but also within populations as popula- tions are affected by mutation, migration, random ge- netic drift and natural selection (HARTL € CLARK 1997). Mutation is the source of genetic variation, which arises and increases divergence between mtDNA genotypes. The input of mutations creates and maintains a certain degree of variability within populations (AVISE 2000). Within finite populations in sexually reproducing organisms, new mutations can become fixed after some time without the effect of selection. The rate at which random genetic drift changes allele frequencies or hap- lotypes varies inversely with the effective population size (N.) (WRIGHT 1931; FISHER 1930). The smaller a population, the stronger are effects of random genetic drift and the faster the populations will loose its genetic variation. The effective population size is much smaller for mtDNA than for genomic DNA because of its hap- loid state and its generally maternal inheritance. Now imagine a population with a large distributional range. Gene flow within this population is restricted and con- sequently, it will be geographically structured. This will affect genetic differentiation between demes (defined as geographical units within which mating is effectively random). The degree of differentiation depends not only on the effective population sizes of demes but also on the pattern of gene flow among them (WRIGHT 1951). Given a finite Mendelian population, which splits into t allopatric subpopulations we can expect to see genetic differences within several loci after a certain number of generations without the force of natural selection. The consequence is genetic variability between populations driven by chance. In addition, natural selection plays a role in differentiation between populations of a widely distributed species. Let us assume the beginning of a speciation process within such a species. If populations become separated, genetic variability will be inherited to subpopulations. In this situation, haplotypes within subpopulations are potentially paraphyletic or even polyphyletic (FUNK & OMLAND 2003). Until monophyly of haplotypes will be achieved time must pass by, at least more than 4N, generations, as estimated in simula- tions (AVISE 1987). Lineage sorting refers to a related problem. A reciprocal monophyly of haplotypes will be generated after complete lineage sorting has been achieved (ENGLBRECHT et al. 2000; cf. below). Not in every case of recent speciation events do genetic distinctness and population independence go in parallel. We intuitively assume that the formation of species will arrange the total amount of genetic variation among them into non overlapping packages of genetic varia- tion. This assumption is indeed appealing, but not nec- essarily realized. DNA taxonomy has to deal with issues of population genetics, as 1t often relies on presumably selectively neutral gene loci. Paraphyly of selectively neutral haplotypes or the absence of genetic differentia- tion among those pose major problems to taxonomy and clearly restrict the applicability of molecular tools. Imagine speciation events, which lead to a clear differ- entiation of phenotypes due to, for example, sexual se- lection but without fixation of selectively neutral alleles in species taxa, because either populations had been large or there was no time yet to fix allele frequencies. The incongruence of genetic and phenotypic distinct- ness is not an artificial construction. Examples are given by TAUTZ et al. (2003) who mentioned the situation of the African great lakes cichlid fauna. In Lake Victoria, cichlids are strongly differentiated in phenotypic groups but lack a clear genetic differentiation in sequenced mi- tochondrial markers (MEYER et al. 1990, compare also TAKAHASHI et al. 2001). These considerations illustrate the limits of DNA taxonomy quite nicely; since, despite reproductive isolation, there must be enough time to build up genetic variability between populations. 2.2. Interspecific variation The degree of genetic differences between populations, subspecies, or species can be described by using genetic distance methods (e.g. NEI 1978). Dealing with a certain degree of differentiation between populations, some re- searchers are going to classify species by the percentage of genetic divergence (for example MALTAGLIATI et al. 2001; JIGGINS & DAVIS 1998). This requires a constant 124 Bonner zoologische Beitráge 53 (2004) rate of sequence differentiation at neutral loci, which 1s independent of population genetic factors mentioned above. However, the influence of small population sizes on the rate of evolution (OHTA 1976) was assessed in a study by DE SALLE & TEMPLETON (1988). They com- pared interspecific levels of sequence divergence within a Drosophila taxon, comprised of several species, sup- ‘posed to be generated by a founder event and another Drosophila taxon, again comprised of several species, which were thought to have originated from a large population. They showed that the second species group presented a much less pronounced molecular differen- tiation then the first one. Another example in which in- traspecific sequence divergence is as great as such among distinct species is given by WAYNE et al. (1990) who analyzed genetic distances of Jackal populations. In summary, the degree of genetic differences will not al- ways match the biological species status one to one. It is indeed tempting to say that there is still a rough correla- tion between genetic distance and extend of reproduc- tive isolation. If this holds true, a convention on a minimal distance measure could be successful in identi- fying new species. Alpine snails of the species Arianta arbustorum (HAASE et al. 2003) show COI haplotypes of more than 20% differences within a population. A genetic distance of 20% as a general convention to de- limit species is useless in many other taxa. This issue will be taken up later again. 3. SPECIES AND SPECIATION “The primary aims of taxonomy are to name, circum- scribe, describe and classify species” (SEBERG et al. 2003, p. 63). We probably all agree that the generation and maintenance of biodiversity is closely linked with the establishment of evolutionary units and our under- standing of the processes will rely on our ability to rec- ognize these evolutionary units. In sexually reproducing organisms, species are exactly those evolutionary units without which biological diversity is only incompletely understood. It is therefore paramount to reflect what processes lead to the formation of evolutionary units and what characteristics can be used to identify them. The discussion on species and speciation traditionally includes the following questions: 1) What are proper species definitions and what is the ontological status of species? 2) What are reproductive isolation and isolat- ing mechanisms? 3) What is the predominant mode of speciation and why are there species? We add the taxo- nomically relevant question of how species are best classified. Subsequently, we will pick up the issues and comment on them from the perspective of a taxonomist. The history of species definitions is legacy. MAYR (1982) gave a most authoritative and readable account on species definitions in taxonomy and evolutionary bi- ology. Definitions are either acceptable or not, but do not represent explanations of the more or less distinctly separate arrays of organisms (DOBZHANSKY 1937). We think that the disputes over proper species definitions are not interesting and we will not follow up this route. Secondly, we will deliberately omit the discussion whether species are classes, natural kinds, or whether species constitute individuals (for a review, see again MAYR 1982). The discussion on the ontological status of species is not important here. (However, it is clear that the question of whether species constitute natural kinds or individuals is much more interesting than dis- putes over definitions as it confronts us with a problem in need of an explanation). With the modern synthesis in evolutionary biology, species in sexually reproducing groups of organisms are seen as groups of individuals constituting the largest possible inclusive Mendelian population (DOBZHANSKY 1935, 1937). In contrast to prevailing typological species concepts, representatives of the modern synthesis shifted the emphasis from pat- terns to processes. Inclusive Mendelian populations do not exist in parthenogenetic and asexual organisms and therefore, the term species in the biological sense is not applicable for these groups of organisms. Many tax- onomists and natural scientists felt this limitation of the biological species concept as much too restrictive and opted for more inclusive concepts covering also asexu- als and parthenogenetic forms. To our knowledge, the most farfetched example is the general lineage concept of species, which applies the term population to sexual and asexual groups of organisms (DE QUEIROZ 1999). We will restrict the term population to a group of sexu- ally reproducing organisms; anything else blurres the essential differences and produces the illusion of unified theories of species. Of course, taxonomy is also dealing with asexually reproducing individuals, but for now, we will deliberately exclude asexuals and parthenogenetics from our discussion. The formulation of the biological species concept shifted the emphasis from definitorial problems to real scientific inquiry. As DOBZHANSKY put it so easily the reasoning runs as following: first is the observable pat- tern of distinctive separate arrays of organisms and sec- ondly there is the theory that these separate arrays seem to be maintained by established reproductive barriers between groups. Mayr added a new twist to this theory. He concluded, starting from the question of how charac- ters can evolve despite mixing of genotypes in sexually reproducing organisms, that the formation of Mendelian groups which are reproductively isolated against other such groups should be clearly advantageous. The new twist is the explicit inclusion of the phenomenon of se- lection albeit in a slightly diffuse way. MAYR was not the first emphasizing isolating mechanism in defining species. Credit has to go to DOBZHANSKY who intro- duced the term isolating mechanism in his book Genet- Bernhard Misor, Cornelya F. C. KLUTSCH, Oliver NIEHUIS & Alexandra PATT: Of Phenotypes and Genotypes ics and the Origin of Species (1937) (The idea of recog- nition of “proper” reproductive partners is central to species and even much older. It finds its explicit prede- cessors in Buffon (in MAYR 1982).) DOBZHANSKY (1937) reserved an entire chapter for the discussion of isolating mechanisms. He mentions that “Darwin and Lamarck pointed out that interbreeding of groups of in- dividuals, which are hereditarily distinct, results in dis- solution and swamping of the differences by crossing.” (DOBZHANSKY 1937, p. 228). He correctly notes that the statement is somewhat misleading since single allelic differences will not diffuse in crosses but segregate again in the offspring. Only distinct character com- plexes of polygenic origin can break down in crosses leaving the hybrids less competitive compared to par- ents. For DOBZHANSKY, isolating mechanisms are mechanisms, which prevent the production of hybrid zygotes reaching the reproductive stage. Most impor- tantly, citing Mayr again, isolating mechanisms are seen as “biological properties of individuals which prevent the interbreeding (fusion) of populations” (MAYR 1970). Therefore, sole geographical isolation of populations does not constitute a necessary and sufficient criterion for recognizing reproductively isolated species. The ba- sic principle behind this idea is the insight that Mende- lian populations are the units of evolution. (It is interest- ing to see that the origin of panmictic populations is not important in this respect.) Scientists like DARWIN, LA- MARCK, DOBZHANSKY, MAYR, and many more have been convinced that differential adaptation to heteroge- neous environments and ecological needs would have been impossible without the partitioning of panmictic populations. For that reason, we can call species inde- pendent modules in the evolutionary process. An analogous picture can be drawn from the field of de- velopmental biology. There it became clear after ground breaking work starting in the early 20th century (BER- TALANFFY 1932; WADDINGTON 1957) that evolvability of complex characters can only be maintained by reduc- ing pleiotropic interactions. For example, imagine genes, which effect the formation of epithelia. Specific alleles might promote the formation of a thick kerati- nous outer layer, thus reducing the risk of dehydration in dry habitats. This could clearly be an advantage for the bearer of this allele. However, thick keratinous lay- ers on bronchial epithelia are certainly counterproduc- tive as they restrict the intake of oxygen. The pleiotropic interactions prevent the fixation of an allele in this case. The situation leads to a reduced adaptability under strong pleiotropic interactions. The solution is the break up of pleiotropic interactions and the recovery of adaptability. The break up of pleiotropic interactions re- sults in the formation of genetic and developmental units, which we also can call modules. The evolution of modules is the key invention (compare for example bt Kn RIEDL 1978; BONNER 1988; RAFF 1996; WAGNER 1996; WAGNER & ALTENBERG 1996). Similarly, species can be seen as genetically independent modules protecting their harmonious gene pool and thus, maintaining distinctness. The biological species concept refers to exactly these properties of inclusive Mendelian popula- tions. Biological species are characterized by distinct- ness and independence. Whether distinctness or inde- pendence comes first is asking for the priority of the chicken or egg. MAYR himself favored for a long period allopatric speciation processes as the predominating mode of speciation, which implies that independence is acquired first and distinctness is a secondary phenome- non. MAYR’s idea predicts a linear order of phenomena from geographic isolation to reproductive isolation. However, theoretical considerations suggest that parapa- tric speciation and its derivatives (peripatric up to sym- patric speciation) are at least conceivable, which makes the decision less obvious (TURELLI et al. 2001). Polytypic species are species in which individual sub- populations can be characterized by clear phenotypic differences, often given subspecific rank. It is assumed that a reduction of gene flow between geographically distant populations maintains local differences. How- ever, most of the genotypic composition within the spe- cies is still homogeneously distributed throughout the entire range. Such a situation is also found in species with clinal variation along an ecological gradient. This even distribution of phenotypic variation can be stabi- lized by the effects of natural selection, resulting in lo- cal fixation of alleles under stabilizing selection (local adaptation). However, selectively neutral alleles will travel freely across large ranges (DIECKMAN & DOEBELI 1999; DOEBELI & DIECKMAN 2003). In such a case, par- tial independence due to restricted vagility of individu- als is driving the evolution of distinctness despite the absence of reproductive isolation. This example illus- trates that reproductive isolation is neither a sufficient nor a necessary condition for adaptive change in popula- tions. Local adaptive changes can be accomplished by the action of natural selection alone, even under para- (sym)patric conditions (DOEBELI & DIECKMAN 2003). Speciation is the maintenance of phenotypic differences by means of reproductive isolation. Reproductive isolat- ing mechanisms are biological properties of individuals preventing the fusion of gene pools. A compatible defi- nition would be “biological properties, which lead to the interruption of the gene flow between populations”. In the example given above, a population differentiates along an ecological gradient. This gradual differentia- tion is maintained by the effects of natural selection. Reproductive isolation is not necessary to maintain the gradual difference. But as soon as, for example, female preferences for certain male characteristics become ge- netically coupled, the gradual differentiation of pheno- 126 Bonner zoologische Beitráge 53 (2004) types can be translated into an abrupt change in pheno- typic properties within now genetically separated popu- lations. This may eventually lead to reproductive isola- tion (DOEBELI & DIECKMANN 2003). In such a situation, individual selection favors the acquisition of reproduc- tive isolation. Starting from distinct phenotypes, the ac- quisition of genetic independence becomes a secondary phenomenon. The example above illustrates quite well that reproductive isolation is not necessarily a phe- nomenon of single gene loci but is best seen as a feature at the organismic level (compare MAYR 2001; WU 2001; WU & TING 2004). In some cases, only a small number of loci might suffice to lead to complete reproductive isolation, in some cases not. Albeit the notion of harmo- nious gene pools implies a genome wide differentiation, this 1s not even intended in MAYR’s writings (MAYR 1996). We could formulate that genes responsible for reproductive isolation are taking the lead in the differen- tiation process of the genotype and it will take some time until unique alleles will be fixed for every locus as soon as reproductive isolation has been acquired. Within sexually reproducing populations individuals will have genes (alleles) in common defining the biological prop- erties of reproductive isolation. We call these “species producing genes”. It might indeed be possible to give a comprehensive genetic delimitation of a species taxon if based on these species producing genes (MAYR 1996). The situation is somewhat different in allopatric popula- tions. Change of allele frequencies are expected to be randomly distributed within the genome assuming com- parable selection regimes in allopatric populations. From a theoretical point of view, this process, well known under the term random genetic drift inevitably reshapes the composition of genotypes and gene pools, finally generating incompatible gene pools between al- lopatric populations; incompatible in the sense that after secondary contact of populations, out-crossing individu- als will suffer reduced fitness. The acquisition of spe- cies status by random drift is assumed to be slow and probably of polygenic origin. Again, it might still be possible to give a comprehensive genetic species de- limitation based on genes causing isolating properties. The phenomenon of genetic incompatibility deserves closer attention since it has been considered as the most important factor of the speciation processes. Random genetic incompatibilities have been called Dobzhansky — Muller incompatibilities (ORR & TURELLI 2002). ORR (1995, 2001) showed that if the number of genetic dif- ferences between individuals of separated populations increases linearly with time, the number of incompati- bilities should increase as the square of their divergence times. This has been called the “snow ball” effect (ORR 1995). The snow ball effect predicts that we will proba- bly often overestimate the number of genetic incom- patibilities between populations responsible for postzy- gotic reproductive isolation. Therefore, we would ex- pect to see many cases of clear genetic incompatibility in experimental crosses of sister species. This is clearly not the case (compare experiments in CHARLESWORTH 1995). It appears that even an occurrence of multiple bottle necks is unable to produce enough genetic in- compatibilities between populations to establish effec- tive postzygotic isolation. This experimental evidence casts considerable doubt on the importance of random genetic drift in speciation and it also sheds some light on the interpretation of genetic variation recorded for selectively neutral marker loci. The experimental results tell us to expect substantial differentiation without postzygotic isolation. We have reviewed the cases in which “species defining genes” take the lead in the genetic differentiation proc- ess of newly formed populations. This does not have to be the case. Depending on speciation rates, selectively neutral loci might even take the lead in the differentia- tion process without causing reproductive isclation. This phenomenon is quite common and leads to geo- graphically structured gene pools with range limited mi- tochondrial haplotypes, for example, in the Eurasian ground dwelling fish Cottus gobio (ENGLBRECHT et al. 2000): several old lineages of Cottus gobio recolonized central Europe and readily mixed where they met. In Cottus gobio, genetic distinctness and even monophyly of haplotype groups restricted to geographical areas do not indicate reproductive isolation based on biological properties of the individuals. The above mentioned ex- perimental results and theoretical considerations should caution us in accepting selectively neutral marker loci as ideal tools of DNA taxonomy. However, all of the mo- lecular taxonomic work so far is based on selectively neutral marker loci (compare HEBERT et al. 2001). 4. CHARACTERS: CLASSES AND RELATIONS Populations of polytypic species and allopatric popula- tions in general, capable of fusing after secondary con- tact force one to accept that the biological properties of individuals, which prevent fusion of populations, are the hallmark of biological species. Therefore, it would be ill advised to base species delimitations on the recognition of any apomorphic character set shared by a number of individuals. This practice has been introduced by people defending derivatives of the phylogenetic species con- cept (in MAYDEN 1997; WHEELER & MEIER 2000). It is based on the argument that every reproductively iso- lated population will evolve autapomorphic character states. This is certainly true in the long run, but com- pletely ignores the initial conditions within reproduc- tively isolated populations. There is clearly a different quality to sets of characters whether they constitute “species producing” characters or not. The phylogenetic species concept does not take into account this differ- Bernhard Misor, Cornelya F. C. KLUTSCH, Oliver NIEHUIS & Alexandra PATT: Of Phenotypes and Genotypes 27 ence in character qualities. If different quality of infor- mation in characters is ignored, any observed difference gains weight. Representatives of typological species concepts refer to the apparent success in applying typo- logical criteria in delimitating species taxa (compare for example HEBERT et al. 2001). We have already shown that there are many conceivable situations in which DNA based taxonomies will lead to erroneous conclu- sions. Reproductive isolation is a quality of relations between populations and ultimately of individuals. This means that character attributes of reproductive isolation are from a different world compared to characters displayed by lets say pinned individuals in an insect collection. Imagine a pinned insect in front of you displaying a cer- tain suite of characters: relatively small and elongated, red with black dots on its back. In the drawer, in which the pinned insect was spotted, we discover several other individuals looking alike. In some other drawers, we manage to locate additional specimens with similar phenotypes. A decent hypothesis would be that all those specimens belong to one species characterized as rela- tively small, elongated, and red with black dots on its back. We could now perform a thought experiment and search for all other possible combinations of the charac- ters size, shape, and dorsal coloration. For example, we could look for big, stout, black and red spotted insects among the pinned myriads of specimens. This is cer- tainly not the only combination we can think of. Maybe we will even be successful and discover a suite of specimens that look exactly like the predicted organism. If we have been successful, we would have filled the class “big, stout, black and red spotted”. Does this imply that classes are real entities and not just projections of our mental condition? All organisms in a class truly dis- play the class defining characters and for this trivial rea- son there is a first reality to the phenomenon. Neverthe- less, there is more to classes of characters. Phenotypic characters are signs of the interaction of phenotypes and the environment, evolutionarily and ecologically. Con- sequently, a class of characters will share a set of envi- ronmental interactions that is definitely real. The causal relationship of form and function in this case enables the prediction of form from function and vice versa. It is clear that the causal relationship of form and function is not a linear mapping in biological systems, but we think that in principle, this statement will hold. In contrast, reproductive isolation as a quality of individual (popula- tion) interaction emerges as a character of relation sepa- rating two sets of individuals. It is in this sense an emergent quality of organisms with historical-factual properties tied to its bearers. Biological properties of in- dividuals preventing fusion of populations are classes of characters, which can be expected to be mutually exclu- sive in reproductively isolated populations. Traditional taxonomy. tries to identify those classes of characters responsible for the quality of reproductive isolation. By doing this, taxonomists often just intul- tively sort and weight characters drawn from the general pattern (classes) displayed by individuals to approxi- mate the phenomenon of reproductive isolation. Our sensory apparatus and experience as taxonomists makes us successful in identifying and circumscribing biologi- cal species based on phenotypic characters. This is common sense among taxonomists and accepted outside the taxonomic community as well. It would be fascinat- ing to collect complete genetic information on charac- ters leading to reproductive isolation in circumscribing biological species taxa (see also MAYR 1996), but un- fortunately, this would probably be a hopeless activity. The quality of reproductive isolation 1s not restricted to a general class of phenotypic characters and we can ex- pect all kinds of genotypic combinations in this respect. It is obvious that an automated identification and cir- cumscription of species taxa based on the genetics of reproductive isolation is not feasible in the near future. In contrast, DNA taxonomy tries to infer species taxa from analyzing more or less randomly drawn genetic in- formation. With randomly drawn genetic information we refer to the selection of genetic marker systems, which has until now never been guided by the needs of taxonomy. The inclusion of genotypic characters was traditionally based on economical and technical consid- erations. The limits of the genetic markers currently in use are easily highlighted. A monophyly of sequenced haplotypes will coincide with species taxa only if there was enough time for the population to complete fixation before the onset of a new speciation event. It is there- fore an empirical question whether the application of DNA taxonomy becomes restricted to just “old” or equivalently fast divergence processes. FUNK € OM- LAND (2003) estimated the frequency of species-level paraphyly or polyphyly at 23% based on 2319 published reports. This frequent occurrence of paraphyly and polyphyly should warn the DNA-taxonomist. 5. SPECIES TAXON AND SPECIMEN IDENTIFICATION 5.1. Introduction Species taxa are most properly identified by their spe- cies-producing class of characters, called biological properties of the individuals preventing the fusion of populations. This is of course often impossible to achieve. The art in taxonomy is the selection of charac- ter sets which come close to that goal. At the phenotypic level, we operate by comparing character sets. weight- ing the information at hand, and develop a hypothesis about the relevance of different character sets. In an ideal case, we would filter a certain set of characters re- 128 Bonner zoologische Beitráge 53 (2004) sponsible for reproductive isolation. The hypothesis that the selected set of characters is responsible for repro- ductive isolation between a pair of species is at least theoretically open for empirical falsification. This is a most important strength of the biological species con- cept. It separates the biological species concept from all other concepts, which emphasize the population lineage properties of species. The introduction of a historical component in species definitions, like specimens belong to a certain species because they are directly linked with other species by descent, creates an hypothesis which is not directly open for empirical falsification. It is clear that organisms are linked by descent. There is nothing wrong with this. But we are convinced that it is not nec- essary to jumble explanations of historical-narrative with hypothetical-deductive qualities in delimiting spe- cies taxa. It is like mixing of taxonomy and phyloge- netic systematics without noticing the essential different qualities of the disciplines. Admittedly, the empirical test is seldom conducted and therefore, most species de- scriptions have to be regarded as points of views instead of critically evaluated hypothesis. But how can the taxonomic enterprise still be a success? It all rests on the shoulders of the specialists familiar with their group of organisms. These specialists develop a feeling for the relevant characters delimiting species taxa in their groups. The success of the traditional taxonomic method is therefore dependent on life-long experience and intui- tive solutions. These are certainly not ideal precondi- tions for the automated inventarisation of biodiversity. 5.2. Specimen identification by means of molecular tools In general, DNA taxonomy does not deal with species- producing sets of genes. It could be hoped for that “spe- cies-producing genes” are not randomly distributed among at least closely related taxa enabling a semi- automated detection of those genes. But detailed genetic analyses of speciation processes revealed that even among closely related sister taxa of Drosophila, species- specific genes greatly vary and can not be compared (see review in WU & TING 2004). The manifold differ- ences in courtship display alone seem to support this view. The contradicting argument might be raised that several studies document a driving force of sexual selec- tion in speciation processes (for example OWENS et al. 1999; PANHUIS et al. 2001; ARNQVIST et al. 2000; MISOF 2002). If sexual selection is fueling speciation, it will mostly be restricted to pre-mating isolation and varia- tion within courtship displays. The extent of phenotypic species-differences should be directly translated into a set of genotypic differences comparable between spe- cies taxa. To us it appears that we just do not know enough about speciation genes and comparative analy- ses of speciation processes. We will have to wait for an- swers from combined empirical and comparative analy- ses of speciation phenomena until we can develop subtle hypotheses of the “genic view of speciation” (WU 2001). In any reproductively isolated group, genetic differences will accumulate. Eventually, genetic differences will become fixed for selectively neutral loci as well. Subse- quently, the genetic variation for selected loci will gain characteristic autapomorphic characters. But those populations will not by necessity evolve isolating mechanisms. Again, in delimitating and circumscribing species taxa we need to identify the biological proper- ties of individuals causing reproductive isolation. It is not at all sufficient to show even in a sympatric situation that two subgroups of individuals maintain phenotypic differences independently of sex. Just think of geneti- cally fixed polymorphisms within species, text book ex- amples are some orchid flowers or Anopheles mos- quitos. Selectively neutral marker loci will not be among the candidates. We refer the reader to the illumi- nating experimental data on effects of random drift on postzygotic reproductive isolation (CHARLESWORTH 1995). Admittedly, autapomorphic characters of, for ex- ample, mitochondrial haplotypes within a clear biologi- cal species can be used a posteriori to filter specimen identity. This filtering of specimens is equivalent to species assignment. Assignment is restricted in its ap- plicability to situations, in which haplotypes within sis- ter taxa have reached monophyly and diagnostic (auta- pomorphic) character states (for an empirical analysis compare FUNK & OMLAND 2003). It is important to stress the reciprocal monophyly of sister taxa. It is not sufficient to show that the biological species under con- sideration shows a monophyletic set of haplotypes. For example, consider the scenario of a peripatric speciation event. We start from a population with a large distribu- tional range with a certain degree of genetic variability. In a peripatric situation, geographically marginal sub- populations might evolve reproductive isolation. Their haplotypes of selectively neutral marker loci will likely be monophyletic. However, the stem population from which the peripatric population became isolated was driven in the status of paraphyly. Nevertheless, indi- viduals from the stem population will remain reproduc- tively compatible. It will take time until the stem popu- lation again reaches monophyly for the haplotypes under consideration. The paraphyletic situation in the stem population poses clear problems to specimen iden- tification and assignment. Some haplotypes within the stem population will be more closely related to haplo- types of the reproductively isolated population. The dis- tribution of genetic distances will not correspond with species taxa and consequently, can not be used to infer species taxa. It 1s particularly surprising that genetic distance measures are advocated in many recent publi- cations for the delimitation of species taxa in phenotypi- cally indistinguishable forms. The character based as- Bernhard Misor, Cornelya F. C. KLUTSCH, Oliver NIEHUIS & Alexandra PATT: Of Phenotypes and Genotypes 129 signment and identification is the only way to go. If, and only if, a set of diagnostic characters has been correlated with reproductively isolated populations, these diagnos- tic characters can be used in specimen assignment and identification. A theoretical example will hopefully illustrate the ar- gument above. Assume we are naive DNA taxonomists and received an unknown invertebrate larva collected in marine sediments. The assignment of the larva to a known taxon will be of great importance because it can bridge gaps in the analysis of biodiversity. We might use genetic information to assign the larva to a certain taxon for which genetic markers have already been characterized. From this larva, we sequence fragments of three different gene loci and the observed sequence information indicates that the sequences are only 5% different from sequences of specimens within a known taxon. As naive taxonomists, we would very quickly say that the larva is the yet undescribed larva of a new taxon. Another detailed analysis of phenotypes and bio- logical properties of individuals within the species taxon suggests that the species has a very heterogeneous ge- netic distance distribution within its range. We find that genetic distances are indeed observed between 0 - 5 % within a Mendelian group, but a geographically mar- ginal subpopulation appears reproductively isolated judged from additional evidences. Although, genetic distances within this subpopulation are small and the average genetic distance of this subpopulation to mem- bers of the stem population fall within the recorded range of the presumably Mendelian stem population. In this situation, genetic distances will be of no help in specimen identification and assignment. However, if we concentrate our search on the identification of autapo- morphic characters, which describe the recently derived reproductively isolated subpopulation, we could cir- cumvent the problem of overlapping genetic distances and paraphyly of the stem population. The likelihood of specimen assignment will depend on the extent of pre- vious sampling within the stem and subpopulations; the better the sampling, the better our chance of correct re- identification (compare POZHITKOV & TAUTZ 2002). POZHITKOV & TAUTZ (2002) developed an algorithm to select ideally suited probes for specimen identification in large through-put-assays. This strategy seems very promising, in particular for small organisms in great abundance. The assignment of specimens will of course obey the rules of probability. A very nice example has been developed by the www.DNA-surveillance project. It constitutes a web-based molecular identification of whales, dolphins and porpoises (BAKER et al. 2003). Obviously, this DNA based specimen identification tool has helped to review species descriptions and type as- signments in this group of organisms. The taxonomy of cetaceans is plagued by the problem of some extremely rare species for which morphological descriptions are often very incomplete. In this case, DNA based taxon- omy has undoubtedly helped clarifying issues. 5.3. Specimen identification by means of phenotypic tools F There are other approaches for the rapid (re-)iden- tification of species beside DNA based taxonomy. One possibility is automated species identification through digital image analysis (WEEKS et al. 1997; WEEKS et al. 1999a, 1999b; WATSON et al. 2004; ROTH et al. 1999; STEINHAGE et al. 1997; STEINHAGE 2000). In this auto- mated approach, images of insect wings (e. g. of moths, bees, and wasps) are used to distinguish between spe- cies. Images are analyzed concerning venation, color or scale pattern. The system DAISY (Digital Automated Identification System; O'NEILL & GAULD 2001; WAT- SON et al. 2004), for example, was able to identify 83 % of unknown moth species from wing images. Taken into account that digital images were taken in the field, where light conditions are not always standardized and recorded specimens are moving, this is an impressively high success rate. A similar approach is run by the ABIS (Automated Bee Identification System, ROTH et al. 1999; STEINHAGE et al. 1997; STEINHAGE 2000) pro- ject. Here, images of forewings of bees, are taken and their venation is used to identify species on different hi- erarchical levels like genus, species group within a ge- nus and species. Both projects have in common that the programs have to be trained with sets of images to have some kind of “experience” in identifying species. Thus, the systems are only able to discriminate between al- ready digital recorded species. The systems are able to re-identify specimens of known taxa, but not able to recognize new species. In consequence, this can lead to misidentifications, as the system always tries to match an unknown image to a species. GASTON & O ‘NEILL (2004) pointed out that an ideal automated species iden- tification system should be able to reject species identi- fication if similarities to known species in the database are too small. Hence, this could help to identify un- known phenotypes within a set of known species in the future. Please notice, this is not similar to saying that the automated systems can be used for recognizing new species. We suggest that the systems can be used to rec- ognize unknown phenotypic variation, which can then subsequently be evaluated by taxonomists. Further im- provements of the systems could include better images, more morphological characters (e. g. images of addi- tional parts of the body like limb or head structures), and a database with information about distribution, food plants, flight time and other biological data (WATSON et al. 2004). The project DORSA (German Orthoptera Collections, RIEDE et al. 2004) is one attempt to incor- porate distribution maps, links to collections, images of type specimens, and last but not least sound recordings 130 Bonner zoologische Beitráge 53 (2004) in a single database. A rapid assessment tool was devel- oped for automated song recognition and a sound library will be established soon. In this case, it is possible to re- identify orthopteran species due to the characteristic song pattern of single species. The integration of the mentioned distributional and biological data allows a verification of the identified species. But again, this ef- fort is primarily suitable for re-identifications and not for recognition of new species since the only way to confirm a specimen as an unknown species is to exclude all other known species, which can be difficult if not all known species are included in the database. In summary, although auspicious efforts have been made to automate species identification through digital image analysis, it is still a long way to go. Two major aspects have to be emphasized in this context. First, all of the presented computed systems are some kind of semi-automated since the program has to be trained; da- tabase establishment and updates as well as verification of recorded species will always remain in human hands. Second, these projects are tools for re-identification and not for recognition of new species. But computer-based routine re- identification systems could be at least a great facility to speed up the re-identification of species in biodiversity research. Of course, 1f species can only be differentiated by genital morphology, which needs first a preparation step, the automated re-identification system will be again of limited use. 6. CONCLUSIONS The proponents of DNA taxonomy try to convince that without relying on molecular characters we will not be able to register a significant share of biodiversity in rea- sonable time. Automated DNA based taxonomy is the key word here. We have tried to argue that taxonomy can not become a robot activity and speed up taxonomy for several reasons. The most important one is that de- scribing species taxa is a scientific activity resulting in testable hypotheses despite the prevailing absence of empirical falsifications. The hypotheses are based on universal theories (species are reproductively isolated groups of individuals) and identified local realizations (species-specific characters causing reproductive isola- tion). Biological properties of individuals preventing fu- sion of populations are as numerous as there are bio- logical species and are emergent qualities of species taxa. These qualities can not be predicted and conse- quently elude an automatic detection. But still, we ex- pect a significant impetus from molecular studies, which analyze genetic variability within and among popula- tions. New data on genetic variability can potentially di- rect our interests in taxonomic studies if genetic and phenotypic data do not correlate. For example, investi- gations are published, which report on the application of molecular techniques revealing hitherto masked differ- entiation among seemingly identical phenotypes (PROUDLOVE & WOOD 2003). Obviously, DNA based methods help to discover patterns of differentiation that are possibly correlated with reproductive isolation. For some groups of organisms, in which morphology will not be of great help, the application of molecular tools can certainly be advantageous and become mandatory. However, similarly to phenotype-based species descrip- tions, assessments of species status based on molecular characters remain points of view as well and are even more limited in their relevance compared to phenotypic characters. Molecular differentiation based on selec- tively neutral marker loci is definitively not a test of re- productive isolation between populations. Given a set of “well” circumscribed species taxa, DNA taxonomy will speed up descriptive work in biodiversity assessment in the sense that the number of genetically diagnosable species for a given collection event or area can be auto- matically recorded. This work will profit from automa- tion since hypotheses testing is not involved. However, recent developments in automatic specimen identifica- tion based on phenotypes are encouraging as well. These developments should not become underfunded in competition with molecular tools, because the ultimate application of phenotypic identification seems more versatile than any DNA based method. We emphasize that the current practice of using genetic distances to de- limit species is ill advised and should be replaced by character-based methods. We see that DNA taxonomy has its clear limitations in specimen (re-) identifications and species taxa circumscriptions. We should stress that we do not even have a grip on the problems facing the description of the 90% undescribed taxa and we specu- late that many of them are not described yet because they are close sister taxa of known species. If this is true we are gearing towards the area in which DNA taxon- omy is most limited. We should be aware of this prob- lem. ADDENDUM It is clear to us that we largely ignored the distinction between sympatric/parapatric and allopatric situations. Some readers might argue that our argumentions are more or less acceptable in allopatric situations, in which assignment of species status is arbitrary anyway, but are not applicable under sympatric/parapatric conditions. It is indeed tempting to say that in a sympatric situation at least two unlinked selectively neutral markers, which segregate independently of sex, indicate separate spe- cies status of their bearers. We would like to argue that even in a sympatric case just two selectively neutral markers rarely tell us who is who. Theoretical work shows that in a sympatric speciation process the differentiation of selectively neutral markers Bernhard MIsor, Cornelya F. C. KLUTSCH, Oliver NIEHUIS & Alexandra PATT: Of Phenotypes and Genotypes 13] will lag behind. But finally, even selectively neutral markers will reach fixation. In this case they can indeed indicate species status. However, we would assume that phenotypic differentiation will be quite apparent as well and DNA taxonomy will tell us what we already guessed. On the contrary, consider a secondary contact of populations. If populations display differentiation, for example ecological or sexual differentiation, reinforce- ment will further promote the evolution of reproductive isolation. But since reproductive isolation is not abso- lutely perfect, selectively neutral marker loci can still penetrate species borders. In consequence, well differ- entiated selectively neutral markers will become associ- ated with the wrong “species” without dissolving spe- cies borders. We would expect polyphyly or paraphyly of selectively neutral marker loci in relation to species borders and phenotypic differentiation (equivalent to species differentiation genes) will be the better choice in taxonomy. This is exactly what FUNK & OMLAND (2003) found in roughly 23% of published reports. Without phenotypic differentiation in sympatry, DNA taxonomy based on selectively neutral markers can only help if several selectively neutral marker loci display congruent pictures of reciprocal monophyly. We are not aware of a published report applying these considera- tions rigorously. Acknowledgements. We would like to thank all members of the molecular unit of the ZFMK for extremely fruitful discussions. We dedicate this paper to the memory of Clas Michael Naumann who died on the 15'" February 2004. His much to early death has taken away a person who has come to the aid of us whenever it was necessary; we have to grow up now without him. REFERENCES ARNQVIST, G., EDVARDSSON, M., FRIBERG, U. & NILSSON, T. (2000): Sexual conflict promotes speciation in in- sects. Proceedings of the national Academy of Sci- ences of the USA 97(19): 10460-10464. AVISE, J. C. (1987): Intraspecific phylogeography: the mi- tochondrial DNA bridge between population genetics and systematics. Annual Review of Ecology and Sys- tematics 18: 489-522. AVISE, J. C. (2000): Phylogeography — The History and Formation of Species. Harvard University Press, Cam- bridge, Massachusetts BAKER, C. S., DALEBOUT, M. L., LAVERY, S. & Ross, H. A. (2003): www.DNA-surveillance: applied molecular taxonomy for species conservation and discovery. Trends in Ecology and Evolution 18(6): 271-272. BONNER, J. T. (1988): The Evolution of Complexity. Princeton University Press, Princeton, NJ. CHARLESWORTH, B. (1995): Down the bottleneck? Current Biology 5: 995-996. DE QUEIROZ, K. (1999): The general lineage concept of species and the defining properties of the species cate- gory. Pp. 49-89 in WILSON, R. A. (ed.) Species. MIT Press, Cambridge, Massachusetts. DE SALLE, R. & TEMPLETON, A. R. (1988): Founder effects accelerate the rate of mtDNA evolution of Hawaiian Drosophila, Evolution 42: 1076-1084. DOBZHANSKY, T. (1935): A critique of the species concept in biology. Philosophy of Science 2: 344-355. DOBZHANSKY, T. (1937): Genetics and the Origin of Spe- cies. Columbia University Press, New York / DIECKMANN, U. & DOEBELI, M. (1999): On the origin of species by sympatric speciation. Nature 400(6742): 354-357. DOEBELI, M. & DIECKMANN, U. (2003): Speciation along environmental gradients. Nature 421(6920): 259-264. ENGLBRECHT, C. C., FREYHOF, J., NOLTE, A., RASSMANN, K., SCHLIEWEN, U. & TAUTZ, D. (2000): Phylogeo- graphy of the bullhead Costus gobio (Pisces: Teleostel: Cottidae) suggests a pre-pleistocene origin of the ma- jor central European populations. Molecular Ecology 9(6): 709-722. EYUALEM, A. & BLAXTER, M. (2003): Comparison of bio- logıcal, molecular and morphological methods of spe- cies identification in a set of cultured Panagrolaimus isolates. Journal of Nematology 35: 119-128. FISHER, R. A. (1930): The Genetical Theory of Natural Se- lection. Clarendon Press, London FUNK, D. J. & OMLAND, K. E. (2003): Species-Level Para- phyly and Polyphyly: Frequency, Causes, and Conse- quences, with Insights from Animal Mitochondrial DNA. Annual Review of Ecology, Evolution, and Sys- tematics 34: 397-423. GASTON, K. J. & O'NEILL, M. A. (2004): Automated spe- cies identification-why not? Philosophical Transac- tions of the Royal Society London, Biology 359 (1444): 655-667. GODFRAY, H. C. J. (2002): Challenges for taxonomy. Na- ture 417: 17-19, HARTL. D. L. & CLARK, A. G. (1997): Principles of popu- lation genetics, 3rd ed. Sinauer Associates, Sunder- land, MA. HAASE, M., MISOF, B., WIRTH, T., BAMINGER, H., & BAUR, B. (2003): Genetic Evidence for Survival of a Land Snail under Pleistocene Permafrost. Journal of Evolutionary Biology 16: 415-428. HEBERT, P. D. N., CYWINSKA, A., BALL, S. L. & DE- WAARD, J. R. (2001): Biological identifications through DNA barcodes. Proceedings of the Royal So- ciety London B 270, 313-322. HEBERT, P. D. N., PENTON, E. H., BURNS, J., JANZEN, D. J. & HALLWACHS, W. (2004): Ten species in one: DNA barcoding reveals cryptic species in the neotropical skipper butterfly, Astraptes fulgerator. Proceedings of the National Academy of Sciences of the USA 101 (41): 14812-14817. JIGGINS, C. D. & DAVIES, N. (1998): Genetic evidence for a sibling species of Heliconius charithonia (Lepidop- tera; Nymphalidae). Biological Journal of the Linnean Society 64: 57-67. LIPSCOMB, D. L., PLATNICK, N. & WHEELER, Q. (2003): The intellectual content of taxonomy: a comment on DNA taxonomy. Trends in Ecology and Evolution 18: 65-66. MALLET, J. & WILLMOTT, K. (2003): Taxonomy: renais- sance or Tower of Babel. Trends in Ecology and Evo- lution 18: 57-59. MALTAGLIATI, F., CAMILLI, L., LARDICCI, C. & CASTELLI, A. (2001): Evidence for morphological and genetic di- 132 Bonner zoologische Beitráge 53 (2004) vergence in Perinereis cultrifera (Polychaeta, Nereidi- dae) from two habitat types at Elba Island. Journal of the Marine Biological Association of the United King- dom 81: 411-414. MAYR, E. (1970), Populations, Species, and Evolution. Harvard University Press, Cambridge, MA. MAYR, E. (1982): The growth of biological thought. Diver- sity, Evolution and Inheritance. The Belknap Press of Harvard University Press, Cambridge, MA etc. MAYR, E. (1996): What is a species and what is not? Phi- losophy of Science 63: 262-277. MAYR, E. (2001): Wu’s genic view of speciation. Journal of Evolutionary Biology 14: 866-867. MAYDEN, R. L. (1997): A hierarchy of species concepts: the denouement in the saga of the species problem. Pp. 381-424 in CLARIDGE, M. F., DAWAH, H. A. & WIL- SON, M. R. (eds.) Species - the Units of Biodiversity. Chapman & Hall, London MEYER, A., KOCHER, T. D., BASASIBWAKI, P. & WILSON, A. C. (1990): Monophyletic origin of Lake Victoria cichlid fishes suggested by mitochondrial DNA se- quences. Nature 347: 550-553. MISOF, B. (2002): Diversity of Anisoptera (Odonata): In- ferring speciation processes from patterns of morpho- logical diversity. Zoology 105: 355-365. NEI, M. (1978): Estimation of average heterozygosity and genetic distance from a small number of individuals. Genetics 89: 583-590. O'NEILL, M. A. & GAULD, I. D. (2001): DAISY. Auto- mated insect identification project. http://www.chas- seuer.usc.edu./pups/projects/daisy.html OHTA, T. (1976): Role of slightly deleterious molecular evolution and polymorphism. Theoretical Population Biology 10: 254-275. ORR, H. A.(1995): The population genetics of speciation: the evolution of hybrid incompatibilities. Genetics 139: 1805-1813. ORR, H. A. (2001): The genetics of species differences. Trends in Ecology and Evolution 16(7): 343-350. Orr, HA. & TURELLI, M. (2002): The evolution of postzygotic isolation: accumulating Dobzhansky- Muller incompatibilities. Evolution 55: 1085-1094. OWENS, I. P. F., BENNETT, P. M. & HARVEY, P. H. (1999): Species richness among birds: body size, life history, sexual selection or ecology? Proceedings of the Royal Society London B 266: 933-939. PANHUIS, T., BUTLIN, R., ZUK, M. & TREGENZA, T. (2001): Sexual selection and speciation. Trends in Ecology and Evolution 16(7): 364-371. POZHITKOV, A. & TAUTZ, D. (2002): An algorithm and program for finding sequence specific oligo-nucleotide probes for species identification. BMC Bioinformatics 3:9. PROUDLOVE, G. & Woop, P. J. (2003): The blind leading the blind: cryptic subterranean species and DNA tax- onomy. Trends in Ecology and Evolution 18(6): 272- 273. RAFF, R. A. (1996): The Shape of Life: Genes, Develop- ment, and the Evolution of Animal Form. University of Chicago Press, Chicago. RIEDE, K., LAMPE, K.-H. & INGRISCH, S. (2004): Deutsche Orthopterensammlungen (DORSA) Aufbau eines vir- tuellen Museums - Verhandlungen Westdeutscher En- tomologentag 2002, Lobbecke Museum, Diisseldorf, 63 (DORSA: German Orthoptera Collections; http://www.dorsa.de/) RIEDL, R. (1978): Order in Living Organisms. Wiley Press, New York. ROTH, V., POGODA, A., STEINHAGE, V. & SCHRODER, St. (1999): Pattern Recognition Combining Feature- and Pixel-based Classification Within a Real World Appli- cation. 21. Jahrestagung der Deutschen Gesellschaft fiir Mustererkennung, DAGM '99, Bonn, Sept. 15-17, 1999, Informatik aktuell: 120-129. RUNDLE, H. D. (1999): Experimental tests of founder- flush: a reply to Templeton. Evolution 53: 1632-1633. SEBERG, O., HUMPHRIES, C. J., KNAPP, S., STEVENSON, D., PETERSEN, G., SCHARFF, N., ANDERSEN, N. M. (2003): Shortcuts in Systematics? - A commentary on DNA- based Taxonomy. Trends in Ecology and Evolution 18: 63-65. STEINHAGE, V. (2000): Automated Identification of Bee Species in Biodiversity Information Systems. Com- puter Science for Environmental Protection 2000, UI 2000, Bonn, Oct. 4-6, 2000, Metropolis Publ. 1: 339- 344. STEINHAGE, V., KASTENHOLZ, B., SCHRODER, S. 8 DRE- SCHER, W. (1997): A Hierarchical Approach to Clas- sify Solitary Bees Based on Image Analysis. Muster- erkennung 1997, 19. DAGM-Symposium, Informatik aktuell: 419-426. STOECKLE, M. (2003): Taxonomy, DNA, and the Bar Code of Life. Bioscience 53 (9): 2-3. TAKAHASHI, K., TERAI, Y., NISHIDA, M. & OKADA, N. (2001): Phylogenetic relationships and ancient incom- plete lineage sorting among cichlid fishes in Lake Tanganyika as revealed by analysis of the insertion of retroposons. Molecular Biology and Evolution 18(11): 2057-2066. TAUTZ, D., ARCTANDER, P., MINELLI, A., THOMAS, R. H. & VOGLER, A. P. (2002): DNA points the way ahead in taxonomy. Nature 418: 479. TAUTZ, D., ARCTANDER, P., MINELLI, A., THOMAS, R. H. & VOGLER, A. P. (2003): A plea for DNA taxonomy. Trends in Ecology and Evolution 18: 70-74. TURELLI, M., BARTON, N. H. & COYNE, J. A. (2001): The- ory and speciation. Trends in Ecology and Evolution 16(7): 330-343. VON BERTALANFFY, L. (1932): Theoretische Biologie. 2 vols. Gebriider Borntrager, Berlin. WADDINGTON, C. H. (1957): The strategy of genes. Allen & Unwin, London. WAGNER, G. P. (1996): Homologues, natural kinds and the evolution of modularity. American Zoologist 26: 36— 43. WAGNER, G. P. € ALTENBERG, L. (1996): Complex adap- tations and the evolution of evolvability. Evolution 50: 967-976. WATSON, A. T, O'NEILL, M. A. € KITCHING, I. J. (2004): Automated insect identification of live moths (Macro- lepidoptera) using Digital Automated Identification System (DAISY): Systematics and Biodiversity 1(3): 287-300. WAYNE, R. K., MEYER, A., LEHMAN, N., VAN VALKEN- BURGH, B., KAT, P. W., FULLER, T. K., GIRMAN, D. & O’BRIEN, S. J. (1990): Large sequence divergence among mitochondrial DNA genotypes within popula- tions of eastern African black-backed jackals. Proceed- Bernhard Misor, Cornelya F. C. KLUTSCH, Oliver NIEHUIS & Alexandra PATT: Of Phenotypes and Genotypes 133 ings of the National Academy of Sciences of the USA 87: 1772-1776. WEEKS, P. J. D., GAULD, I. D., GASTON, K. J. & O'NEILL, M. A. (1997):Automating the identification of insects: a new solution to an old problem. Bulletin of Entomo- logical Research 87: 203-211. WEEKS, P. J. D., O'NEILL, M.A., GASTON, K. J. & GAULD, I. D. (1999a): Automating insect identification: explor- ing the limitations of a prototype system. Journal of Applied Entomology 123: 1-8. WEEKS, P. J. D., O'NEILL, M.A., GASTON, K. J. & GAULD, I. D. (1999b): Species-identification of wasps using principle component associative memories. Image and Vision Computing 17: 861-866. WHEELER, Q. D. & MEIER, R. (eds) (2000): Species Con- cepts and Phylogenetic Theory: a Debate. Columbia University Press, New York WRIGHT, S. (1931): Evolution in Mendelian populations Genetics 16: 97-159. WRIGHT, S. (1951): The genetical structure of populations Annals of Eugenics 15: 323-354. Wu, C.-I. (2001): The genic view of the process of speci- ation. Journal of Evolutionary Biology 14: 851-865. Wu, C.-L. & TING C.-T. (2004): Genes and speciation. Na- ture Reviews, Genetics 5: 114-122. Authors’ address: Bernhard MISOF (corresponding au- thor), Cornelya F. C. KLUTSCH, Oliver NIEHUIS & Alex- andra PATT, Zoologisches Forschungsmuseum Alexan- der Koenig, Adenauerallee 160, 53113 Bonn, Germany, e-mail: b.misof.zfmk(@uni-bonn.de 2 7 ] j e & - . 7 a ' Ei Bonner zoologische Beitrage Band 53 (2004) | Heft 1/2 Seiten 135-148 3onn, Juni 2005 A New Lizard of the Genus Gekko Laurenti, 1768 (Squamata: Sauria: Gekkonidae) from the Phong Nha - Ke Bang National Park, Quang Binh + Province, Vietnam! Herbert RÓSLER”, Thomas ZIEGLER”, VU Ngoc Thanh”, Hans-Werner HERRMANN’ ) & Wolfgang BOHME”’ 2 . . . S . 4 N > Staatliche Naturhistorische Sammlungen Dresden, Museum für Tierkunde, Forschungsstelle, Dresden, Germany AG Zoologischer Garten Köln, Germany CRES, Centre for Natural Resources and Environmental Studies, Vietnam National University, Hanoi, University of Science, Faculty of Biology, Department of Vertebrate Zoology, Zoological Museum, Thanh Xuan, Hanoi, Viet- nam Conservation and Research for Endangered Species (CRES), Zoological Society of San Diego, USA % Zoologisches Forschungsmuseum Alexander Koenig, Bonn, Germany Abstract. We describe a new species of the genus Gekko from the karst forest of the Phong Nha - Ke Bang National Park, Quang Binh Province, Vietnam. Gekko scientiadventura sp. n. is distinguished from all other Gekko species occur- ing in Vietnam by the lack of dorsal tubercles. From the four species of Gekko that do not occur in Vietnam and that have likewise no dorsal tubercles (athymus, melli, subpalmatus, and tawaensis), the new species may be distinguished as follows: G. athymus has more preanal pores than G. scientiadventura sp. n. whereas in G. tawaensis both preanal and femoral pores are completely lacking. G. melli and G. subpalmatus are apparently closer related to G. scientiadventura sp. n., but in these two species the nasorostrals are separated from each other by internasals. We provide first data on habitat and natural history of the new species and furthermore provide a key for the Vietnamese Gekko species. Key words. Gekko scientiadventura sp. n., karst forest; taxonomy, natural history. 1. INTRODUCTION According to BOBROV (1995), six species of the genus Gekko exist in Vietnam: G. badenii Szczerbak & Nek- rasova, 1993, G. chinensis (Gray, 1842), G. gecko (Lin- naeus, 1758), G. japonicus (Schlegel, 1836), G. pal- matus Boulenger, 1907 and G. ulikovskii Darevsky & Orlov, 1994. However, according to the investigations of OTA et al. (1995) the occurrence of G. japonicus in northern Vietnam (see also NGUYEN & HO 1996) must be regarded as doubtful. Not yet contained in BOBROV's (1995) list is G. grossmanni Gúnther, 1994. SCHLUTER (1999) claims G. petricolus Taylor 1962 to occur in the “triangle of Thailand, Vietnam, and Cambodia”, but definite Vietnamese records of this species are - to our best knowledge - still lacking. The incomplete state of knowledge of Vietnam's gekkonid fauna is also high- lighted by the recent description (1993-1994) of the three species of the genus Gekko mentioned above (for the correct year of description of G. badenii see GUN- THER 1994). When we started our herpetological investigations of the Phong Nha - Ke Bang karst forest, in the Quang Binh I In commemoration of Clas Michael Naumann zu Königsbrück (26.06.1939 — 15.02.2004) Province close to the Laotian border seven years ago, aside from Gekko gecko only three species of Hemidac- tvlus were known (ZIEGLER & HERRMANN 2000). Dur- ing our subsequent studies of the herpetological diver- sity of this area we were able to record three additonal species in the National Park: Gehyra mutilata (Wieg- mann, 1834) and an undescribed species of each of the genera Cyrtodactylus and Gekko (ZIEGLER et al. 2004). The former has been described already by ZIEGLER et al. (2002), but the latter, listed still as Gekko sp. by ZIEGLER et al. (2004), will be diagnosed and described below. Because in the aforementioned papers the Phong Nha - Ke Bang National Park, its herpetofauna and the respec- tive conservation needs have already exhaustively been described, we restrict ourselves in the present paper on the diagnosibility of the new species of Gekko. 2. MATERIAL AND METHODS Voucher specimens are deposited in the Vietnam Na- tional University (HNUV), Hanoi, in the Zoologisches Forschungsmuseum Alexander Koenig (ZFMK), Bonn, and in the personal collection of the senior author (CPHR). Comparative material was provided (cadens according to LEVITON et al. 1985) by the Natural His- 136 Bonner zoologische Beitráge 53 (2004) tory Museum (BMNH), London, and the Museum fiir Naturkunde (ZMB), Berlin. Biometric and scalation characters as well as colour pat- tern data were evaluated in both living and preserved specimens. Measurements were taken with a dial calip- per (to the nearest mm). The following abbreviations are used: a.s.l. = above sea level, SVL = snout vent length, TL = tail length, SE = distance from snout tip to front of eye, EE = distance between hindmargin of eye to hind- margin of ear, HW = maximum head width, HH = maximum head height, HL = head length (from tip of snout to hindmargin of ear). The following scale counts were taken: supralabials (SPL), sublabials (SBL), nasals (N); in direction from rostral to labial: nasorostrals, su- pranasals, postnasals, internasals (IN); postmentals (PM), gulars bordering the postmentals (GP), interorbi- tals (IO), ventrals (V), scales around midbody (SB), subdigital lamellae under the first (LT1) and fourth toe (LT4), preanal pores (PP), postanal tubercles (PAT), dorsal scale rows in the middle of the third caudal whorl (S3R). In living specimens symmetrical scale counts were taken only on the lizard's right side. Colouration is described according to the standard plates published by GRALLERT & ROLAND (1960). 3. RESULTS 3.1. Description of Gekko scientiadventura sp. n. 3.1.1. Diagnosis. A small-bodied species of Gekko. The biggest specimen (captive) has a total length of 154 mm, of which 81 mm are made up of the tail. Habitus slen- der, head distinctly broader than neck, head and body slightly depressed. Unregenerated tail always longer than head-body length, not constricted at its base, not thickened, slighty depressed. Lateral fold weak, upper side of head, body and tail without tubercular scales. Digits and toes slightly webbed at the base, only inter- space between toes 4 and 5 not webbed. All digits and toes, except the inner ones, clawed. 14-17 subdigital lamellae below fourth toe. 5-8 preanal pores. Nostril touches rostral. No internasals. Posterior ciliaries spiny. Upper side yellowish to brownish in life. Dorsally seven large light spots that may be expanded to lateral narrow wavy bands. Tail with 7-10 light crossbands. Gular re- gion and parts of venter marbled. From all species of Gekko known to occur in Vietnam (G. badenii, G. chinensis, G. gecko, G. japonicus, G. palmatus, G. ulikovskii), G. scientiadventura sp. n. iS invariably distinguished by the lack of middorsal tuber- cles. For more detailed comparison, also with other Gekko-species see below (3.4. and 3.5.). 3.1.2. Holotype. ZFMK 76198, adult male, Figs. 1-3, primary limestone forest, Phong Nha - Ke Bang, sur- roundings of 17.32 N, 106.16 E, ca. 50-150 m a.s.l., Quang Binh. Province, Vietnam, coll. H.-W. HERRMANN, VU NGOC THAN & T. ZIEGLER, 27-29 Au- gust 2001. Fig. 1: Live holotype of Gekko scientiadventura sp. n. (ZFMK 76198). Phot. T. ZIEGLER 3.1.3. Description of the holotype. Total length 128 mm (for more measurements and indices see Table 1). Rostral twice as broad as high, narrower than mental, without a median suture. 13/12 supra- and 11/10 sublabials. Nostril touches rostral and Ist supralabial. 3/3 nasals, nasorostrals in contact medially. Nasorostrals twice as big as postnasals. Pupil vertical. Tympanum obliquely oval, ca. half of the eye diameter. Interorbital scale count 44. Mental triangular, as broad as high. Two postmentals, twice as long as broad and of subequal length with the Ist supralabial. Postmentals posteriorly touched by six scales. Lateral scales of snout about three times bigger than those on the dorsal side of the snout, roundish, juxtaposed. Head scales granular, ciliaries posteriorly spiny. Gular scales granular, about the same size as upper head scales. Dorsal scales round, smooth, juxtaposed, convex, ca. 1.5 times bigger than head scales. Ventral scales flat, smooth, imbricate, three times as big as dorsals, 41 in an oblique row between the weakly developed lateral folds. Midbody scale count 140. Scales on the upper surface of the upper fore- and hindlimbs flat, smooth and imbricate, granular on the lower. Lower parts of all limbs with granular scales. Fingers and toes basally weakly webbed, only toes 4 and 5 without webbing. 14/12 lamellae under the Ist and 17/15 under the 4th toe. 5 preanal pores arranged in an angularly bent series, 3/3 postanal tubercles. Upper caudal scales flat, subimbricate, as big as the dorsals, ar- ranged in regular transverse rows and weakly expressed whorls, the 3rd whorl being composed of (dorsally) 10 scale rows. Subcaudals flat, smooth, imbricate, 2.5 times wider than high, with a median row of regularly arranged plates; each 3rd plate ca. 3 times as broad as high, marking the beginning of whorl. Laterally, the subcaudals are bordered by alternating scales (2-2-3-2- H. ROSLER, T. ZIEGLER, VU N.T., H.-W. HERRMANN & W. BOHME: A New Gekko Species from Vietnam 3 2-3-2-2 ...) where scales each are bordering the respec- tive broader subcaudal plate. Fig. 2: Ventral view of the live holotype of Gekko scientiad- ventura sp. n. (ZFMK 76198). Phot. T. ZIEGLER Hemipenial morphology. Right hemipenis (Fig. 3: terminology after RÖSLER 1998) 2.2 mm wide, 4.6. mm long, clavate, forked. Pedicel and truncus not dif- ferentiated from each other; basally on the sulcal side ıs a small wrinkled adminiculum, running laterocranially and not reaching the sperm groove. Above the admini- culum a ribbed area. Apex bilobed, the lobes being sub- equal in size. Sperm groove originates from the latero- caudal side of the organ and runs straight along the truncus, where it divides into two branches that termi- nate on the upper side of each lobe in a slit-like concay- ity. Apically, the sperm groove is shallow and is ac- companied by weakly developed sulcal lips. The sperm groove gets deeper and widens towards the proximal part of the truncus. The epidermal tissue of the lobes appears to be rough, although no calyces can be distin- guished. Colour of preserved holotype. Upper side of head and body dark grey. Head grey-brown, with darker vermicu- Fig. 3: Right hemipenis of the holotype of Gekko scientiad- ventura sp. n. (ZFMK 76198): a) sulcal, b) asulcal view. Dra- wing: H. ROSLER. lations and flecks, additionally small light dots. Dorsum with seven median light flecks that continue laterally into small transverse bands with strongly waved anterior and posterior margins. These transverse bands are ante- riorly bordered by brown-blackish lines and dissolve laterally. Between the single bands light flecks and nu- merous small spots are interspersed. Limbs above dark grey with grey-brown stripes. Throat, chest and belly buff, the throat bearing a reticulate pattern. Belly later- ally with light spots. Forepart of tail above and below dark grey, hindpart grey-black. Nine grey-brown trans- verse bands on the upper side of tail, the hindmost five reaching the lower side of the tail and nearly form closed rings. 3.1.4. Paratypes. ZFMK 76174 (Fig. 4), adult female, ZFMK 76175-179, juveniles, Phong Nha - Ke Bang, surroundings of 17.32 N, 106.16 E, 50-150 m a.s.l., Quang Binh Province, Vietnam, coll. H.-W. HERRMANN, VU NGOC THANH & T. ZIEGLER, 26 August - | September 2001. ZFMK 80651-652, adult females, same locality data, coll. VU NGOC THANH & T. ZIEGLER, 29 August - 10 September 2003. Paratypes ZFMK 76179 and 80652 will be permanently transferred to the scientific collection of the Vietnamese National University, Hanoi (HNUV). 3.1.5. Description of paratypes. Individual body meas- urements and proportions can be seen in Table |. For the latter, the paratypes have the following mean values and standard deviations (n being variable): SVL:TL 0.87+/-0.07, SVL:HL 3.49+/-0.24, HL:HW 1.28+/-0.03, HL:HH 2.28+/-0.03, SE:EE 1.33+/-0.09. All paratypes possess invariably 3 nasals, no internasals and 2 postmentals. Further, females and juveniles are lacking preanal pores. The postmentals are differently shaped, the maximal relative length is the left post- mental of ZFMK 76176 which is three times as long as 138 Bonner zoologische Beitráge 53 (2004) Fig. 4: Live female paratype of Gekko scientiadventura sp. n. (ZFMK 76174). Phot. T. ZIEGLER Fig. 5: Portrait of one of the captive males of Gekko scien- tiadventura sp. n. (see also fig. 28 in ZIEGLER et al. 2004) from 22 April 2003, measuring a SVL of 67 mm and a TL of 70 mm at the time; note the spiny posterior ciliaries as well as the bordering nasorostrals. Phot. T. ZIEGLER broad. The right postmental is divided in ZFMK 76176 and 76178. The posterior ciliaries are spiny in all para- types. The remaining scalation characters vary to a small degree only (see Table 2). Range (mean and stan- dard deviation) of the number of supralabials is 12-14 (13.00+/-0.82), of the sublabials 9-13 (9.94+/-1.26), of the scales behind postmentals 5-7 (5.67+/-0.71), of the interorbitals 41-49 (43.75+/-3.59), of the ventrals 38-48 (41.80+/-3.90), of the scales around midbody 118-139 (132.20+/-8.23), of the lamellae under the Ist toe 12-15 (13.86+/-1.03), of the lamellae under 4th toe 14-17 (15.64+/-1.15), of the postanal tubercles 2-3 (2.50+/- 0.53) and of the number of dorsal scales in the 3rd tail whorl 10-11 (10.29+/-0.49). All juvenile paratypes (as well as two embryos, see 3.1.6.) have still two egg teeth which touch each other apically as it is typical for geckos with relative thick and calcified eggshells (ROSLER 2001b). The tooth crown of the egg teeth has a shape similar to that of G. gecko (see SLUITER 1893). Colour of preserved paratypes. The pattern of the pre- served material corresponds to that of the living ani- mals, the overall colouration being more greyish. 3.1.6. Other material. Next to the type series, there are two male embryos (ZFMK 76199 and CPHR 2028) and three eggshells (ZFMK 76180) from the same locality. The embryonic males exibit egg teeth as decribed above, their hemipenes are completely everted, deeply forked and show two subequal apical lobes. In one of them (CPHR 8028) the dark pigments (6 crossbands) start to develop. In addition there are three males and one female (Figs. 5-7, see also fig. 28 in ZIEGLER et al. 2004) that had hatched in September/October 2001 from eggs found in a karst cave at the type locality in August 2001, and which are kept alive in the Cologne Zoo. 3.1.7. Colouration in life. Variable in all adults. Basic colour either more brownish marbled with yellowish flecks, or yellowish with brownish marbling. Upper side of head with large, blackish-brown flecks or spots which may be present also on the snout. Some speci- mens additionally possess some greyish flecks on the same head regions. Ciliary scales black with yellow margins. Iris metallic green-yellowish, pupil with bronze-coloured margin. A row of 7 middorsal grey flecks which may be extended to wavy crossbands of the same colour towards the flanks. These crossbands may be dissolved into flecks. The middorsal flecks are anteriorly bordered with a blackish brown pattern that is w-shaped in the neck region. The grey crossbars on the tail are likewise anteriorly bordered by blackish brown pattern elements. The grey flecks on the limbs are big- ger on the hind legs. Throat and belly with more or less distinct lemon to buff colouration, with more or less dis- tinct brown-olive marbling. Colour pattern of underside of tail variable, with pale or dark grey to ochre or black- ish brown flecks. H. ROSLER, T. ZIEGLER, VU N.T., H.-W. HERRMANN & W. BOHME: A New Gekko Species from Vietnam 139 Fig. 6: Gular region of the male of Gekko scientiadventura sp. n. depicted in Fig. 5 (10 November 2003). Phot. T. ZIEGLER Fig. 7: Captive female of Gekko scientiadventura sp. n. (10 November 2003); note the regenerated tail. Phot. T. ZIEGLER 3.1.8. Etymology. The species name is a patronym for a scientific magazine programme of the German Televi- sion channel ZDF called “Abenteuer Wissen” (adven- ture of knowledge) in order to acknowledge the first documentation of the Phong Nha - Ke Bang National Park biodiversity for the German public. First live pic- tures of Gekko scientiadventura sp. n. were shown on this TV programme. The name is derived from the Latin words “scientia” (knowledge, science) and “adventura” (literally = things that will come = noun in apposition. adventure) and is a 3.2. Distribution / So far, Gekko scientiadventura sp. n. is known only from the type locality (Fig. 8). Although currently re- garded as endemic for Vietnam, its occurrence in Laos cannot be excluded, considering the proximity of the Laotian border from the type locality. CHINA VIETNAM Hanoi O South China LAOS Sea CAMBODIA Fig. 8: Position of the type locality within Quang Binh Prov- ince. 3.3. Natural history All specimens of the type series were collected at night in primary forest, either directly on or in the immediate neighbourhood of the karst limestone outcrops that were partly overgrown with vegetation (Fig. 9). The geckos were mostly seen on the vegetation in low height (1.0- 2.5 m) rather than on the bare rocks. In contrast, these bare rocks were mostly populated by the syntopic and only recently described bent-toed gecko Cyrtodactylus phongnhakebangensis which, in turn, tended to avoid 140 Bonner zoologische Beitráge 53 (2004) the vegetational parts of the microhabitat (ZIEGLER et al. 2002). Fig. 9: Habitat of Gekko scientiadventura sp. n. near the type locality. Phot. T. ZIEGLER Apart from the voucher material of our type series no further sightings of Gekko scientiadventura sp. n. were made, with the exception of a mass egg-laying site in a dark, manifold rock crevice system at the type locality. Fig. 10 shows this site (in about 2.5 m height) at the be- ginning of the rainy season (end of August) in 2001. The four geckos cited under “Other material” originate from these eggs; they hatched in September/October 2001 which correlated with the beginning rains. One year later, in August 2003, we found two glued eggs just at this mass egg-laying site. In addition, two further communal egg-laying places were found in this cave system which seemed to be used also already since a longer period; they contained, however, much less eggs and were situated much higher (ca. 5 m high). On August 30, 2003, we measured at the lower egg- laying site, at 14.30 h, a relative humidity of 85% and a temperature of 27.3 °C. At 21.40 h the same day the temperature was 26.3 °C and the relative humidity had Fig. 10: Mass egg-laying site of Gekko scientiadventura sp. n. in a karst rock crevice at the type locality. Phot. T. ZIEGLER increased to 93% (with a recorded minimum of 70% be- tween 14.30 and 21.40 h). In the primary forest, outside the rock crevice system, we measured with a minimum- maximum thermometer on August 28, 2001 24.4-30.2 °C and one day later 24.3-27.8 °C. For further (long- term) climatic data from this area see HERRMANN et al. (2002). One of the authors (TZ) was able to record some behav- ioural and autecological observations of the captive- raised geckos: The growling calls of Gekko scientiad- ventura sp. n. were mostly uttered during dawn and night, but could also be heard in summer afternoons (e.g. 22-26 July, 2002), 1.e. during the light phase of the terrarium. Obviously, also the black-and-white annula- tion of the tail plays a role in the infraspecific commu- nication between both sexes, because in an interacting couple the tails were raised and displayed with undulat- ing movements. A male that was taken out of its terrar- ium used to wag its tail in a much faster manner. Once an acinetic reaction could be observed, a lethisimulation with a recurved tail and extremities that were raised from the ground and standing off from the body. The geckos were fed mainly with crickets of various size classes that were powdered with a vitamin-mineral mix- ture, however, they were repeatedly observed to lick on and even to bite off small pieces of bananas. In 2003, two egg-layings occured but the paired, partly deformed clutches proved to be unfertilised. 3.4. Comparison with other Vietnamese Gekko species Next to the lack of dorsal tubercles, Gekko scientiadven- tura sp. n. can be distinguished from its Vietnamese congeners by the combination of the characters size, number of internasals and preanal pores as well as dor- sal pattern: G. badenii reaches a SVL up to 76.5 mm and has 1-3 in- ternasals; males have 14-18 preanal pores, and the dor- sal pattern consists of 4-8 very narrow and sometimes H. ROSLER, T. ZIEGLER, VU N.T., H.-W. HERRMANN & W. BOHME: A New Gekko Species from Vietnam 14] even interrupted dorsal bands (SZCZERBAK & NEK- RASOVA 1994). G. chinensis (synonym: G. semipalmatus Stejneger, 1932) reaches a SVL up to 70 mm. Always | internasal which exceeds the nasorostral in size. Males have 18-24 preanal pores. Dorsally 4 indented, cloudy crossbands with light flecks in between (OTA et al. 1995). G. semi- palmatus has 10 more or less regularly arranged longi- tudinal rows of dorsal tubercles and the males have 23 preanal pores (STEJNEGER 1932). G. gecko of Vietnamese origin reach a SVL up to 173 mm. 0-1 internasal. Males have 13-20 preanal pores. Dorsum with transversely and more or less regularly ar- ranged light flecks and dark (brownish to reddish) bands, flecks and stripes (own data). We recognize the following synonyms of Gekko gecko (sensu lato): G. verticillatus Laurenti, 1768; G. teres Laurenti, 1768; G. perlatus Houttuyn, 1782; G. guttatus Daudin, 1802; G. verus Merrem, 1820; G. annulatus Kuhl, 1820; G. in- dicus Kuhl, 1857. All taxa considered as synonyms here have (according to their descriptions) dorsal tubercles. We are uncertain in synonymizing G. aculeatus Hout- tuyn, 1782 with G. gekko (see also KLUGE 1993, 2001). DAUDIN (1802) considers this name to be synonymous with Tarentola mauritanica Linnaeus, 1758 (see also ROSLER 2001 a). Moreover, we recognize the subspe- cies G. g. azhari Mertens, 1955. Validity and status of G. reevesii Gray, 1831 will be discussed elsewhere (ROSLER & ZIEGLER in prep.). Both last-named forms differ by their size, the dorsal tubercles and the colour pattern unequivocally from G. scientiadventura sp. n. G. grossmanni reaches a SVL of 89 mm. 0-1 internasal. Males with 12-14 preanal pores. Dorsum with several rows of more or less transversally arranged light flecks (GUNTHER, 1994). G. japonicus reaches a SVL length of 74 mm and has 1- 2 internasals. Males have 6-9 preanal pores. Dorsum with 6 indented dark crossbands which are medially in- terrupted by light flecks (STEINEGER 1907; ZHAO et al. 1999; own data). BOURRET (1937) described a male G. japonicus from Ngan Son, Tonkin (northern Vietnam) with a SVL of 71 mm as “identique a la description originale, avec une ligne de 10 pores préanaux de chaque cóté” (see also BOURRET 1939 and his unpub- lished manuscript, pp. 34-36). A second locality, Bac Thai, was recorded by NGUYEN & Ho (1996). Accord- ing to OTA et al. (1995), the gecko described by BOUR- RET belongs to G. palmatus. Synonyms of G. japonicus are Platydactylus jamori Temminck & Schlegel, 1838 and Hemidactylus nanus Cantor, 1842. The original de- scription of A. nanus contains only notes on the colour pattern (CANTOR 1842), but both types (BMNH 1946.8.26.9-10) have tubercles on back, limbs and tail (pers. comm. C. MCCARTHY). G. palmatus reaches a SVL of 79 mm and has 0-2 inter- nasals. Males have 24-27 preanal pores. Dorsum includ- ing occipital and nuchal region with 6 rows of dark flecks with lighter interspaces (OTA et al. 1995). G. ulikovskii reaches a SVL up to 108 mm and has one internasal. Males have 10-15 preanal pores. Dorsum with up to eight narrow, light dorsal bands (DAREVSKY & ORLOV 1994). 3.5. Comparisons with the remaining Gekko species According to the original descriptions of all Gekko spe- cies not occurring in Vietnam, the lack of dorsal tuber- cles sets G. scientiadventura sp. n. unequivocally apart from: G. albofasciolatus Gúnther, 1867; G. auriverru- cosus Zhou & Liu, 1982; G. gigante Brown & Alcala, 1978; G. hokouensis Pope, 1928 (synonym: Luperosau- rus amissus Taylor, 1962); G. kikuchii Oshima, 1912: G. liboensis Zhou & Li, 1982; G. mindorensis Taylor, 1919; G. monarchus Schlegel, 1836 (synonyms: Platy- dactylus burmeisteri Giebel, 1861; P. deissneri Giebel, 1861); G. palawanensis Taylor, 1925: G. petricolus Taylor, 1962; G. porosus Taylor, 1922; G. romblon Brown & Alcala, 1978; G. scabridus Liu & Zhou, 1982; G. siamensis Grossmann & Ulber, 1990 (synonym: G. tavlori Ota & Nabhitabhata, 1991); G. similignum Smith, 1923; G. smithii Gray, 1842 (synonyms: Platydactylus stentor Cantor, 1847; P. albomaculatus Giebel, 1861): G. swinhonis Günther, 1864; G. verreauxi Tytler, 1864; G. vittatus Houttuyn, 1782 (synonyms: Lacerta unistri- ata Shaw, 1792; Stellio bifurcifer Schneider, 1792: Platydactylus bivittatus Duméril & Bibron, 1836: G. trachylemus Peters, 1872); G. yakuensis Matsui & Okada, 1968. According to the English summary of the Chinese origi- nal description of G. taibaiensis Song, 1985, this spe- cies is closely related to G. swinhonis and G. japonicus (SONG 1985). From these, G. taibaiensis differs by fewer preanal pores (4-6) which are arranged in an in- terrupted order. The maximum SVL is 69 mm, and there are 7-8 subdigital lamellae under the fourth toe (ZHAO et al. 1999). G. taibaiensis differs from G. scientiadven- tura by its significantly lower number of subdigital la- mellae under the 4th toe and the series of preanal pores which is medially interrupted by a smooth scale. The specimens figured by SONG (1985) and HUANG & ZONG (1998) do not show a contrasting dorsal pattern as does G. scientiadventura sp. n. There are only few species in the genus Gekko that have a lack of dorsal tubercles in common with G. scientiad- ventura sp. n. These are G. athymus Brown & Alcala, 1962, G. subpalmatus Gúnther, 1864, and G. tawaensis Okada, 1956. G. athymus reaches a SVL of 100-120 mm, and has one internasal. Digits webbed for 1 third of their length. 19- 142 Bonner zoologische Beitráge 53 (2004) 21 subdigital lamellae under 4th toe. Males with 22-23 preanal and femoral pores; subcaudals transversely broadened (BROWN & ALCALA 1962, 1978). Fig. 11: Dorsal view of the holotype of Gekko subpalmatus (BMNH 1946.8.2592). Phot. T. ZIEGLER G. subpalmatus reaches a SVL of 78 mm and has 1-2 internasals. Toes basally narrowly webbed. 7-10 sub- digital lamellae under 4th toe. Males with 5-11 preanal pores. Subcaudals transversely broadened (GUNTHER 1864; POPE 1935; ZHOU et al. 1989; ZHAO et al. 1999). For a better comparison with G. scientiadventura sp. n. we studied the holotype of Gekko subpalmatus (BMNH 1946.8.2592) and the lectotype and the paralectotype of Gekko melli Vogt, 1922 (ZMB 27659 A & B) which is considered to be synonymous with G. subpalmatus (e.g. ZHAO & ADLER 1993) (Figs. 11-13): The rostral of the holotype of G. subpalmatus has a short upper transverse suture, the posterior ciliaries are pointed, and there are five distinct tubercles anterior of the tympanum on the right side of the head. The dorsum is lilac-grey to violet- grey, a light postocular stripe and four longish grey- brown vertebral flecks are barely visible, and the tail has several broad light transverse bands that are bordered by darker pigment. The lectotype and the paralectotype of G. melli lack a rostral suture, the posterior ciliaries are likewise pointed, and pretympanic tubercles are lacking. The dorsum of ZMB 27659 A is olive-brown. There is a dark naso-ocular and a light postocular stripe. Two dark stripes are running from the ear openings to the axillae, and the shoulder region shows two sepia-coloured stripes. The dorsum has three broad, irregular and partly confluent, grey-brown bands with sepia-coloured mar- gins, the tail has four broad pale brown bands that are likewise bordered by darker pigment. The regenerated tail tip has fine darker dots. Apart from their colour pat- tern, the types of both taxa differ also in size and in body proportions (SVL/TL index) and in the number of interorbitals, subdigital lamellae under the 4th toe, and scale rows in the third caudal whorl (see Tab. 3). Pend- ing future studies on more material we consider G. melli therefore as valid. Fig. 12: Portrait of the holotype of Gekko subpalmatus (BMNH 1946.8.2592). Phot. T. ZIEGLER Fig. 13: Dorsal view of the lectotype of Gekko melli (ZMB 27659 A). Phot. H. ROSLER G. tawaensis reaches a SVL up to 70 mm and has inter- nasals. The toes are basally narrowly webbed, and there are 12 subdigital lamellae under the 4th toe. The males have no preanofemoral pores, and the subcaudals are anteriorly mesially divided (OKADA 1956; SENGOKU 1989; UTSUNOMIYA et al. 1996). G. scientiadventura sp. n. differs from the other Gekko species without dorsal tubercles by the combination of following characters: (a) SVL, (b) internasals, (c) exten- sion of webbing between toes, (d) subdigital lamellae under 4th toe, (e) number of preanal pores, and (f) shape of subcaudals. G. scientiadventura sp. n. differs thus from G. athymus in the characters a, b, d, e; from G. subpalmatus in the characters a, b, c, d; and from G. tawaensis in the characters b, e, and f. 4. DISCUSSION The genus Gekko Laurenti, 1768 contains, according to the last overview by KLUGE (2001) 28 species. We add to this number five more species which we consider as valid: G. liboensis Zhou & Li, 1982, G. melli Vogt, 1922, G. mindorensis Taylor, 1919 G. scabridus Liu & Zhou, 1982, and G. scientiadventura Sp. n. H. RÖSLER, T. ZIEGLER, VU N.T., H.-W. HERRMANN & W. BOHME: A New Gekko Species from Vietnam 143 ZHAO & ADLER (1993) formally synonymize G. liboen- sis with G. hokouensis. KLUGE (1993), WELCH (1994), MATSUI & OTA (1995) and ZHAO et al. (1999) regard G. hokouensis as valid. It has 10 irregular rows of dorsal tubercles and 5-9 rows of prenanal pores (ZHOU et al. 1989; ZHAO et al. 1999). According to BAUER (1994) who refers to a personal communication by R. CROMBIE, G. mindorensis is a synonym of G. kikuchii. Both possess dorsal tubercles (TAYLOR 1919). However, G. mindorensis has more preanofemoral pores and longer limbs. Moreover, it has broad, medially divided, dark dorsal crossbands (FERNER et al. 2001; GAULKE 2003), whereas G. kiku- chii has a double series of dark flecks on the dorsum (see the figs. in LU et al. 1999 and in HENKEL & SCHMIDT 2003). ZHAO & ADLER (1993) formally synonymize G. scabridus with G. chinensis, whereas MATSUI & OTA (1995) and ZHAO et al. (1999) regard the former as valid. G. scabridus has 17-21 irregular, longitudinal rows of dorsal tubercles and 10-15 preanal pores (ZHOU et al. 1989). The presence or lack of dorsal tubercles does not occur solely in the genus Gekko. Also other gekkonine genera, e.g. Hemidactylus Gray, 1825, and Pachydactylus Wiegmann, 1834, contain species with or without dorsal tubercles (SMITH 1935; FITZSIMONS 1943; LOVERIDGE 1947). The majority of Gekko species has these tuber- cles (SENGOKU 1989). Additionally, species-specific tu- bercles may also exist on the upper surfaces of head, limbs and tail. The tubercles may be rather big, as in G. monarchus, or very small, as in G. ulikovskii. Neither their relative size nor their complete absence are geo- graphically correlated. Of the four tubercleless species, only G. scientiadventura sp. n., G. melli and G. subpal- matus are mainland species (Vietnam, southern China), whereas G. fawaensis is an endemic of Japan (southern Honshu und some small satellite islands) and G. athy- mus is an endemic of Palawan Island (Philippines). Apart from the lack of dorsal tubercles, the four species have only few characters in common. The dorsal and tail pattern of G. scientiadventura sp. n. resembles most G. tawaensis (see fig. in UTSONOMIYA et al. 1996). Also in G. subpalmatus, the dorsal pattern consists of medi- ally arranged light flecks, and the tail is banded (see fig. in ZHAO & ADLER 1993). The dorsal pattern of G. athymus consists of indistinct, broad, wavy bands (BROWN & ALCALA 1962). However, light vertebral ar- eas or flecks/bands respectively occur more or less con- spicuously also in other Gekko species (among else in G. auriverrucosus, G. chinensis, G. hokouensis, G. ja- ponicus, G. similignum, G. swinhonis) so that closer re- lationships among the tubercleless Gekko species cannot be deduced from this character. Phenotypically G. scientiadventura differs among the four tubercleless species most from G. athymus which is the geographically remotest (see above). From G tawaensis, it differs by the anteriorly undivided subcau- dals. Transversally widened (plesiotypic) versus divided (apotypic) subcaudals are not an exclusive character for Gekko, but are also common in the genera, e.g., Lygo- dactylus Gray, 1854 and Phelsuma Gray, 1825 (PAS- TEUR 1964; LOVERIDGE 1942). A closer relationship of G. scientiadventura sp. n. to the two insular species G. athymus and G. fawaensis seems unlikely to us. Rather, our new species seems to be closer to G. melli and G. subpalmatus which has to be proven by future (molecular) studies. Both species, however, lack the striking spiny ciliary scales, and they have much broader webbings between the toes. G. sub- palmatus differs strikingly from G. scientiadventura sp. n. by its uniform colour pattern (see figs. in ZHAO & ADLER 1993; HUANG & ZONG 1998). Less uniform is G. melli, but its dorsal bands are more roughly shaped as compared with the finely structured dorsal pattern of G. scientiadventura sp Nn. 5. KEY TO THE GEKKO SPECIES OF VIETNAM I Dorsaltubereleslackine 2.2... 22. a Gekko scientiadventura sp. n. I RD ORSALNUBERCIES PRESEME tabla 2 SD Pe eee G. gecko De E A A OS 3 3 Broad webbing between toes ....oooocinocccnocccooccccocccconos: + 3” Narrow webbing between toes ..............::ccssceeeeeeeeees 5 4 1 Internasal, larger than nasorostrals....... G. chinensis 4” 0-2 Internasals, smaller than nasorostrals.. G. palmatus 5 Males with less than 10 preanal pores.... G. japonicus 5” Males with more than 10 preanal pores .................... 6 6: Dorsum with blotches: 2-5-2230... 4. G. grossmanni GD Orsini With DADOS cuate tia nennen 7 7 30-37 Interorbitals, southern Vietnam ....... G. badenii 7° 40-46 Interorbitals, central Vietnam ....... G. ulikovskii Acknowledgements. We thank Prof. Vo Quy and Prof. Dr. Truong Quang Hoc (Centre for Natural Resources Man- agement and Environmental Studies, Vietnam National University, Hanoi) for their continuous assistance and en- couragement. The People's Committee of Quang Binh, the Forestry Planning Department of Quang Binh and the Phong Nha - Ke Bang National Park administration issued work and collecting permits. We are especially indebted to the National Park director Neuyen Tan Hiep and his vice directors Cao Xuan Chinh and Luu Minh Thanh, as well as Dinh Huy Tri, the director of the Science Research Center. The National Park staff and our guide Bui Ngoc Thanh helped to make fieldwork successful. We wish to thank Dr. Rainer Gúnther (ZMB, Berlin) and Dr. Colin MeCarthy 144 Bonner zoologische Beitráge 53 (2004) (BMNH, London) for the loan of specimens under their care. We are greatful to Dr. Ivan Ineich (Muséum National d'Histoire Naturelle, MNHN, Paris), Dr. Hidetoshi Ota (Tropical Biosphere Research Center, University of the Ryukyus, Okinawa) and Dr. Franz Tiedemann (Naturhis- torisches Museum Wien, NMW, Vienna) for their support and for fruitful discussions. The field work of H.-W.H. and T.Z. was funded in large by the Zoological Garden Co- logne, the Kölner Kulturstiftung der Kreissparkasse Köln and BIOPAT. Last but not least we are grateful to the German Television channel ZDF for supporting biodiver- sity research and nature conservation in the Phong Nha - Ke Bang National Park. REFERENCES BAUER, A. M. (1994): Gekkonidae (Reptilia, Sauria), Part I Australia and Oceania. Das Tierreich, 109: I-XII, 1- 306. Walter de Gruyter, Berlin, New York. BOBROV, V. V. (1995): Checklist and bibliography of the Lizards of Vietnam. Smithsonian Herpetological In- formation Service 105: 1-28. BOULENGER, G. A. (1907): Description of new lizard in the British Museum. Annals and Magazine of natural His- tory, London 19: 486-489. Bourret, R. (1937): Notes herpétologiques sur l'Indochine francaise. XII. Les lézards de la collection du Labora- toire des Sciences Naturelles de l'Université. Descrip- tion de cinq especes nouvelles. Annexe au Bulletin General de l’Instruction Publique, Hanoi 9: 3-22. BOURRET, R. (1939): Notes herpétologiques sur I'Indochine francaise. XX. Liste des reptiles et batraciens actuel- lement connus en Indochine francaise. Annexe au Bul- letin Général de l’Instruction Publique, Hanoi 4 (dé- cembre): 49-60. BOURRET, R. (unpubl.): Les lézards de l'Indochine. Un- published manuscript, Museum National d’Histoire Naturelle, Paris. BROWN, W. C. & ALCALA, A. C. (1962): A new lizard of the genus Gekko from the Philippine Islands. Proceed- ings of the Biological Society of Washington 75: 67- 70. Brown, W. C. & ALCALA, A. C. (1978): Philippine lizards of the family Gekkonidae. Silliman Univ. Press, Monograph Series, No. 1: 1-146. CANTOR, T. (1842): General features of Chusan, with re- marks on the flora and fauna of that island. Annals and Magazine of natural History, London 9: 481-493. CANTOR, T. (1847): Catalogue of reptiles inhabiting the Malayan Pininsula and islands, collected or observed by Theodore Cantor. Journal of the Asiatic Society of Bengal 16 (2): 1-159. (Reprint 1981, New Delhi, Cosmo Publications.) DAREVSKY, I. S. & ORLOV, N. L. (1994): Eine bemerkens- werte neue, großwüchsige Art der Gattung Gekko: Gekko ulikovskii sp. nov. aus Zentralvietnam. Sala- mandra, Bonn 30 (1): 71-75. DAUDIN, F. M. (1802): Histoire naturelle, générale et parti- culiere des Reptiles. Vol. 4, 397 pp., F. Dufart, Paris. DUMERIL, A. M. C. & BIBRON, G. (1836): Erpétologie gé- nérale ou histoire naturelle complete des reptiles 3: I- IV, 1-517. Roret, Paris. FERNER, J. W., BROWN, R. M., SISON, R. V. & KENNEDY, R. S. (2001): The amphibians and reptiles of Panay Is- land, Philippines. Asiatic Herpetological Research 9: 34-70. FITZSIMONS, V. F. (1943): The lizards of South Africa. Transvaal Museum, Pretoria 1: 1-528. GAULKE, M. (2003): Ubersicht der philippinischen Gecko- Fauna. Gekkota, Quedlinburg 4: 3-24. GIEBEL, C. (1861): Drei neue Arten Platydactylus von der Insel Banka. Zeitschrift für die gesammten Natur- wissenschaften, Berlin, 1861: 58-60. GIRARD, C. (1857): Descriptions of some new reptiles, col- lected by the United States Exploring Expedition, un- der the command of Capt. Charles Wilkes, U. S. N. Part 4: Including the species of Saurians, exotic to North America. Proceedings of the Academy of natu- ral Sciences Philadelphia 9: 195-199. GRALLERT, W. & J. ROLAND (1960): Lipsia — Philate- listische Farbtafeln. Philatelistische Schriftreihe, Heft 6, VEB Graphische Werkstatten, Leipzig, 11 S, Farb- taf. unpagin. GRAY, J. E. (1831): A synopsis of the class reptilia. Pp. 1- 110 in: GRIFFITH, E. & PIDGEON, E. (eds.): The animal kingdom arranged in conformity with its organization, by the Baron Cuvier. Vol. 9, Whittaker, Treacher, and Co., London. GRAY, J. E. (1842): Description of some new species of reptiles, chiefly from the British Museum collection. The Zoological Miscellany, London 1842: 57-59. GROSSMANN, W. & ULBER, T. (1990): Ein neuer Gecko aus Zentral-Thailand: Gekko siamensis sp. nov. (Rep- tilia: Sauria: Gekkonidae). Sauria, Berlin 12 (3): 9-18. GUNTHER, A. (1864): The reptiles of British India. I- XXVII, 1-452, Hardwicke, London. GUNTHER, A. (1867): Additions to the knowledge of Aus- tralian reptiles and fishes. Annals and Magazine of na- tural History, London 20: 45-57. GUNTHER, R. (1994): Eine neue Art der Gattung Gekko (Reptilia, Squamata, Gekkonidae) aus dem Súden Vietnams. Zoologischer Anzeiger 233(1/2): 57-67. HENKEL, F. W. & SCHMIDT, W. (2003): Praxis Ratgeber Geckos. 159 pp., Edition Chimaira, Frankfurt am Main HERRMANN, H.-W., ZIEGLER, T., STUART, B. L. & ORLOV, N. L. (2002): New findings on the distribution, mor- phology and natural history of Triceratolepidophis sieversorum (Serpentes: Viperidae). Herpetological Natural History, La Sierra 9(1): 89-94. Houttuyn, M. (1782): Het onderschied der salamanderen van de haagdissen in t algemeen, en van de gekkos in t byzonder, aangetood. Verhandelingen van het Ge- nootschap der Wetenschappen Vlissingen, Middelburg 9: 305-336. HUANG, Z. Y. £ Y. ZONG (1998): Zhongguo Techan de Paxing Dongwu. 116 pp., Fudam Universitat, Shang- hal. KLUGE, A. G. (1993): Gekkonoid lizards taxonomy. 245 pp., International Gecko Society, San Diego. KLUGE, A. G. (2001): Gekkotan lizard taxonomy. Hama- dryad 26 (1): 1-209. KUHL, H. (1820): Beitráge zur Kenntnis der Amphibien. Pp 77-132 in: Beitráge zur Zoologie und vergleichenden Anatomie 1. Herrmansche Buchhandlung, Frankfurt am Main. LAURENTI, J. N. (1768): Specimen medicum, exhibens sy- nopsin reptilium emendatam cum experimentis circa venena et antidota Reptilium Austracorum. 216 pp., H. ROSLER, T. ZIEGLER, VU N.T., H.-W. HERRMANN & W. BÖHME: A New Gekko Species from Vietnam 145 Joan Thomae, Viennae, (Reprint 1966, A. Asher & Co, Amsterdam). LEVITON, A. E., GIBBS, R. H., HEAL, E., DAWSON, C. E. (1985): Standards in herpetology and ichthyology: part I. Standard symbolic codes for institutional resource collections in herpetology and ichthyology. Copeia, Washington 1985: 802-832. LINNAEUS, C. (1758): Systema nature per regna tria nature, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. 10. Edition, Vol. I. L. Salvii, Holmiae. Liu, Y-Z. € ZHOU, K-Y. (1982): Gekko scabridus. Pp.438- 440 (445) in: ZHOU, K-Y., Liu, Y-Z. & Li, D-J. Three new species of Gekko and remarks on Gekko hokouen- sis (Lacertiformes, Gekkonidae). Acta Zootaxonomica Sinica 7 (4): 438-446. LOVERIDGE, A. (1942): Revision of the afro-oriental geckos of the genus Phelsuma. Bulletin of the Mu- seum of comparative Zoology, Cambridge, Mass. 89: 439-482. LOVERIDGE, A. (1947): Revision of the African lizards of family Gekkonidae. Bulletin of the Museum of com- parative Zoology, Cambridge, Mass. 98: 1-469. Lu, G., DU, M. & SHI, X. (1999): A field guide to the rep- tiles and amphibians of Taiwan. 343 pp., Taipeh. MATSUI, T. & OKADA, Y. (1968): A new species of Gekko found in Yakushima, one of the small island south of Kyushu. Acta Herpetologica Japonica 3: 1-4. MATSUI, M. & OTA, H. (1995): On chinese herpetology. Herpetologica, Lawrence 51 (2): 234-250. MERREM, B. (1820): Versuch eines Systems der Amphi- bien. I-XV, 1-78, Johan Christian Krieger, Marburg. NEKRASOVA, O. D. & SZCZERBAK, N. N. (1993): The geckoes from South of Vietnam. 7th Ordinary General Meeting Societas Europaea Herpetologica, Barcelona, Programme and Abstracts, 104. NGUYEN VAN SANG & HO THU Cuc (1996): Danh luc bo sat va ech nhai Viet Nam. Nha xuat ban khoa hoc va ky thuat. 264 pp., Ha Not. OKADA, Y. (1956): A new species of gekko from Shikoku, Japan. Annotationes Zoologicae Japonenses 29: 239- 241. OsHIMA, M. (1912): Description of a new gecko from Botol Tobago Island. Philippine Journal of Science 7: 241-242. OTA, H., LAU, M. W., WEIDENHÓFER, T., YASUKAWA, Y. de BOGADEK, A. (1995): Taxonomic review of the geckos allied to Gekko chinensis Gray, 1842 (Gekkonidae, Reptilia) from China and Vietnam. Tropical Zoology 8: 181-196. OTA, H. & NABHITABHATA, J. (1991): A new species of Gekko (Gekkonidae: Squamata) from Thailand. Co- peia 1991(2): 503-509. PASTEUR, G. (1964): Recherches sur l'évolution des Lygo- dactyles, lézards Afro-Malgaches actuels. Travaux de l'Institut Scientifique Cherifien, Rabat 29: 1-132. PETERS, W. C. H. (1872): Uber eine, zwei neue Gattungen enthaltende, Sammlung von Batrachiern des Hrn. Dr. O. Wucherer aus Bahia, so wie tiber einige neue oder weniger bekannte Saurier. Monatsberichte der König- lichen Preussischen Akademie der Wissenschaften Berlin 1872 (Nov.): 768-776. POPE, C. H. (1928): Four new snakes and a new lizard from south China. American Museum Novitates 325: 1-4. Pope, C. H. (1935): The reptiles of China. Turtles, Croco- dilians, Snakes, Lizards. Natural History of Central Asia, New York 10: 1-604. ROSLER, H. (1998): Studien an den Begattungsorganen der Geckos (Reptilia: Gekkota) 1. Externmorpho- logische Merkmale und ihre Terminologie sowie móg- liche Ursachen für Fehldeutungen in der Merkmafs- analyse an evertierten Hemipenes. Gekkota 1: 36-64. RÖSLER, H. (2001 a): Studien am Tokeh: 1. Gekko gecko azhari Mertens, 1825 (Sauria: Gekkonidae). Gekkota 3: 33-46. ROSLER, H. (2001 b): Die Eızähne einiger Geckos (Sauria: Gekkota). Gekkota 3: 99-124. SCHLEGEL, H. (1836): Le Platydactyle de Japon. Platydac- tylus Japonicus. Pp. 337-338 in: DUMERIL, A. M. C. & BIBRON, G. Erpétologie générale ou histoire naturelle complete des reptiles. Vol. 3, Roret, Paris. SCHLEGEL, H. (1836): Le Platydactyle Monarque. Platy- dactylus Monarchus. Pp. 335-336 in: DUMERIL, A. M. C. & BIBRON, G. Erpétologie générale ou histoire na- turelle complete des reptiles. Vol. 3, Roret, Paris. SCHLÜTER, U. (1999): Haltung und Zucht von Laos- Geckos (Gecko petricolus). Deutsche Aquarien- und Terrarien-Zeitschrift 52 (9): 35-37. SENGOKU, S. (1989): Notes on a gekko population in the Danjo Group, Nagasaki Prefecture, Japan. Pp. 180-184 in: MATSUI, M., HIKIDA, T. & Goris, R. C. (eds.): Current herpetology in East Asia. Kyoto. SHAW, G. (1792): The single-striped lizard. Naturalist’s Miscellany 3: [1-2]. SLUITER, C. P. (1893): Uber den Eizahn und die Eischwiele einiger Reptilien. Morphologisches Jahrbuch 20: 75- 89. SMITH, M. A. (1923): On a collection of reptiles and batra- chians from the island of Hainan. Journal of the Natu- ral History Society of Siam 6: 195-212. SMITH, M. A. (1935): The fauna of British India, including Ceylon and Burma. Reptilia and Amphibia. | Loricata, Testudines. 185 pp., Taylor & Francis Ltd., London. SONG, M. (1985): A new species of Gekko from Shaanxi. Acta, Herpetologica Sinica, Chengdu, 4(4): 329-330. STEJNEGER, L. (1907): Herpetology of Japan and adjacent territory. Bulletin / Smithsonian Institution, United States National Museum, Washington, D.C. 58: I- XVII, 1-577. STEJNEGER, L. (1932): The Chinese lizards of the genus Gekko. Proceedings of the United States National Mu- seum 132: 1-6. SZCZERBAK, N. N. & NEKRASOVA, O. D. (1994): A contri- bution to the knowledge of gekko lizards of southern Vietnam with description of a new species (Reptilia, Gekkonidae). Vestnik zoologii 1: 48-52 (in Russian). TAYLOR, E. H. (1919): New or rare Philippine reptiles. Philippine Journal of Science 14: 105-125. TAYLOR, E. H. (1922): Additions to the herpetological fauna of the Philippine Islands. I. Philippine Journal of Science 21: 161-206. TAYLOR, E. H. (1925): Additions to the herpetological fauna of the Philippine Islands. IV. Philippine Journal of Science 26: 97-111. TAYLOR, E. H. (1962): New Oriental reptiles. University of Kansas Science Bulletin, Lawrence 43: 209-263. TYTLER, R. C. (1864): Oberservations on a few species of gecko alive in the possesion of the author. Journal of the Asiatic Society Bengal 33: 535-548. 146 Bonner zoologische Beitráge 53 (2004) UTSONOMIYA, T., UTSONMIYA, Y., OH KAWA, H., OKADA, J. &NAITOW, J. (1996): The amphibians and reptiles of Hiroshima prefecture. 168 pp., Hiroshima. VOGT, T. (1922): Zur Reptilien- und Amphibienfauna Süd- chinas. Archiv fiir Naturgeschichte, Berlin 10: 135-146. WELCH, K. R. G. (1994): Lizards of the world. 1. Geckos. 165 pp., The Longdunn Press, Bristol. ZHAO, E. & ADLER, K. (1993): Herpetology of China. Con- tributions to Herpetology, St. Louis 10: 1-521. ZHAO, E., ZHAO, K. & ZHOU, K. (1999): Reptilia. Pp. I-IIl, 1-394 in: HONGFU, Z., ZUOXIN, Z. & DAXIANG, S. (eds.): Fauna Sinica. Science Press, Beijing. ZHOU, K-Y. & Li, D-J. (1982): Gekko liboensis. Pp. 440- 441 (445) in: ZHOU, K-Y., Liu, Y-Z. & Li, D-J. Three new species of Gekko and remarks on Gekko hokouen- sis (Lacertiformes, Gekkonidae). Acta Zootaxonomica Sinica 7 (4): 438-446. ZHOU, K-Y. & LIU, Y-Z. (1982): Gekko auriverrucosus. Pp. 443-446 (446) in: ZHOU, K-Y., Liu, Y-Z. & LI, D-J. Three new species of Gekko and remarks on Gekko hokouensis (Lacertiformes, Gekkonidae). Acta Zootaxonomica Sinica 7(4): 438-446. ZHOU, K., Liu, Y. & Li, D. (1989): Three new species of Gekko and remarks on Gekko hokouensis (Lacertifor- mes, Gekkonidae). Smithsonian Herpetology Informa- tion Service 77: 1-10. ZIEGLER, T. & HERRMANN, H.-W. (2000): Preliminary list of the herpetofauna of the Phong Nha - Ke Bang area in Quang Binh province, Vietnam. Biogeographica, Paris 76(2): 49-62. ZIEGLER, T., HERRMANN, H.-W., VU NGOC THANH, LE KHAC QUYET, NGUYEN TAN HIEP, CAO XUAN CHINH, Luu MINH THANH & DINH Huy TRI (2004): The am- phibians and reptiles of the Phong Nha - Ke Bang Na- tional Park, Quang Binh Province, Vietnam. Hama- dryad, Tamil Nadu 28 (1-2): 19-42. ZIEGLER, T., ROSLER, H., HERRMANN, H.-W. & VU NGOC THANH (2002): Cyrtodactylus phongnhakebangensis sp. n., ein neuer Bogenfingergecko aus dem anna- mitischen Karstwaldmassiv, Vietnam. Herpetofauna, Weinstadt 24 (141): 11-25. Authors’ addresses: Herbert ROSLER, Staatliche Natur- historische Sammlungen Dresden, Museum fiir Tier- kunde, Forschungsstelle; A.B.-Meyer-Bau, K6nigsbrii- cker Landstr. 159, D-01109 Dresden, Germany; Corresponding address: F.-Freiligrath-Str. 51, D-06502 Thale am Harz, Germany; Dr. Thomas ZIEGLER (corre- sponding author), AG Zoologischer Garten Köln, Riehler Str. 173, D-50735 Köln, Germany, E-mail: tziegler@zoo-koeln.de; VU Ngoc Thanh, CRES, Centre for Natural Resources and Environmental Studies, Viet- nam National University, Hanoi, University of Science, Faculty of Biology, Department of Vertebrate Zoology, Zoological Museum, 334 Nguyen Trai Str., Thanh Xu- an, Hanoi, Vietnam; Dr. Hans-Werner HERRMANN, Conservation and Research for Endangered Species (CRES), Zoological Society of San Diego, 15600 San Pasqual Valley Road, Escondido, CA 92027-7000, USA; Prof. Dr. Wolfgang BOHME, Zoologisches For- schungsmuseum Alexander Koenig, Adenauerallee 160, D-53113 Bonn, Germany H. RÖSLER, T. ZIEGLER, VU N.T., H.-W. HERRMANN & W. BOHME: A New Gekko Species from Vietnam 147 Table 1: Measurements and proportions of Gekko scientiadventura sp. n. For abbreviations see Material and Methods, Section 2 * = holotype; ** = paratype; *** = live specimen. SVE SLE BL HWrHHN SE EE SVL:TL SVL: HL -HL:HW HL: HH SE : El ZFMK 76174** 69.0 78.7 182 140 80 8.2 6.5 0.88 3.79 1.30 2.28 1.26 ZFMK 76175** 33.0: -41.5. 9:5. 7.4 4.0 4.5 3.0 0.84 3.68 1.28 2.38 1.50 4 ZFMK 76176** 2.0 29:0. 86 7.0 3.8 3.9 2.8 0.93 3.14 1,23 2.26 1.39 ZFMK 76177** 30.0 363 90 70 44 4.3 3:2 0.83 3.33 1.29 2.05 1.34 ZFMK 76178** 29:5; 36:0: 9:0. O 4:1) 83.9 3:2 0.82 3.28 1.29 2.20 1.22 ZFMK 76179** 29.0: 33.0. 84 64.40: 4.0 3.0 0.88 3.45 1.31 2.10 1:33 ZFMK 76198* 60.0 68.0 15.5 12.2 5.0 7.7 5.1 0.88 3.87 1.27 3.10 1.51 ZFMK 76199** 26.5 26.5 80 64 - - ~ 1.00 3.31 1.25 ZFMK 80651** 6650 117.6: 13-4 70.2080 6.5 - 3.78 1.3] 2.51 1.23 ZFMK 80652** 39:8 27,3:07 16.3 129° 7:0" 77.9 5.8 0.80 3.62 1.28 2.36 1.36 male*** 70.0. 15:00: 1.923, 15:0. 8.5 9.0 7.0 0.93 3.59 1.30 2.29 1.29 female*** 68.0 - 180 135 75 85 7.0 — 3.78 1.33 2.40 1.21 male*** 66.0 70.0 19.0 140 80 8.5 7.0 0.94 3.47 1:36 2.38 1.21 male*** 73.0 81.0 20.0 15.0 8.0 9.0 7.0 0.90 3.65 1.33 2.50 1.29 Table 2: Pholidosis of Gekko scientiadventura sp. n. For abbreviations see Material and Methods, Section 2. * = holotype: ** = paratype; *** = live specimen. SPL SBL N IN PM GP IO V SB Er LT4 PP PAT S3R ZFMK 76174** 13/13 18/12: 3/3 0 2 5 41 38 139 12/14 14/14 0 3/3 10 ZFMK 76175** 13/12 9/9 3/3 0 2 6 ~ 48 118 14/12 17/16 0 3/3 10 ZFMK 76176** 14/13 11111 3/3 0 2 6 - 43 13/14 15/17 0 - 10 ZFMK 76177** 14/14 9/9 3/3 0 2 3 = 40 133 14/- 15/- 0 - 11 ZFMK 76178** 13/14 9/9 3/3 0 2 5 - 43 136 13/15 17/- 0 - ZFMK 76179** 12/12 9/9 3/3 0 2 7 - 40 135 15/- 16/16 0 — 10 | ZFMK 76198* 13/12 11/10 38 0 2 6 44 4] 140 14/12 17/15 5 3/3 10 | ZFMK 76199** 14/12 10/9 3/3 0 2 5 - _ = 0 | ZFMK 80651** - 11/11. 3/3 - 2 6 42 43 131 14/14 15/14 0 2/2 11 | ZFMK 80652** 12/13 10/9 3/3 0 2 6 49 42 133: 15/15. 17/16 0 2/2 10 male*** 13 11 3 0 2 5 47 - - 14 14 6 3 female*** 14 12 3 0 2 7 48 - - 14 16 0 3 = male*** 13 11 3 0 2 5 49 - - 14 14 8 3 male*** 14 11 3 0 2 5 5] > - 14 14 6 3 148 Bonner zoologische Beitráge 53 (2004) Table 3: Measurements, proportions and pholidosis of Gekko subpalmatus (holotype BMNH 1946.8.2592, female) and Gekko melli (lectotype ZMB 27659 A, female; paralectotype ZMB 27659 B, juv.). For abbreviations see Material and Methods, Section 2. BMNH 1946.8.2592 ZMB 27659 A ZMB 27659 B SVL 55.0 84.6 29.0 TL 56.0 76.0* 26.5 HL 14.3 22.0 9.0 HW 11.0 15.8 6.6 HH 6.2 9.2 4.0 SE 7.1 9.3 4.0 EE 3.9 8.2 3.1 SVL: TL 0.98 1.11 1.09 SVL: HL 3.85 3.85 3.22 HL: HW 1.30 1.39 1.36 HL: HH 3] 2.39 2.29 SE:EE 1.29 1.13 1.29 SPL 10/10 11/13 11/10 SBL 10/10 11/12 11/9 N 3/3 3/3 3/3 IN | 1 l PM 2 2 2 GP 5 4 10 45 39 35 V 42 43 49 SB 148 147 158 LT] 12/11 11/11 11/10 LT4 10/10 14/12 14/13 PP 0 0 0 PAT 1/1 1/1 1/1 S3R 7 9 9 * 25.0 mm regenerated Bonner zoologische Beiträge Band 53 (2004) Heft 1/2 Analyse der biologisch-ökologischen Ursachen der Evolution der gastroneuralen Metazoa — Testen einer phylogenetischen Hypothese' Klaus Peter SAUER & Harald KULLMANN Institut fiir Evolutionsbiologie und Okologie, Rheinische Friedrich-Wilhelms-Universitát Bonn, Germany Abstract. The goal of our paper is to test the validity of the phylogenetic hypothesis built by AHLRICHS (1995) concern- ing the evolution of the gastroneuralian metazoa. While phylogenies are historical-narrative explanations of the evolu- tionary history of groups of organisms, the evolutionary theory and the mechanisms and processes of the evolutionary change are nomological-deductive explanations of the organismic evolution. Because phylogenies completely depend on the working evolutionary mechanisms and processes, phylogenetic hypotheses must be based on and tested against the theory of evolution. Answering the question “Why are gastroneuralian metazoa?” requires a functional-adaptive (no- mological-deductive) analysis of the key characters evolutionary responsible for building the ecological zone of the gas- troneuralian stem species. Those functional-adaptive analyses are an essential part for testing the validity of phyloge- netic hypotheses. By our analysis of the key characters of the gastroneuralian metazoa the hypothesis of the monophyletic origin of the Gastroneuralia could not be falsified. Key words. Gastroneuralia, Bilateria, phylogeny, evolution, nomological-deductive explanation, historical-narrative ex- Seiten 149-163 | Bonn, Juni 2005 planation, functional-adaptive analysis, key characters 1. PROLOG Wir verfolgen hier das Ziel, die Evolution des Taxon Gastroneuralia (= Nemathelminthes + Spiralia, sensu AHLRICHS 1995) kausal zu erkláren. Der geneigte Leser möge jedoch keine vollständige und abgeschlossene Er- klärung der Evolution der Gastroneuralia erwarten. Der nachfolgende Aufsatz ist vielmehr als Einübung in eine umfangreichere Bearbeitung des Problems zu sehen. Die Kausalanalyse der Evolution höherer Taxa umfasst nach BOCK (1981) zwei wichtige Schritte: 1. die Ent- wicklung einer phylogenetischen Hypothese und 2. die funktional-adaptive Analyse der Schlüsselmerkmale des Taxon, dessen Evolution erklärt werden soll. Vor allem der zweite Schritt — die Merkmalsanalyse — ist von be- sonderer Bedeutung, da diese einen Test der phylogene- tischen Hypothese ermöglicht und die Grundlage für die Abschätzung der Validität der phylogenetischen Hypo- these darstellt. Die in den Phylogenien verschiedener Taxa zum Ausdruck kommenden „Genealogien“ sind Abstraktionen der zurückliegenden Evolutionsgeschich- te. Phylogenien sind daher eine Form von historisch- narrativen Erklärungen (NAGEL 1961; HEMPEL 1965, 1977; BOCK 1981) der Evolutionsgeschichte. Bei der Rekonstruktion einer Phylogenie treffen wir zu- nächst auf ein rational-empirisches Verfahren, das nicht kausalanalytisch einsetzt, sondern vergleichend zur Er- I Clas Michael Naumann zu Königsbrück (26.06.1939 — 15.02.2004) zum Gedenken kenntnis gelangt. Dieser vergleichend formenkundliche Frageansatz beruht auf der abgestuften Gestaltähnlich- keit der Organismen und beinhaltet die Ursachen dieser Gestaltähnlichkeit sowie deren historisches Geworden- sein zunächst nicht. Vielmehr arbeitet der morphologi- sche Frageansatz mit der Vorstellung von der Ver- gleichbarkeit der Gestalten und mit der Möglichkeit, einzelne Arten in Gruppen und diese Gruppen wieder- um in umfassenderen zusammenfassen zu können, wo- bei ein hierarchisch-enkaptisches System entsteht, in dem jede Gruppe durch gemeinsame Gestaltmerkmale gekennzeichnet ist. Die auf diese Weise formulierten Abstammungshypothesen sind, wie alle wissenschaftli- chen Hypothesen, theoriebelastet. Seit Charles DARWIN (1859) seine Theorie der organismischen Evolution in seinem revolutionären Werk On the Origin of Species by Means of Natural Selection veröffentlicht hat, müs- sen phylogenetische Hypothesen auf der Theorie der or- ganismischen Evolution begründet sein. In vielen kladistischen Untersuchungen gehen die Auto- ren dagegen immer noch davon aus, dass phylogeneti- sche Systematisierung und Evolutionsbiologie vonein- ander unabhängig sind. Dieser Ansicht scheint auch AX zu sein (1984, S. 59), wenn er schreibt: „Das Verfahren der phylogenetischen Systematisierung von Organismen ist unabhängig von jeglicher Vorstellung über kausale Mechanismen der Evolution, gleichgültig, ob es sich um das Hypothesengebäude der ‚synthetischen Theorie der Evolution’ ... handelt, um Erweiterungen und Verände- rungen dieser Theorie oder um Alternativen zu ihr...” 150 Bonner zoologische Beitráge 53 (2004) und er fährt fort (AX 1984, S. 136): „Die Ordnung der Produkte der Phylogenese und die Wiedergabe ihrer Verwandtschaftsbeziehungen in einem phylogeneti- schen System ist rational und methodologisch unabhän- gig von der Kenntnis der Mechanismen, welche die evo- lutiven Veränderungen in den Populationen von Arten und den Prozess der Phylogenese steuern.“ Wir befinden uns in scharfem Gegensatz zu dieser Auf- fassung. Wir sind vielmehr davon überzeugt, dass es wenig befriedigend ist, ja gegen die Regeln der Natur- wissenschaft verstößt, eine Abstammungshypothese nur auf dürren Apomorphien — manchmal nur einer einzigen — zu begründen, ohne diese Hypothese an der beobacht- baren Realität und den dem evolutiven Wandel zugrun- de liegenden Mechanismen (= Prozessen) zu testen. Es ist unbestritten, dass man ausschließlich auf der Grundlage des vergleichend formenkundlichen Frageansatzes (der Suche nach Homologien und ihrer Wertung) die genea- logıschen Beziehungen der Organismengruppen unter- einander rekonstruieren kann (s. o.). Heraus kommt ein formales System von Dichotomien, dessen Knotenpunk- te mehr oder weniger gut durch Apomorphien gestützt sind. Die Ermittlung eines solchen Systems ist der Ver- such, die Folge von Freignissen und Änderungen zu be- schreiben und in eine chronologische Ordnung zu brin- gen. Dabei entsteht jedoch ein Baum ohne Blüten und Blätter, ein Skelett ohne Fleisch und Blut. Kladisten, vor allem aber solche, die sich molekularer Methoden bedienen, begnügen sich allzu oft mit diesem Stand der historisch-narrativen Erklärungen (s. u.). Phylogenien sind wissenschaftliche Erklärungen (NA- GEL 1961; HEMPEL 1965, 1977; BOCK & VON WAHLERT 1963; BOCK 1981; SZALAY & BOCK 1991) eines Satzes von Beobachtungen und miissen auch als solche auf der zugrunde liegenden Theorie der organismischen Evolu- tion beruhen und vor allem auch gegen diese getestet werden (BOCK 1981; SZALAY & BOCk 1991). Es ist selbstverstandlich entscheidend fiir eine Phylogenie, wie die Organismen evolviert sind. Eine Phylogenie beruht vollstandig auf den bekannten Mechanismen der Evolu- tion und nicht lediglich auf der Hypothese, dass evoluti- ver Wandel stattgefunden hat. Die Evolutionstheorie und die Mechanismen des evolu- tiven Wandels sind nomologisch-deduktive Erklárungen der organismischen Evolution (BOCK & VON WAHLERT 1963, Bock 1981; eine ausführliche Diskussion der nomologisch-deduktiven Erklárung findet man bei NA- GEL 1961 und HEMPEL 1965, 1977). Mit der von ihm entwickelten Hypothese der Phyloge- nie der Gastroneuralia versucht AHLRICHS (1995) die Evolutionsgeschichte dieser Gruppen zu erkláren. Da phylogenetische Hypothesen auf der Theorie der Evolu- tion beruhen, miissen sie auch gegentiber dieser getestet werden, d.h. nomologisch-deduktive Erklárungen die- nen dazu, historisch-narrative Zu testen (s.u.). Damit ge- langen wir zu einer erklärenden (evolutionären) Phylo- genetik. BOCK & VON WAHLERT (1963) haben dieses Konzept der erklärenden Phylogenenetik entwickelt (siehe dazu auch OSCHE 2002). Sie unterscheiden drei Aspekte evolutionsbiologischer Untersuchungen: 1. Die reine Beschreibung von phyletischen Linien von Orga- nismen und ihre chronologische Ordnung. Eine solche historische Studie kann man als Phylogenetik bezeich- nen. 2. Die Untersuchung der Mechanismen des evolu- tiven Wandels einschließlich ihrer Ursachen und Kon- sequenzen. Diese Analyse der Mechanismen des evolutiven Wandels haben BOCK & VON WAHLERT (1963) als Studium der Evolutionsprinzipien bezeichnet. Dieser Aspekt ist, obgleich mit historischen Gegenstän- den befasst, keine historische Untersuchung. 3. Die möglichst vollständige, auf den Mechanismen der Evo- lution beruhende Erklärung der Ereignisse und des evo- lutiven Wandels in den phyletischen Linien der Orga- nismen. Diesen Aspekt bezeichnen BOCK & VON WAHLERT (1963) als erklärende (evolutionäre) Phylo- genetik. Diese ist eine historische Disziplin. Wenn man die Entstehung neuer Gruppen, wie hier z.B. die der Gastroneuralia erkláren will, dann muss man nach den Ursachen von deren Ursprung suchen; dabei miissen die bekannten Mechanismen der Evolution be- rúcksichtigt werden. Nur wenn eine solche Erklárungs- prozedur vorliegt, ist man berechtigt, eine Untersuchung „Die Evolution der ...“ zu nennen. Wir untersuchen hier die Evolution der Gastroneuralia. 2. DIE EVOLUTION DER GASTRONEURALIA 2.1. Der Ursprung der Gastroneuralia Bei der Frage nach dem Ursprung der Gastroneuralia gilt es zunächst die Frage zu beantworten: „Woher kommen die Gastroneuralia? Diese Frage ist gleichbe- deutend mit der Suche nach der Schwestergruppe der Gastroneuralia und der Stammart dieser beiden Schwestertaxa. Es gilt aber auch die Frage zu klären: „Warum gibt es Gastroneuralia?“ Die Beantwortung dieser Frage ist gleichbedeutend mit dem Versuch, die Selektionswirkungen, welche die Schlüsselmerkmale der Gastroneuralia begünstigt haben, kausal zu erklären. Wir suchen also nach autapomorphen Schlüsselmerk- malen, welche die „ökologische Zone“ (GÜNTHER 1949, 1950) der Gastroneuralia erschließen halfen. Wir müssen daher die adaptive Bedeutung folgender Merk- male begründen (funktional-adaptive Analyse nach Bock 1981): l. eines lediglich Millimeter großen Körperhabitus, 2. eines kurzen und dünnen, wurmförmigen und äußer- lich nicht weiter untergliederten Körpers, Klaus Peter SAUER & Harald KULLMANN: Analyse der biologisch-ókologischen Ursachen der Evolution 15] 3. einer Lebenslaufgeschichte der Individuen, die durch eine direkte Ontogenese und friihe Geschlechtsreife gekennzeichnet ist (r-Strategie), 4. eines dorsofrontalen Cerebralganglions, 5. von mehreren Längsnervensträngen, mit einem Paar ventrolateral gelegener Hauptnervenstränge, 6. Partnerfindung und -erkennung. Diese Merkmale sind allen ursprünglichen Gastroneu- ralia (Gastrotricha [Nemathelminthes], Catenulida, Gnathostomulida [Spiralia] u.a.) gemeinsame Homolo- gien, d.h. diese Merkmale traten bei der Stammart der Gastroneuralia erstmals bei allen Individuen gemeinsam auf (Autapomorphien sensu HENNIG 1982; Abb. | und 2). Zu diesem Ergebnis fiihrt die Morphologie, die ihre Erklárung aus dem formenkundlichen Vergleich ge- winnt. Die Erklárungen, die durch die vergleichende Formenkunde, die Morphologie, gewonnen werden, be- ruhen auf dem Homologieprinzip: „Homolog sihd Strukturen, deren nicht zufällige Übereinstimmung auf gemeinsamer Information beruht“ (OSCHE 1973). In unserem Fall handelt es sich um Information, die nur von den Eltern auf die Kinder übertragen worden sein kann (das gilt in der Regel nur für genetische Informati- on). „So o ® N S lo) 9 ® ® 37 ON I > Oe So : se Ro xe S < x S PS S 9 Q So oe N x Sa $ © © Se E g x $ < S& E > RD © ys < © SS © © S S Plathelminthes Gnathifera Plathelminthomorpha Cycloneuralia Spiralia Nemathelminthes Merkmale 1-5 Gastroneuralia Abb. 1: Verwandtschaftsbeziehungen innerhalb der Gastroneuralia kombiniert nach AHLRICHS (1995) und EHLERS (1985). Zur Klärung der Frage: „Woher kommen die Gastroneu- ralia? * bedienen wir uns der Regeln der vergleichenden Formenkunde. Die auf dem Homologieprinzip beruhende Morphologie führt hier za dem partikulären Allsatz: Alle ursprünglichen Gastroneuralia (= ursprüngliche Ne- mathelminthes + ursprüngliche Spiralia) sind durch die oben genannten Merkmale 1 bis 5 gekennzeichnet. Damit gewinnt in der historischen Evolutionsforschung das Homologieprinzip eine besondere Erklärungsqualität. Das gilt hier aber lediglich für die Erklärung des histori- schen Ursprungs der Gastroneuralia. Die Regeln der Morphologie entsprechen denen der de- duktiven Logik und erlauben Schlussfolgerungen (The- oreme) aus Prämissen (Axiomen) hervorgehen zu las- sen. Die Morphologie ist durch ihre syllogistische Schlussweise ausgezeichnet. Der Syllogismus der tradi- tionellen aristotelischen Logik besteht aus drei Sätzen, wobei zwei die Prämissen, der dritte Satz die Konklusi- on bilden. In unserem Fall können wir aus den beiden Vorsätzen: l. Wenn ein Organismus die Merkmale | bis 5 besitzt, ist er ein ursprünglicher Gastroneuralier, Gastrotricha haben die Merkmale | bis 5, schlie- Ben, dass 3. Gastrotricha ursprüngliche Gastroneuralia sind. [59] Der Mittelbegriff „Merkmale 1 bis 5“ lässt die Gastro- trıcha als ursprüngliche Gastroneuralia erkennen. Die Beantwortung der Frage: „Warum gibt es Gastro- neuralia?“, also die Suche nach den Selektionswirkun- gen, welche die oben genannten Schlüsselmerkmale | bis 5 der ursprünglichen Gastroneuralia begünstigt ha- ben, ist lediglich durch eine historisch-narrative Erklä- rung (NAGEL 1961; HEMPEL 1965, 1977) zugänglich. Bevor wir dieser Erklärung nachgehen, wollen wir zu- nächst die Abstammungshypothese, wie sie AHLRICHS (1995) entwickelt hat, näher betrachten und ein mégli- ches Schwestertaxon ermitteln (Abb. I und 2). Bonner zoologische Beitráge 53 (2004) Sé S x NO 2 XS S S S SN 0 $ > o = > 9 x o N x 0 ros ¡O x < e D x N o? E AS N d O S S & ¥ 2 SS OP & N > e ES N > Q Q 163 ES x 6 > AY o Spiralia Merkmale 1-5 a ae Gastroneuralia Radialia Bilateria Acrosomata Eumetazoa Metazoa Abb. 2: Verwandtschaftsbeziehungen innerhalb der Metazoa verándert nach AX (1995) und AHLRICHS (1995). 2.2. Die Grundmuster der Stammarten der Bilateria, Gastroneuralia und Radialia und ihre genealogischene Verkniipfung Stellt man die Frage nach den Ursachen, die zur Evolu- tion des Grundmusters der Gastroneuralia geführt ha- ben, so ist es zunáchst notwendig, dieses Grundmuster zu beschreiben und genau zwischen abgeleiteten Merk- malen, die in der Stammart der Gastroneuralia erstmals auftreten, und plesiomorphen Merkmalen zu unterschei- den, denn nur abgeleitete Merkmale können uns konkre- te Hinweise auf die speziellen selektiven Bedingungen in der Stammlinie eines Taxon geben. Der Vergleich der grundplannahen Gruppen der Gastroneuralia liefert uns ein eindeutiges Bild: Sowohl bei den Gastrotricha (= Nemathelminthes), als auch bei den Gnathostomulida und Catenulida (= Spiralia) handelt es sich um mikro- skopisch kleine Bewohner des Sandlückensystems (Me- sopsammal) mit einem wurmförmigen Körper. Diese Infauna-Lebensgemeinschaft wird als Mesopsammon bezeichnet. Das Nervensystem besteht aus einem fronta- len Gehirn und Längsnerven, wobei vor allem ein Paar ventrolateraler Hauptnervenstränge auffällt. Weiterhin zeigen alle basalen Gastroneuralia eine direkte Entwick- lung ohne Larvenstadium. Um zu entscheiden, ob es sich bei diesen Merkmalen um evolutive Neuerwerbungen der Gastroneuralia oder um das evolutive Erbe aus dem Grundmuster eines übergeordneten Taxon handelt, soll zunächst das Grund- muster des nächst übergeordneten Taxon, der Bilaterier, untersucht werden. Glaubt man der überwiegenden Mehrzahl der aktuellen Lehrbücher zum Thema Spezielle Zoologie (z.B. AX 1995), so ist die Frage nach dem Grundmuster der Bila- terier heute relativ eindeutig beantwortet. Das dort geschilderte Grundmuster gleicht bis auf ganz wenige Details dem oben geschilderten Grundmuster der Gas- troneuralia. Der letzte gemeinsame Vorfahr aller Bilate- rier war demnach ein kleiner, bilateralsymmetrischer, wurmförmiger Organısmus, der das Benthal vorzeitli- cher Meere bewohnte. Als Konsequenz seiner wurm- förmigen Gestalt hatten sich ein Vorder- und ein Hinter- ende ausgebildet, wobei das Vorderende zum einen durch die Mundöffnung, zum anderen durch eine Kon- zentration von Sinnes- und Nervenzellen („Gehirn“) ge- kennzeichnet ist. Paarige Protonephridien erlaubten dem Urbilaterier eine verbesserte Osmoregulation und somit das Leben in Medien mit wechselnder Salinität. Eine neue Qualität erhält in der Stammlinie der Bilate- rier das Mesoderm, also die Strukturen zwischen Ekto- und Entoderm (BARTOLOMAEUS 1993). Während die Mesogloea der Cnidarier und Ctenophoren noch weit- gehend aus extrazellulärer Matrix mit einzelnen, aus den angrenzenden Epithelien eingewanderten Zellen (z.B. i-Zellen = Stammzellen) besteht, findet sich bei den Bi- lateriern an dieser Stelle eine echte Gewebe formende Zellage, welche unter anderem für die Ausbildung eines Klaus Peter SAUER & Harald KULLMANN: Analyse der biologisch-ókologischen Ursachen der Evolution 153 subepidermalen Hautmuskelschlauches aus (im ein- fachsten Falle) einer äußeren Ring- und einer inneren Längsmuskulatur verantwortlich ist. Auch über den Le- benszyklus scheint Klarheit zu bestehen: Die Stammart der Bilaterier hatte eine direkte Entwicklung ohne pela- gisches Larvenstadium. Unklarheit über das Grundmuster der Bilaterier besteht nach diesem Szenario nur in einigen wenigen Punkten, vor allem in der Frage nach dem Vorhandensein oder Fehlen von After oder primären bzw. sekundären Lei- beshöhlen. Während sich einige dieser Merkmale bei einem Ver- gleich der ranghöchsten Taxa der Bilaterier tatsächlich widerspruchsfrei für die Stammart aller Bilaterier postu- lieren lassen, so gibt es andere Merkmale, die bei ge- nauem Hinsehen unserer Ansicht nach einer kritischen Überprüfung bedürfen. Widerspruchsfrei sind das Vor- handensein von Protonephridien, die neue Qualität des Mesoderms als gewebebildende Zellage und die grund- sätzlich bilateral angelegte Körpersymmetrie. Problema- tisch scheinen uns vor allem die folgenden Punkte: Der Gesamthabitus (klein, wurmförmig, im Interstitium le- bend), die Cephalisation am Vorderende und das Fehlen eines pelagischen Larvenstadiums. Der Vergleich der beiden Gruppen Radialia und Gastro- neuralia (die AHLRICHS 1995 ohne Begründung als Schwestergruppen annimmt; Abb. 2) zeigt auf den ers- ten Blick, dass sich wurmförmige, mikroskopisch kleine Sandlückenbewohner nur innerhalb des Taxon Gastro- neuralia finden lassen. Auch das Fehlen eines pelagi- schen Larvenstadiums finden wir nur bei den grund- plannahen Gruppen der Gastroneuralia. Anders verhält es sich hingegen bei den Radialia. Phoroniden und Deu- terostomia sind makroskopische, sessil oder hemisessil lebende Organismen, deren Körper keinesfalls als wurmförmig beschrieben werden kann. Vielmehr erin- nern diese Gruppen in Körperbau und Lebensweise an die Polypen der Cnidarier, und wie diese ernähren sie sich filtrierend mit Hilfe eines Tentakelapparates. Wei- terhin zeigen Phoroniden und Deuterostomier, wie die Cnidarier, einen pelago-benthischen Lebenszyklus (JÄGERSTEN 1972) mit einer Schwimmlarve als Propa- gationsstadium. Auch eine Cephalisation mit vorne lie- gendem Gehirn ist nicht deutlich ausgeprägt, das Ner- vensystem ist als diffuser Nervenplexus ausgebildet (BULLOCK & HORRIDGE 1965; SAUER & HOCH 2002). Es ist kein zwingender Grund zu erkennen, warum diese Merkmale Neuerfindungen der Radialia sein sollten. Vielmehr scheint es uns plausibel, dass vom Grundmus- ter der Eumetazoa bis hin zum Grundmuster der Deute- rostomia ein makroskopischer, polypenähnlicher Habi- tus, ein pelago-benthischer Lebenszyklus und eine mit Hilfe eines Tentakelapparates filtrierende Lebensweise kontinuierlich beibehalten wurden. Das einzige proble- matische Taxon bei einem solchen Szenario sind die nicht sessilen Ctenophoren (Abb. 2). Zwar sind auch sie makroskopisch, auch sie ernähren sich mit Hilfe von Tentakeln und auch thr Habitus lässt sich, analog der Meduse der Cnidarier, von einem polypenähnlichen Typ ableiten, allerdings besitzen sie eine direkte Entwick- lung ohne Larve. Ihre von dem „normalen“ radiáren Furchungstyp der Cnidaria und Radialia abweichende biradiale Furchung lässt jedoch vermuten, dass die Ent- wicklung der Ctenophoren ein abgeleitetes Eigenmerk- mal der Ctenophoren darstellt und unserer Argumentati- on somit nicht im Wege steht. Durch das hier dargestellte Szenario erhält auch eine al- te und von vielen für überholt angesehene Überlegung in leicht abgewandelter Form neue Aktualität. Im Laufe der Anagenese der Radialia könnten die Coelome, so wie bereits in der Archicoelomaten-Theorie (MASTER- MAN 1898; REMANE 1950; JÄGERSTEN 1955; SIEWING 1969, 1980) vorgeschlagen, aus einer Trennung der hydrostatischen und der verdauenden Funktion des Gastrovaskularsystems, also aus Gastraltaschen hervor- gegangen sein, wie sie bei Cnidariern ausgebildet sind und auch für die Stammart der Eumetazoa hypothetisiert werden. Die Bildung der Coelome aus Entodermzellen bei den Tentaculaten und aus Darmabfaltungen bei Deu- terostomiern kann als Indiz hierfür gewertet werden. Anders verhält es sich bei den Coelomen der Euspira- lier. Dass ihre Coelome nicht homolog zu den Coelo- men innerhalb der Radialer sind, ist weitgehend aner- kannt (z.B. BARTOLOMAEUS 1993). Vielmehr entstehen innerhalb der Euspiralia, vermutlich unabhängig, Coe- lome als Eigenbildungen der Taxa Nemertini, Mollusca und Articulata. Für ein Vorhandensein oder den Verlust von Coelomen im Grundmuster der Gastroneuralia gibt es keinerlei Hinweise. Folgt man den hier ausgeführten Überlegungen, stellt sich das Grundmuster der Bilaterier wie folgt dar: Der Urbilaterier war ein makroskopisch großer, polypenähn- licher Organismus, der sich mit Hilfe eines Tentakel- kranzes, auf Sedimentböden siedelnd, filtrierend ernähr- te. Das bedeutet, es hat in der Stammlinie der Bilateria ein Lebensortwechsel vom Hartsubstrat, wo die ur- sprünglichen Eumetazoa siedelten, auf Sedimentböden stattgefunden. Das Nervensystem ist diffus organisiert (BULLOCK & HORRIDGE 1965: SAUER & HOCH 2002) und der Lebenszyklus beinhaltet ein pelagisches Lar- venstadium. Autapomorphien der Bilaterier sind eine bi- lateral angelegte Körpersymmetrie, Protonephridien und ein Gewebe bildendes Mesoderm, welches u.a. für die Ausbildung eines Hautmuskelschlauches verantwortlich ist. Dieses Grundmuster wird von den Deuterostomiern mit einigen wenigen Abwandlungen, vor allem der Ausbildung von Coelomen, übernommen. 154 Bonner zoologische Beitráge 53 (2004) Starke evolutive Veránderungen finden hingegen in der Stammlinie der Gastroneuralia statt. Ihre auffálligste Autapomorphie ist ein mikroskopisch kleiner, wurm- fórmiger Habitus, verbunden mit einen Wechsel des Le- bensortes in das Sandlúckensystem. Als Folge dieser Veránderungen ist es weiterhin zur Cephalisation und der Ausbildung von Lángsnerven gekommen. Die Ursa- chen und der Verlauf der Evolutionsgeschichte der Bila- teria, Gastroneuralia und Radialia sowie deren genealo- gische Verknüpfung können nur historisch-narrativ er- klart werden. 3. DIE HISTORISCH-NARRATIVE ERKLARUNG 3.1. Charakterisierung des Erklárungstyps Zur Erklárung von Phánomenen des Lebendigen miissen in der Evolutionsbiologie historische Faktoren beriick- sichtigt werden. Aus diesem Grund treffen wir in dieser Disziplin auf den historisch-narrativen, den entwick- lungsgeschichtlichen Frageansatz (NAGEL 1961; HEM- PEL 1965, 1977). Jede phánotypische Gestalt, aber auch jede Form von ökologischer Interaktion der Individuen mit ihrer Umwelt ıst das augenblickliche Endprodukt eines einmaligen historischen Prozesses. Die diesem Prozess zugrunde liegende Kausalitát entzieht sich der unmittelbaren experimentellen Prüfung. Aus diesem Grund ist in diesem Zusammenhang eine andere als die nomologisch-deduktive Erklärungsprozedur erforder- lich, nämlich die historisch-narrative. Dieses Erklä- rungsverfahren beschreibt ein zu erklärendes Merkmal, also auch jeden Typ von ökologischer Interaktion, als letztes Ergebnis einer einmaligen Entwicklungsfolge. Damit wird das betrachtete Phänomen durch Erzählen der aufeinander folgenden Entwicklungsstadien erklärt. Ziel einer historisch-narrativen Erklärung ist es, nach- zuweisen, wie „ein Zustand zum nächsten führt“ (HEMPEL 1965, 1977), wie also aufeinanderfolgende Zustände (Merkmalsausprägungen) durch allgemeine Gesetzmäßigkeiten miteinander verknüpft sind. Damit wird z.B. die Evolution eines jüngeren Zustands eini- germaßen wahrscheinlich gemacht. Im Falle der Beant- wortung der Frage: „Warum gibt es Gastroneuralia?“ gilt es, die Selektionswirkungen zu rekonstruieren, wel- che die Entstehung der oben genannten Merkmale 1 bis 5 begünstigt haben. Historisch-narrative Erklärungen beinhalten also einerseits einen gewissen Anteil nomo- logischer Verknüpfungen, andererseits aber auch einen Anteil an Beschreibung. Dieses von NAGEL (1961) und HEMPEL (1965, 1977) entwickelte Erklärungsverfahren hat einerseits Ähnlichkeit mit dem nomologisch- deduktiven Verfahren und andererseits mit dem rein be- schreibenden Verfahren. In der Praxis wird man diese beiden Verfahren nicht strikt trennen können. Auch wird man die aufeinander folgenden Zustände einer Entwicklungsfolge nicht vollständig auflösen können. Daher wird eine historisch-narrative Erklärung eine Vielzahl von Tatsachen und Ereignissen beschreiben, die sich über einen bestimmten Zeitraum verteilen. Wir wollen hier prüfen, ob die Evolution der abgeleite- ten Grundmustermerkmale der Gastroneuralia einer his- torisch-narrativen Erklärung zugänglich sind. Ist dies der Fall, dann gewinnt die phylogenetische Hypothese von AHLRICHS (1995) an Wahrscheinlichkeit. Um diese phylogenetische Hypothese zu testen, unterziehen wir die Schliisselmerkmale der Gastroneuralia einer funkti- onal-adaptiven Analyse (BOCK 1981). 3.2. Die funktional-adaptive Analyse der Schliisselmerkmale der Gastroneuralia Die Evolutionstheorie und die Mechanismen des evolu- tiven Wandels (z.B. Anderungen von Anpassungen) sind nomologisch-deduktive Erklárungen. Eine Phylo- genie ist dagegen ein Versuch, die evolutionáre Ge- schichte einer bestimmten Gruppe — hier die der Gastro- neuralia — zu rekonstruieren. Jedes dabei genutzte Merkmal ist das augenblickliche Endprodukt eines ein- maligen historischen Prozesses. Die diesem Prozess zugrunde liegende Kausalität entzieht sich, wie bereits festgestellt, der unmittelbaren experimentellen Prüfung. Aus diesem Grund muss die Erklärungsprozedur von den Evolutionsmechanismen zur Phylogenie fortschrei- ten, d.h. nomologisch-deduktive Erklärungen können zum Testen historisch-narrativer Erklärungen herange- zogen werden, niemals aber umgekehrt (BOCK 1981). Im folgenden werden wir die Randbedingungen unter- suchen, die zur Kleinheit, direkten Entwicklung und frü- hen Geschlechtsreife (Merkmale | bis 3) der Stammart der Gastroneuralia geführt haben. Dazu müssen wir die Umweltprobleme kennen, welche die Stammart der Gastroneuralia zu lösen hatte. Wir müssen versuchen, den ursprünglichen Lebensort und die ursprüngliche Lebensweise der Gastroneuralia zu rekonstruieren. 3.3. Rekonstruktion des ursprünglichen Lebensortes der Gastroneuralia Zur Rekonstruktion des ursprünglichen Lebensortes der Stammart der Gastroneuralia müssen wir uns den rezen- ten Lebensort der Individuen der ursprünglichen Grup- pen der Gastroneuralia näher ansehen. Die ursprüngli- chen Gruppen der Gastroneuralia sind die Gastrotricha, Catenulida, Gnathostomulida u.a. (siehe Abb. 1). Die Individuen der Arten dieser Gruppe sind lediglich 0,1 bis | mm lang und leben überwiegend in Sedimenten mit limitierter Sauerstoffverfügbarkeit. Die zu lösenden Umweltprobleme der Individuen dieser interstitiellen Fauna — der Meiofauna — gehen von der Korngröße der Partikel der Sedimente, vor allem aber von der limitier- ten Sauerstoffverfügbarkeit aus. Klaus Peter SAUER & Harald KULLMANN: Analyse der biologisch-ökologischen Ursachen der Evolution 5 In den Sedimenten entwickeln sich anoxische Zonen. FENCHEL & RIEDL (1970) nannten diese anoxischen Zonen und die sie besiedelnden Organismen das „Sul- fid-Biom“. Dieses „Sulfid-Biom“ wird vor allem durch die urspriinglichen Gastroneuralia gebildet. BOADEN & PLATT (1971) haben diese Lebensgemeinschaft das „Thiobios“ genannt, ein Begriff, der allgemein ange- nommen worden zu sein scheint. Wir haben hier nicht den Raum, um auf die umfangrei- che Diskussion über die ursprüngliche Anaerobiose ver- schiedener Meiofauna-Arten bzw. der Metazoa über- haupt einzugehen (z.B. BRYANT 1991; FENCHEL & FINLAY 1995). Wir stimmen jedoch mit REISE & AX (1979) überein und weisen die Hypothesen, dass die Metazoen während des Präkambriums in anoxoischer Umwelt evolviert sein sollen, bzw. dass sich unter den ursprünglichen Gastroneuralia primär anaerobe Meio- fauna-Arten befinden, entschieden zurück. Dieses Prob- lem berührt unsere Überlegungen auch in keiner Weise (siehe unten). Das sogenannte „Sulfid-Biom“ hat eine scharfe Ober- grenze, wo das Redox-Potential mit der Sedimenttiefe rapide von positiven zu negativen Werten übergeht. Diese Grenze wird als Redox Diskontinuitäts-Zone (RDZ) bezeichnet. Unterhalb dieser Grenze werden die Bedingungen anoxisch. Wenn das Redox-Potential ne- gative Werte annimmt, so geht man davon aus, dass im allgemeinen kein O, mehr vorhanden ist. Im sauerstoff- armen Bereich dieser Zone leben allerdings zahlreiche Vertreter der ursprünglichen Gastroneuralia (FENCHEL & RIEDL 1970; HUMMON 1972; LASSERRE & RENAUD- MORNANT 1973; OTT & SCHIEMER 1973, WIESER et al, 1974; BOADEN 1975, 1977; HOGUE 1978; BOADEN & ELHAG 1984). Ein solch harscher Lebensraum stellt seine Bewohner vor schwierige ökologische Probleme, vor allem durch die limitierte O>-Verfúgbarkeit. Welches ist das präa- daptive Plateau, das die Nutzung dieses an organischer Substanz und an „ungenutzten“ Ressourcen reichen, je- doch starken Sauerstoffschwankungen unterworfenen Lebensraumes möglich gemacht hat? Wie mag eine erb- liche phänotypische Varianz in der Effizienz der Nut- zung von O», bei geringem O>-Partialdruck [pO>] ent- standen sein? Um diese Fragen zu beantworten, müssen wir die Aus- bildung und Entwicklung der Atmosphäre der Erde ge- nauer ansehen (KASTING et al. 1992). Die Zunahme des O,-Partialdrucks der Atmosphäre der Erde wurde in den letzten 50 Jahren intensiv diskutiert (NICHOLAS 1991, FENCHEL & FINLAY 1995). Bei welchem pO» in der Ge- nese der Atmosphäre der Erde haben sich eine aerobe Biochemie und erste aerobe Organismen eingestellt? Unter welchem pO, sind die Stammarten der Metazoa, Eumetazoa, Bilateria, Gastroneuralia und Radialia ent- standen”? Die rezente O,-Konzentration beträgt | PAI O), = 20.9% = 0.2 Atm [PAL 0, O»-level]. Vor 1.5 Milliarden Jahren hatte sich eine Oo- Konzentration von 0.01 PAL O, = 0.2% =2 x 10” Atm eingestellt. Dieser PO>-Wert entspricht dem sogenann- ten Pasteur-Punkt, das ist die Oo-Konzentration, bei der amphiaerobe Organismen von aeroben auf anaeroben Stoffwechsel umschalten. Vor I Milliarde Jahren betrug der pO,-Wert etwa 0.1 PAL O, = 2 % (KASTING et al. 1992). present atmosphere Es ist nicht unrealistisch, die Evolution der Bilateria vor einer Milliarde Jahren anzusetzen (BENTON & AYALA 2003). Damit hat die noch früher anzusetzende Evoluti- on der friihen Metazoa wie die der Bilateria bei gerin- gerer Oo-Verfiigharkeit stattgefunden. Diese geringe O>-Verfügbarkeit ist ein Umweltproblem, auf das es verschiedene organismische (adaptive) Ant- worten gibt. „Ziel“ eines aeroben Organismus muss es sein, den limitiert verfügbaren Sauerstoff an die Er- folgsorgane zu bringen. Dieses „Ziel“ kann grundsátz- lich auf mehreren Wegen erreicht werden: 1. dadurch, dass die Diffusionsstrecke des Sauerstoffs durch diinne, einschichtige Epithelien, die allseits von sauerstoffhalti- gem Meerwasser umgeben sind, so gering wie móglich gehalten wird (z.B. bei den Cnidaria, die auch in ihrem Gastrovaskularsystem Meerwasser enthalten; Abb. 2), 2. durch Kleinheit des Organismus (z.B. ursprüngliche Gastroneuralia; Abb. 1) oder 3. durch die Entwicklung eines O>-Transportsystems, eines Kreislaufsystems, bel größeren Bilateria mit deutlich entwickeltem Haut- Muskel-Schlauch (z.B. bei den Phoronida, den altertüm- lichsten rezenten Radialia; Abb. 2). Den Zusammen- hang zwischen pO», Respiration durch Diffusion des O» durch Gewebe und die Größen- bzw. Gewebsdickebe- grenzung bei Organismen können wir quantifizieren und nomologisch-deduktiv erkláren. Dazu bedienen wir uns einer Formel, die HILL (1929) entwickelt hat, welche den Radius eines Zylinders bestimmt, der O>-Diffusion bis in das Zentrum erlaubt. In unserer Betrachtung set- zen wir dabei voraus, dass ein möglicher zylinderförmi- ger Organismus, wie z.B. ein ursprünglicher Gastroneu- ralier a) eine konstante Stoffwechselrate besitzt und b) der pO, konstant ist. Diese Formel lautet Akxy, nen a wobei I, = Radius k= Diffusionskoeffizient von O» ins Gewebe (n. ATKINSON 8.4 x 10” em /atm/h) Yo = O>-Partialdruck (atm) a= Stoffwechselrate in cm’ Osem’ Gewebe'h (konst.) 156 Bonner zoologische Beitráge 53 (2004) Die Stoffwechselrate ist aber ihrerseits größenabhängig. Diese Beziehung folgt nach NICHOLAS (1984, 1991) folgender Formel Y -=a(b=1) X Y= O>-Verbrauch X= Gewicht a und b = Konstanten ATKINSON (1976, 1980) nutzte publizierte O>- Verbrauchswerte (Y) von freilebenden Nematoden und setzte b = 0.79 um die Rate a des O>-Verbrauchs in Ab- hángigkeit von der Körpergröße zu bestimmen. Diese Betrachtung führt zu dem Ergebnis, dass ein Meio- benthos-Organismus mit einem Radius von 25 um etwa 0.6 nl 0>/mg Feuchtgewicht/h nutzt (NICHOLAS 1984, 1991 ). Das wiederum bedeutet, dass die aerobe Respi- ration eines Organismus mit einem Radius von 50 um oder weniger solange nicht O>-limitiert ist, wie der O>- Partialdruck 15 mm Hg (= 10% = 0.1 PALO;) der atmo- sphärischen Luft nicht unterschreitet (vor 0.5 bis 1.0 Milliarden Jahren; NICHOLAS 1991). Durch unvorhersagbare O>-Schwankungen war der Le- bensort der ursprünglichen Gastroneuralia sehr instabil. Das führte dazu, dass es in den Populationen der diesen Lebensort bewohnenden Arten zu dichteunabhängigen und unvorhersagbaren Mortalitátsepisoden kam. Das hatte einen tiefgreifenden Einfluss auf die Lebenslauf- geschichte der Individuen der Arten, die einen solchen Lebensort nutzten. Solche unvorhersagbaren und dich- teunabhángigen Mortalitátsepisoden führen zu r-selek- tierten Merkmalsgefügen (PIANKA 1970; SOUTHWOOD 1980): kleine, kurzlebige Organismen mit früher Ge- schlechtsreife und schneller Entwicklung sowie kurzer Generationszeit. Sogenannte r-Selektion ist dann zu er- warten, wenn starke, regelmäßige, aber unvorhersagbare Umweltschwankungen regelmäßige Mortalitätsereignis- se nach sich ziehen, aber gleichzeitig mit einer hohen Nahrungsverfügbarkeit verknüpft sind (PIANKA 1970; SOUTHWOOD 1980). Solche Umweltbedingungen be- günstigen eine hohe Fekundität, frühe Geschlechtsreife und schnelle Entwicklung eines kleinen Organismus. Mit der r-Selektion ist auch Progenese verbunden (GOULD 1977). Progenese ist die in der Ontogenese zeitlich vorverlegte Reproduktion. In den Populationen der Stammlinien der Acrosomata, Bilateria und Gastroneuralia (Abb. 2) muss eine erbliche phánotypische Variabilitát in bezug auf den pelago- benthischen Lebenszyklus (JÄGERSTEN 1972) der In- dividuen der Populationen dieser Stammlinien existiert haben. Zunächst muss es eine erbliche phänotypische Variabilität der Habitat-Selektion der Larven der Indivi- duen der Populationen der Stammlinie der Acrosomata gegeben haben. Während die Individuen der Populatio- nen der Stammlinie der Ctenophora zu einem rein pela- gischen Lebenszyklus übergegangen sind, haben die In- dividuen der Populationen der Stammlinie der Bilateria und Radialia (Abb. 2) den ursprünglichen pelago- benthischen Lebenszyklus beibehalten, was man daran erkennen kann, dass die Phoroniden als ursprüngliche Radialia einen pelago-benthischen Lebenszyklus mit ei- nem sessilen Adultus und einer der Propagation dienen- den mobilen Larve haben. Die Individuen der Populati- onen der Stammlinie der Gastroneuralia sind dagegen zu einem rein benthischen Lebenszyklus übergegangen. Der Übergang zu einem rein benthischen Lebenszyklus mit direkter Entwicklung muß durch eine erbliche phä- notypische Variabilität zwischen den Individuen der Populationen der Stammlinie der Gastroneuralia in be- zug auf den lebenslaufgeschichtlichen Zeitpunkt der Geschlechtsreife und Reproduktion möglich geworden sein. In diesen Populationen — so hypothetisieren wir — gab es in bezug auf die Lebenslaufgeschichte zwei Phä- notypen: zum einen Individuen, die während ıhrer Le- benslaufgeschichte zweimal geschlechtsreif wurden, einmal in der Juvenilphase und dann noch einmal als sogenannte Adulti; dieses Phänomen der Dissogonie ist z.B. von Ctenophora bekannt. Der zweite Phänotyp wurde durch Progenese (GOULD 1977) nur noch einmal in einem frühen Ontogenesestadium geschlechtsreif. Diese Individuen der Populationen der Stammlinie der Gastroneuralia, die sich unter den r-selektierenden Randbedingungen schließlich nur noch einmal zu einem sehr frühen Ontogenesezeitpunkt reproduziert haben, müssen gegenüber den Individuen mit Dissogonie den nicht zufällig höheren Reproduktionserfolg gehabt ha- ben, also selektionsbegünstigt gewesen sein. In diesem Zusammenhang ist es wichtig zu berücksich- tigen, dass alle Arten der ursprünglichen Gastroneuralia durch eine direkte Entwicklung ausgezeichnet sind. Wenn Larven bei den Gastroneuralia auftreten (z.B. die Müllersche oder die Goettesche Larve bei den Polycla- dida), so sind diese nicht den primären Larven aus dem pelago-benthischen Lebenszyklus der Metazoa, Eume- tazoa, Bilateria und Radialia homolog, sondern sind se- kundáre Larven (JÄGERSTEN 1972). Vielmehr spricht einiges dafür, dass die Larven des primär pelago- benthischen Ontogeneseverlaufs in der Stammlinie der Gastroneuralia geschlechtsreif geworden sind (Progene- se) und damit gewissermaßen ein neuer, sich direkt oh- ne Larve entwickelter Adultus entstanden ist. Die Synthese all dieser beobachtbaren Evidenz sowie der hinzugefügten, für die Erklärung der Evolution der Gastroneuralia wichtigen Überlegungen, erlaubt den Schluss, dass die geringe Größe, direkte Entwicklung und der ontogenetisch frühe Fortpflanzungszeitpunkt der ursprünglichen Gastroneuralia ursächlich durch die gravierenden pO,-Schwankungen in den an nutzbarer Klaus Peter SAUER & Harald KULLMANN: Analyse der biologisch-Gkologischen Ursachen der Evolution 5 organischer Substanz reichen Sedimenten durch r-Sc- lektion erzwungen wurde. Die Evolution der Gastroneu- ralia ist demnach mit einem Wechsel im Lebensort und in der Lebensweise verbunden. 3.4. Lebensort- und Lebensweisewechsel in der Evolution der Gastroneuralia Um die Evolutionsgeschichte von Art-Taxa zu verste- hen und zu erklären, muss man ihre Ökologie untersu- chen, die das Ergebnis der wirksamen Evolutionspro- zesse ist: Während des historischen Ablaufs, den man „Makroevolution“ nennt, d.h. im Verlauf von Anagene- se (adaptiver phyletischer Wandel) und Cladogenese (adaptive Radiation) evolvieren vollständig neue Um- weltbezüge, neue ökologische Nischen. Nach GÜNTHER (1949, 1950), HUTCHINSON (1957, 1959) und BOCK (1979) versteht man unter der ökologischen Nische ei- ner Art zwei Dimensionssysteme, die organismischen (autozoischen) Dimensionen, die in einem definierten biotischen und abiotischen Umweltausschnitt (dieser bildet die ökischen Dimensionen) „eine Rolle spielen“. Der Deckungsbereich beider Dimensionssysteme wird von GÜNTHER (1950) als ökologische Nische definiert. Die Güte der Passung der Individuen der Populationen einer Art in die für sie spezifischen Umweltbedingun- gen, ihre Adaptiertheit (Adaptivness), entscheidet über ihre Fitness. Während intraspezifische Konkurrenz zur Nischenerweiterung führen kann, verursacht interspezi- fische Konkurrenz Nischendifferenzierung. Die Bildung einer Nische setzt 1. ungenutzte, in der Regel nur be- grenzt verfügbare Ressourcen als „ökologische Lizen- zen“ (GÜNTHER 1950) und 2. Merkmale als „organis- mische Lizenzen“ (OSCHE 1983) voraus, welche die Nutzung der ökologischen Lizenzen ermöglichen. Dabei spielen bereits vorhandene, in einem alten Umweltbe- zug entstandene Anpassungen, Praeadaptationen, eine Rolle, die in dem neuen Nischenbezug eingesetzt wer- den, was zu neuen Selektionsszenarien führt. Dabei sind vielfach Verhaltensänderungen „Schrittmacher der Evo- lution“ (MAYR 1970). Während der Anagenese und Cladogenese bilden „Schrittmacherarten“ „Schlüssel- merkmale“ (organismische Strukturen) aus, die einen Wechsel im Umweltbezug herbeiführen können. Durch einen solchen Wechsel im Umweltbezug können ókolo- gische Nischen entstehen, die wir ex post faktum als neue ökologische Zonen (GÜNTHER 1950; SIMPSON 1953) erkennen können. Ökologische Zonen erlauben die adaptive Radiation der „Schrittmacherart“. Dabei wird die ökologische Zone in zahlreiche ökologische Nischen „aufgeteilt“. Diesen Vorgang nennt man kurz gesagt Makroevolution; das ist 1. der Ursprung und die langfristig umfassende Änderung von Anpassungen und damit verbunden die Entstehung neuer, gegenüber den alten bemerkenswert unterschiedlicher Umweltbezüge sowie 2. den Ursprung und die adaptive Radiation eines Taxon von höherem kategorialem Rang. Der historische Übergang vom Grundmuster der Eume- tazoa zu dem der Bilateria und schließlich zum Grund- muster der Gastroneuralia ist eine solche makroevoluti- onäre Ereigniskette. Während dieses Prozesses, der vollständig auf die Mechanismen reduziert kann, die für mikroevolutive Veränderungen ursächlidh sind (RENSCH 1972), kommt es oft zu einem tiefgrei- werden fenden Lebensort- und Lebensweisewechsel. Lebensort-Typen umfassen nach RIEDL (1966) „Reprä- sentanten geschlossener (und aus Anpassungsreihen gleichen Ursprungs hervorgehender) Verwandtschafts- gruppen, welche jeweils weitgehend einheitlich und zumeist nur mehr historisch zu verstehende Bindungen der Gruppenmerkmale an das gemeinsame Biotop auf- weisen“. Der ursprüngliche Lebensort der Eumetazoa ist das Hartsubstrat im Litoral urzeitlicher Meere. Durch eine Verhaltensänderung der Larve im pelago-ben- thischen Lebenszyklus der Individuen der Populationen in der Stammlinie der Bilateria hat spätestens seit der Evolution der Schrittmacherart der Bilateria ein Le- bensortwechsel stattgefunden. Die Larven dieser Schrittmacherart änderten ihre Habitat-Selektion und siedelten auf Weichböden, die durch eine hohe Verfüg- barkeit ungenutzter Ressourcen, aber auch durch eine geringe O,-Verfügbarkeit (s.o.) ausgezeichnet waren und sind. Der Lebensort der Phoroniden und die Habi- tat-Selektion der Actinotrocha-Larve (HERRMANN 1976, 1979) sind dafür Indizien. Während in der Stammlinie der Radialia der Lebensort-Typ beibehalten wurde, ist in den Populationen der Stammlinie der Gastroneuralia ein weiterer neuer Lebensort-Typ entstanden, der im Sedi- ment lebende Urgastroneuralier, der durch die Merkma- le 1 bis 5 gekennzeichnet war. Mit dem Lebensortwechsel vom Hart- auf den Weich- boden war zunächst kein Lebensweisewechsel verbun- den. Der Urbilaterier muss ein sessiler Filtrierer von makroskopischer Dimension (RIEGER et al. 1991) und polypenähnlichem Habitus gewesen sein. Ob schon bei diesem Urbilaterier die Trennung der hydrostatischen Funktion (Coelom) von der verdauenden Funktion (Darm) des Gastrovaskularsystems der Eumetazoa vor- gelegen hat, wie es bei den Radialia der Fall ist, bleibt offen. Vielleicht können wir einmal dem „/gnorabi- mus * ein mutiges „wir werden wissen“ entgegensetzen. In der Stammlinie der Gastroneuralia hat der Lebens- ortwechsel des Urgastroneuraliers auch zu einem Le- bensweisewechsel geführt. Wieder müssen wir davon ausgehen, dass eine Verhaltensänderung die Schrittma- cherfunktion bei der Erschließung des ursprünglichen Lebensortes der Gastroneuralier hatte. Die Larven im noch pelago-benthischen Lebenszyklus der Individuen der Populationen der Stammlinie der Gastroneuralia be- siedelten das Interstitial mariner Weichböden, wo — wie oben bereits dargestellt — diese Larven durch den Pro- 158 Bonner zoologische Beitráge 53 (2004) zess der Progenese zu frühreifen, frei beweglichen Adulti (GOULD 1977) wurden. Die Adaptation an gerin- ge O>-Verfúgbarkeit sowie an dichteunabhängige und unverhersagbare Mortalitátsepisoden reichte jedoch noch nicht aus, um diesen „unwirtlichen“, aber unge- nutzten Lebensraum zu nutzen. Die Schliisselart der Gastroneuralia muss die mikrophage Lebensweise ihrer progenen Larve als Adultus beibehalten haben. Dafür spricht, dass alle ursprünglichen Gastroneuralia mikrophag sind; z.B. ernáhren sich die Catenulida von Bakterien, Flagellaten und Diatomeen (KAESTNER 1964, S. 230) und die Gastrotricha von Bakterien, Flagellaten, Fora- miniferen und Diatomeen (KAESTNER 1964, S. 272). Da wir auch bei den Planulae der Anthozoa, den grund- planáhnlichsten Eumetazoa, eine Mundóffnung kennen, diirfen wir annehmen, dass auch die Larve der Stammart der Eumetazoa eine solche hatte. Bei diesem Typ Planu- la-Larve handelt es sich grundsátzlich um eine Gastrula mit Gastralraum, deren Blastoporus während der Meta- morphose als Mundöffnung persistiert (SIEWING 1969). Diese planktotrophen Larven sind durch ein funktions- tüchtiges entoblastemales Epithel ausgezeichnet; mit Hilfe der Cilien des Peristoms strudeln diese Larven Nano- und Mikroplankton in ihre Mundöffnung (NY- HOLM 1943). Damit ist naheliegend, dass die Schlüssel- art der Gastroneuralia an die mikrophage Lebensweise práadaptiert war. Zur Lebensweise gehórt auch der Fortpflanzungsmodus. Der Fortpflanzungsmodus der Cnidarier, Ctenophoren, Phoroniden, Echinodermen und Ascidien ist die äußere Besamung. Das spricht sehr stark dafiir, dass die Indivi- duen der Populationen der Stammlinie der Eumetazoa- Bilateria-Radialia-Deuterostomia-Chordata durch den ursprünglichen Fortpflanzungsmodus der äußeren Be- samung gekennzeichnet waren. Der Begriff äußere Be- samung beschreibt im hier besprochenen Zusammen- hang die ungezielte Abgabe von Keimzellen ins freie Wasser. Dabei findet die Syngamie sozusagen „fernab“ der Elterntiere statt. Organismen mit solchermaßen äu- Berer Besamung sind gezwungen, die Verbindung zum freien Wasserkörper zu erhalten. Mit dem Fortpflan- zungsmodus der äußeren Besamung ist daher ein Le- bensortwechsel, wie er in der Stammlinie der Gastro- neuralia stattgefunden hat, nicht zu vollziehen. Es ist leicht einzusehen, dass die ungezielte Abgabe von Eı- zellen und Spermien in die Umgebung bei Bewohnern des Interstitials aufgrund stark reduzierter Begegnungs- wahrscheinlichkeit der Keimzellen den Reproduktions- erfolg von Individuen mit äußerer Besamung an diesem Lebensort stark mindern würde. Um den Lebensort- wechsel in das Interstitial mariner Weichböden vollzie- hen zu kónnen, mussten die Individuen der Populatio- nen der Stammlinie der Gastroneuralia zunáchst einen an diesen Lebensort angepassten Fortpflanzungsmodus entwickeln. 3.5. Lebensort-Typ und Fortpflanzungsmodus Unter den Randbedingungen des von den ursprúngli- chen Gastroneuralia genutzten Lebensortes sind solche Individuen selektionsbegünstigt, die ihre Spermien möglichst nahe an die Eier des Paarungspartners heran- bringen. Die anagenetisch „vollkommenste“ Lösung dieses Problems ist die auf eine Kopulation folgende in- nere Besamung. Dieser Begriff der inneren Besamung beschreibt jedoch keinen einheitlichen, im Tierreich homologen Merkmalskomplex. Vielmehr ist der Kom- plex „Kopulation mit nachfolgender innerer Fertilisati- on“ bei den Metazoa vielfach unabhängig entwickelt worden. Und so kann der Merkmalskomplex „innere Besamung‘“ auch keine Autapomorphie der Gastroneu- ralia sein. Während die ursprünglichen Gruppen der Euspiralia noch den ursprünglichen Fortpflanzungsmo- dus der äußeren Besamung aufweisen, sind die ur- sprünglichen Gruppen der Nemathelminthes und Spira- lia (sensu AHLRICHS 1975; Abb. 1) durch eine innere bzw. quasi innere Besamung ausgezeichnet. Der anagenetische Schritt zur inneren Besamung macht eine Kopulation der Geschlechtspartner notwendig. Eine „echte“ Kopulation, d.h. eine Vereinigung der Genital- armaturen und nachfolgende Injektion von Spermien in den weiblichen Genitalsitus wurde aber erst möglich, nachdem bereits komplizierte Partnerfindungs-, Partner- erkennungs- und Paarungsmechanismen entwickelt worden waren. Die Merkmalskomplexe Partnerfindung und -erkennung sind mögliche Autapomorphien der Gastroneuralia. Die Validität dieser Hypothese gilt es in Zukunft zu prüfen. Die innere Besamung wird bei den ursprünglichen Ver- tretern der Gastroneuralia, nachdem die Individuen der Populationen der Stammlinie Partnerfindungs- und -erkennungsmechanismen entwickelt hatten, auf kon- vergentem Wege durch „parallele evolutive Vervoll- kommnung“ (REGENFUSS 1975) erreicht. Die Art und Weise, wie bei den Macrodasyiden, ursprünglichen Gastrotrichen (REMANE 1936), die Spermien übertragen werden, ist sehr vielfältig. Das spricht für die konver- gente Entwicklung der inneren Besamung auch in dieser Gruppe. Betrachten wir ein Beispiel etwas näher. Bei dem protantrischen Zwitter (die ursprünglichen Gastro- neuralia haben den Hermaphroditismus aus der Stamm- linie Metazoa-Eumetazoa-Bilateria beibehalten; SAUER 1996) Dactylopodola baltica wird zunächst bei einem männlich gestimmten Individuum an der rechten Kör- perseite eine Ausstülpung, die Spermatophore, gebildet, in welche die Spermien einwandern. Diese Sperma- tophore wird auf ein Individuum, welches sich in der weiblichen Phase befindet, übertragen (TEUCHERT 1968). Hat ein männlich gestimmtes Individuum dieser Art ein weiblich gestimmtes gefunden und erkannt, dann heftet es die Spermatophore genau an der Stelle Klaus Peter SAUER & Harald KULLMANN: Analyse der biologisch-ókologischen Ursachen der Evolution 159 an, wo das weiblich gestimmte Individuum in seiner männlichen Phase seine Spermien gebildet hatte. Die Spermien dringen dann an dieser Stelle in den Körper ein und wandern zu den Eiern (TEUCHERT 1968). Für unsere Überlegungen ist die Tatsache, dass bei Gastrotrichen die Eier nach der Fertilisation durch Rup- tur der Körperwand abgegeben werden (TEUCHERT 1968) besonders interessant. Das erinnert an den Modus der Abgabe von Keimzellen bei Cnidaria und Ctenopho- ra. Dieser Befund spricht sehr stark dafür, dass die Gastrotrichen neben der Entwicklung neuer auch ur- sprüngliche Merkmale des Fortpflanzungsverhaltens aus den Grundmustern der Eumetazoa-Bilateria beibehalten haben. Auch bei den Catenuliden werden die Eier nach der Besamung durch Ruptur der Körperwand entlassen (KARLING 1974; EHLERS 1985), d.h. auch die ursprüng- lichen Spiralia haben — wie die ursprünglichen Ne- mathelminthen — alte Merkmale des Fortpflanzungsver- haltens aus den Grundmustern der Eumetazoa-Bilateria beibehalten. Bei den ursprünglichen Nemathelminthes und Spiralia umhüllen keine speziellen Wandzellen die Keimlager (HENDELBERG 1983; EHLERS 1985), was an das Grundmuster der Eumetazoa erinnert. Sowohl in- nerhalb der Nemathelminthen, vor allem aber bei den Spiraliern, wird die innere Besamung, verbunden mit immer komplexer werdenden Genitalsitus, erreicht. Den ursprünglichen Gastroneuralia muss es trotz der Bei- behaltung ursprünglicher Merkmale des Fortpflanzungs- verhaltens aus dem Grundmuster der Eumetazoa schon sehr früh auf verschiedenen Wegen gelungen sein, die innere Besamung zu erreichen. Das bedeutet, dass die In- dividuen der Populationen der Stammlinie der Gastroneu- ralia schon sehr früh Partnerfindungs- und Partnererken- nungsmechanismen entwickelt haben müssen. Das ermöglichte ihnen eine „quasi innere“ Besamung ihrer Eier; sie gaben ihre Keimzellen wahrscheinlich in unmit- telbarem Körperkontakt ins freie Wasser ab. Dieses Ver- halten könnte die Praeadaptation zur Nutzung des In- terstitials mariner Weichböden, dem Lebensort der ursprünglichen Gastroneuralia, gewesen sein. Die Evolution der Merkmale 4 und 5 der ursprünglichen Gastroneuralia (Abb. 2), das gegenüber den Cnidariern und Phoroniden viel komplexere Nervensystem, steht wahrscheinlich mit dem Erwerb von Partnerfindungs- und Partnererkennungsmechanismen in ursächlichem Zusammenhang. Solche Mechanismen setzen eine ent- sprechende Sensorik und neurale Informationsverarbei- tung voraus. 3.6. Konsequenzen der Änderung des Fortpflanzungsmodus der Gastroneuralia für ihre adaptive Radiation Innere Fertilisation erlaubt auch Spermienspeicherung und macht wiederholte Paarungen möglich, ehe die ers- ten Eier besamt und abgelegt werden. Das führt zwangs- läufig zur Konkurrenz der Spermien verschiedener Spermienerzeuger um die begrenzt verfügbaren und be- sambaren Eier (PARKER 1970), was wiederum Wirkun- gen der sexuellen Komponente der Selektion verursacht. J Bevor wir auf die bis heute stark unterschätzte Bedeu- tung der sexuellen Komponente der Selektion für die Steigerung der Geschwindigkeit der ókologisch-adap- tiven Differenzierung (SAUER 1996) während der adap- tiven Radiation einer Gruppe zu sprechen kommen, müssen wir kurz auf die Verknüpfung der Inhalte der Konzepte Selektion, natürliche Selektion und sexuelle Selektion eingehen. Generell ist Selektion nicht zufällig differentielle Zygotenbildung bzw. Reproduktion ver- schiedener Individuen einer Population. Dabei bezieht sich der Ausdruck „nicht zufällig“ auf die selektions- verursachende phänotypische Varianz und ihre Heritabi- lität. FISHERS (1930) fundamentales Theorem der Selek- tion besagt, dass die Änderungsrate eines Merkmals zu jeder Zeit der diesem Merkmal zugrunde liegenden ad- ditiven genetischen Varianz proportional ist. Worin liegt der heuristische Wert einer weiteren Differenzierung der generellen Wirkung der Selektion in eine natürliche und eine sexuelle Wirkungskomponente? DARWIN (1859) hat das Konzept der natürlichen Selektion entwickelt, um mechanistisch zu erklären, wie sich Merkmale in Popu- lationen über Generationen hinweg graduell verändern. Dabei hat er nicht immer strikt zwischen Überlebens- und Fortpflanzungserfolg unterschieden. Ihm war je- doch als erstem klar geworden, dass die unterschiedli- che erbliche phänotypische Ausprägung von Merkmalen ihre Träger zu unterschiedlich guter Auseinanderset- zung mit ihrer Umwelt befähigt (Kampf ums Dasein), was letztlich dazu führt, dass in einer Population ..bes- ser“ angepasste Phänotypen häufiger, „schlechter“ an- gepasste hingegen seltener wurden. Dieses Ergebnis un- terschiedlichen individuellen Erfolgs hat DARWIN (1859) „natürliche Auslese“ genannt. Das Konzept der „sexuellen Selektion“ hat DARWIN (1859, 1871) entwi- ckelt, da gewisse Besonderheiten in der Fortpflanzungs- biologie der Tiere sich einer mechanistischen Erklärung durch sein Konzept der natürlichen Selektion (1859) entzogen. Mit dem Konzept der sexuellen Selektion suchte DARWIN (1859, 1871) dem Dilemma zu ent- kommen, dass die meist männlichen Träger von extra- vaganten Strukturen durch die Wirkung der natürlichen Selektion Erfolgsminderungen zu erwarten haben. Einen Ausweg sah Darwin darin, dass die Träger solcher ex- travaganten Merkmale, wie z.B. das Schwanzgetieder des Pfaus, durch eine höhere Attraktivität für die Weib- chen einen höheren Fortpflanzungserfolg erzielen soll- ten, um dadurch wiederum den durch die Wirkung der natürlichen Selektion erzwungenen Nachteil zu kom- pensieren (DARWIN 1872, S. 64): „Diese Form der Se- lektion wird nicht durch den Kampf ums Dasein in be- 160 Bonner zoologische Beitrage 53 (2004) zug auf andere Lebewesen oder äußere Bedingungen verursacht, sondern durch die Fortpflanzungskonkur- renz zwischen den Individuen des einen Geschlechts, im allgemeinen des männlichen, um den Besitz des anderen Geschlechtes“ (übersetzt von den Verfassern). Diese nicht zufällig differentielle Konkurrenzüberlegenheit beim Partnergewinn hat DARWIN (1871, 1872) „sexuelle Selektion“ genannt. Die Theorie der Lebenslaufgeschichte besagt, dass sich Individuen zu jedem Zeitpunkt ihres Lebenslaufes so verhalten müssen, dass sie am Ende eine maximal mög- liche Zahl von Nachkommen erzielen. Auf diesem Weg müssen die Individuen verschiedene ökologische Prob- leme lösen, d.h. Selektionsfilter passieren. Alle bioti- schen und abiotischen Widerstände der Umwelt, welche nicht zufällig differentielles Überleben und/oder nicht zufällig differentiellen Ressourcengewinn bewirken, sind Faktoren, welche natürliche Selektion verursachen. Haben die Individuen schließlich den Reproduktions- zeitpunkt erreicht, wird durch nicht zufällig differentiel- len Paarungspartnergewinn die sexuelle Komponente der Selektion wirksam. Die nicht zufällig differentielle Reproduktion bzw. Zygotenbildung (Selektion) ergibt sich also aus dem nicht zufällig differentiellen Überle- ben und dem nicht zufällig differentiellen Ressourcen- gewinn, der in Reproduktion investiert werden kann (natürliche Komponente der Selektion) und dem nicht zufällig differentiellen Partnergewinn (sexuelle Kompo- nente der Selektion). Unter dem hier behandelten Gesichtspunkt der kausalen Erklärung von phylogenetischen Mustern und der ihnen zugrunde liegenden Mechanismen der ökologischen Differenzierung ist von besonderem Interesse, dass LANDE (1982) die Möglichkeit untersucht hat, ob ent- lang von Merkmalsklinen Artbildung durch die Wir- kung der sexuellen Komponente der Selektion möglich ist. Er zeigte, dass es durch den Fisher-Prozess grund- sätzlich möglich ist, dass sich parapatrische Populatio- nen in verschiedene Richtungen entwickeln und dadurch starke Unterschiede in Geschlechtspartner-Erkennungs- systemen herausbilden, die schließlich zu neuen Arten führen können. Im Zusammenhang mit der ökologischen Differenzie- rung einer Art während des Prozesses der Radiation ist das Indikator-Modell (ZAHAVI 1975; MAYNARD-SMITH 1991; IWASA et al. 1991) der sexuellen Selektion beson- ders wichtig. Modelle dieses Mechanismus gehen davon aus, dass die „Pracht“ der extravaganten Merkmale mit der Güte der Angepasstheit ihrer Träger korreliert. Die Weibchen erkennen also am sexuellen Signal (Indika- tor) die genetisch-ókologische Qualität ihres potentiel- len Partners. Dadurch wird der Einfluss des Zufalls bei der Vereinigung der väterlichen und mütterlichen Gene, der bei äußerer Besamung extrem hoch ist, sehr stark reduziert. Durch die Wirkung der sexuellen Komponen- te der Selektion wird die ökologisch-adaptive Differen- zierung von Arten stark beschleunigt (SAUER 1996). Artengruppen, deren Mitglieder starker sexueller Selek- tion unterliegen, sind in der Regel artenreicher, also ö- kologisch stärker differenziert, als solche, deren Mit- glieder im wesentlichen der natürlichen Komponente der Selektion unterliegen (SAUER 1996). Während von den Nemathelminthen (sensu AHLRICHS 1995) etwa 17.000 Arten bekannt sind, kennen wir von den Spira- liern (ohne Arthropoden) ca. 160.000 Arten. Die Arthro- poden mit komplizierten Partnerwahlmechanismen ha- ben alleine über zwei Millionen Arten hervorgebracht. Von den ,,Tentaculata* und Echinodermata, zwei subor- dinierten ursprünglichen Taxa der Radialia, sind insge- samt lediglich 12.000 Arten bekannt. Dieser grobe Vergleich der Artenzahlen macht den Zu- sammenhang deutlich, der zwischen der ókologischen Differenzierung (Artenzahl) eines Taxon und dem Ausmaß der in den Populationen seiner Mitgliedsarten wirksamen sexuellen Komponente der Selektion besteht (SAUER 1996). Arten, die starker sexueller Selektion unterliegen, soll- ten weniger gefahrdet sein, auszusterben. Fiir diese Hypothese haben wir einen indirekten Hinweis. Die Mitglieder der basalen Gruppen der Radialia haben alle eine áufere Besamung. Die Wirksamkeit der sexuellen Komponente der Selektion ist also in diesen Gruppen äußerst gering. Die „Fremdheit“, mit der die Gruppen der „Tentaculata“, Echinodermata, „Hemichordata“ und Tunicata nebeneinander stehen, lässt eine genealogische Verbindung nur schwer erkennen und macht wahr- scheinlich, dass von der adaptiven Radiation der basalen Radialia nur artenarme Restgruppen überlebt haben. Die Mehrzahl muss ausgestorben sein. Leider gibt es dazu keinen Hinweis im Fossilbericht. Diese beobachtbare Realität unterstreicht erneut die Notwendigkeit einer er- klärenden (evolutiven) Phylogenetik, d. h. die Berück- sichtigung der wirksamen Mechanismen bei der histo- risch-narrativen Erklärung der Phylogenie einer Gruppe. 4. EPILOG Die funktional-adaptive Analyse (BOCK 1981) der Merkmale, durch welche die ursprünglichen Vertreter der Gastroneuralia (= Nemathelminthes + Spiralia, sen- su AHLRICHS 1995) ausgezeichnet sind (Merkmale 1 bis 5: Abb. 1), legt nahe, die Nemathelminthes und Spiralia auf eine gemeinsame Stammlinie zurückzuführen, wie es AHLRICHS (1995) mit seinem Gastroneuralia- Konzept vorgeschlagen hat. Durch die Synthese der beobachtbaren Tatsachen und der erzählend hinzugefügten, für die Beantwortung der Frage „Warum gibt es Gastroneuralia? “ wichtigen Be- Klaus Peter SAUER & Harald KULLMANN: Analyse der biologisch-ókologischen Ursachen der Evolution 161 funde und Uberlegungen, ist es nicht gelungen, das Gastroneuraliakonzept von AHLRICHS (1995) zu falsifi- zieren. Allerdings muss hier ausdrúcklich angemerkt werden, dass die Hypothese von AHLRICHS (1995), die Schliisselart der Gastroneuralia hatte ihren millimeter- großen, kurzen, wurmförmigen und äußerlich nicht wei- ter untergliederten Körperhabitus aus dem Grundmuster der Bilateria übernommen, entschieden zurückgewiesen werden muss. Diese Auffassung AHLRICHS (1995) ist durch die mehrheitlich vertretene und akzeptierte Auf- fassung belastet, der Urbilaterier sei ein Organismus von millimetergroßem Habitus und direkter Entwick- lung (Zusammenstellung der relevanten Literatur siehe bei RIEGER et al. 1991). Wir teilen hier vielmehr die Auffassung von RIEGER et al. (1991), dass die Stammart der Bilateria ein Organismus mit pelago-benthischem Lebenszyklus und einer acoelomaten, kurzlebigen Larve von mikroskopischer Dimension gewesen sein muss. Aus einer solchen Larve mit Mundöffnung und Gastral- raum (SIEWING 1969; NYHOLM 1943) entwickelte sich in der Ontogenese ein (vielleicht coelomater, s.o.) sessi- ler Filtrierer von makroskopischer Dimension (cm- Bereich). Die Frage „Warum gibt es Gastroneuralia? * beantwor- ten wir auf folgende Weise: Die mikroskopische Kör- pergröße, die direkte Entwicklung sowie die ontogene- tisch frühe Geschlechtsreife und der damit verbundene ontogenetisch frühe Fortpflanzungszeitpunkt des Ur- gastroneuraliers wurde durch einen Lebensortwechsel der Larve eines ursprünglich noch pelago-benthischen Lebenszyklus in eine r-selektierende Umwelt erzwun- gen. Dem Lebensortwechsel ging also eine Verhaltens- änderung, die Habitat-Selektion der Larve betreffend, voraus. Mit dem Habitatwechsel in das Sediment ge- langte die Larve in einen an ungenutzten Ressourcen reichen Lebensort. Das r-selektierende Umweltszenario an diesem Lebensort wurde von unvorhersagbaren pO>- Schwankungen und damit verbundenen unvorhersagba- ren und dichteunabhängigen Mortalitátsepisoden be- stimmt. Dieser Lebensortwechsel des Urgastroneuraliers in das Sediment mariner Weichböden, wo äußere Besa- mung (der ursprüngliche Fortpflanzungsmodus der Bila- teria) aufgrund der reduzierten Begegnungswahrschein- lichkeit der Keimzellen selektionsbenachteiligt sein muss, war den Individuen der Populationen der Stamm- linie der Gastroneuralier nur durch den Erwerb von Partnerfindungs- und Partnererkennungsmechanismen möglich geworden. Diesen Merkmalskomplex der Part- nerfindung hypothetisieren wir als Schlüsselmerkmal (Autapomorphie) der Gastroneuralia. Dieser Merkmalskomplex der Partnerfindung war das präadaptive Plateau (OSCHE 1962), von dem aus die In- dividuen der Populationen der Stammlinie der Gastro- neuralia durch Ablaichen auf kleinstem Raum zunächst zur „quasi internen“ Besamung ihrer Eier gelangten. oO Das änderte das Selektionsszenario. Durch parallele Vervollkommnung wurde dann in den verschiedenen Gruppen die „innere Besamung“ durch die Entwicklung der unterschiedlichsten Genitalsitus erreicht. Die Evolu- tion eines dorsofrontalen Cerebralganglions, verbunden mit mehreren Längsnervensträngen, von denen zwei ventrolateral gelegene besonders auffallen (Merkmale 4 und 5), dürfte in ursächlichem Zusammenhang mit der umfangreiche Informationsverarbeitung notwendig ma- chenden Partnerfindung und -erkennung stehen. Part- nerfindung und -erkennung sowie „quasi innere” bzw. „innere“ Besamung waren zwingende Voraussetzungen zur Bildung der ökologischen Zone der Gastroneuralia. Danksagung. Herrn Prof. Dr. Dr. h. c. Günther Osche. Freiburg, danken wir für die kritische Durchsicht des Ma- nuskriptes und für zahlreiche wertvolle Hinweise. Frau Monika Schmied sind wir für ihre sorgfältige Reinschrift des Manuskriptes zu besonderem Dank verpflichtet. LITERATUR AHLRICHS, W. H. (1995): Ultrastruktur und Phylogenie von Seison nebeliae (Grube 1859) und Seison annulatus (Claus 1876). Hypothesen zur phylogenetischen Ver- wandtschaftsverhältnisse innerhalb der Bilateria. Diss. Universität Göttingen, Cuvillier Verlag Göttingen. ATKINSON, H. J. (1976): The Respiratory Physiology of Nematodes. Pp. 243-272 in: CROLL, N. A. (ed.): The Organisation of Nematodes. Academic Press, New York. ATKINSON, H. J. (1980): Respiration in nematodes. Pp. 101-142 in: ZUCKERMANN, B. N. (ed.): Nematodes as Biological Models, Vol. 2. Academic Press, New York. Ax, P. (1984): Das phylogenetische System. Gustav Fi- scher Verlag Stuttgart, New York. Ax, P. (1995): Das System der Metazoa I. Ein Lehrbuch der phylogenetischen Systematik. Gustav Fischer Ver- lag, Stuttgart, Jena und New York. BARTOLOMAEUS, T. (1993): Die Leibeshöhlenverhältnisse und Nephridialorgane der Bilateria-Ultrastruktur, Ent- wicklung und Evolution. Habilitationsschrift, Georg- August-Universität Göttingen. BENTON, M. J. & AYALA, F. J. (2003): Dating the Tree of Life. Science, Vol. 300: 1698-1700. BOADEN, P. J. S. (1975): Anaerobiosis, Meiofauna and early metazoan. Zoologica Scripta 4: 21-24. BOADEN, P. J. S. (1977): Thiobiotic Facts and Fancies (As- pects ot the Distribution and Evolution of Anerobic Meiofauna). Mikofauna Meeresboden 61: 45-63. BOADEN, P. J. S. & PLATT, H. M. (1971): Daily migration patterns in an intertidal meiobenthic community. Tha- lassia Jugoslavica 7: 1-12. BOADEN, P. J. S. £ ERHAG, E. A. G. (1984): Meiobenthos and the oxygen budget of intertidal sand beach. Hy- drobiologia 118: 39-47. Bock, W. J. (1981): Functional-Adaptive Analysis in Evo- lutionary Classification. American Zoologist 21: 5-20. Bock, W. J. (1979): The synthetic explanation of macroevo- lutionary change — a reductionistic approach. Bulletin the Carnegie Museum of Natural History 13: 20 — 69. 162 Bonner zoologische Beitráge 53 (2004) Bock, W. J. & VON WAHLERT, G. (1963): Two evolution- ary theories — a discussion. British Journal for the Phi- losophy of Sciences 14: 140-146. BULLOCK, T. H. & HORRIDGE, G. A. (1965): Structure and Function in the Nervous Systems of Invertebrates. W. H. Treeman and Co., San Francisco - London. BRYANT, C. (ed., 1991): Metazoan Life without Oxygen. Chapman and Hall, London etc. DARWIN, C. (1859): On the Origin of Species by Means of Natural Selection on the Preservation of Favoured Races in the Struggle for Life. Murray, London. DARWIN, C. (1871): The Descent of Man, and Selection in Relation to Sex. 1“ ed. Murray, London. DARWIN, C. (1872): The Origin of Species by Means of Natural Selection. 6" ed. Murray, London. EHLERS, U. (1985): Das Phylogenetische System der Plathelminthes. Gustav Fischer Verlag. Stuttgart - New York. FENCHEL, T. M. & RIEDL, R. J. (1970): The Sulfid System: a new biotic community underneath the oxidized layer of marine sand bottoms. Marine biology 7: 255-268. FENCHEL, T. M. & FINLAY, B. J. (1995): Ecology and Evo- lution in Anoxic Worlds. Oxford University Press, Ox- ford etc. FISHER, R. A. (1930): The Genetical Theory of Natrual Se- lection. Clarendon Press, Oxford. GOULD, S. J. (1987): Ontogeny and Phylogeny. The Belknap Press, Harvard University Press. Cambridge, Mass. - London. GUNTER, K. (1949): Uber Evolutionsfaktoren und die Be- deutung des Begriffs ökologische Lizenz. Pp. 23-54 in MAYR, E. € SCHUZ, E. (eds.) Ornithologie als biologi- sche Wissenschaft. Festschrift Stresemann. C. Winter Universitatsverlag, Heidelberg. GUNTHER, K. (1950): Okologische und funktionelle An- merkungen zur Frage des Nahrungserwerbs bei Tief- seefischen mit einem Exkurs über die ökologischen Zonen und Nischen. Pp. 55-93 in GRÜNEBERG, H. & ULRICH, W. (eds.): Moderne Biologie. Festschrift Nachtsheim. F. W. Peters, Berlin. HEMPEL, C. G. (1965): Aspects of scientific explanation. The Free Press, New York. HEMPEL, C. G. (1977): Aspekte wissenschaftlicher Erklä- rung. Walter de Gruyter, Berlin - New York. HENDELBERG, J. (1983): Plathelminthes-Turbellaria. Pp. 75-104 in: ADIYODI, K. G. & ADIYODI, R. G. (eds.): Reproductive biology of Invertebrates. Vol. 2: Sper- matogenesis and sperm function. J. Wiley and Sons Ltd. Chicester. HENNIG, W. (1982): Phylogenetische Systematik. Verlag Paul Parey, Berlin - Hamburg. HERRMANN, K. (1976): Untersuchungen über Morphologie, Physiologie und Ökologie der Metamorphose von Phoronis mülleri (Phoronida). Zoologische Jahrbücher Anatomie 95: 354-426. HERRMANN, K. (1979): Pharonis psammophila Cori (Pho- ronida, Tentaculata): Larvenentwicklung und Meta- morphose. Helgoländer Wissenschaftliche Meere- suntersuchungen 32: 550-581. HILL, A. V. (1929): The diffusion of oxygen and lactic acid through tissues. Proceedings of the Royal Society B 104: 39-96. HOGUE, E. W. (1978): Spatial and temporal dynamics of a subtidal estuarine gastrotrich assemblage. Marine Bi- ology 49: 211-222. HUMMON, W. D. (1972): Dispersion of Gastrotricha in a marine beach of the San Juan Archipelago, Washing- ton. Marine Bulletin 16: 349-355. HUTCHINSON, G. E. (1957): Concluding remarks. Cold Spring Harbor Symposia on Quantitative Biology 22: 415-427. HUTCHINSON, G. E. (1959): Homage to Santa Rosalia, or why are there so many kinds of animals? American Naturalist 93: 145-159. IWASA, Y ., POMIANKOWSKY, A. & NEE, S. (1991): The evo- lution of costly mate preferences. II. The “handicap” principle. Evolution 45: 1431-1442. JAGERSTEN, G. (1955): On the early phylogeny of the Metazoa. Zoologiska Bidrag fran Uppsala 39: 321— 354. JAGERSTEN, G. (1972): Evolution of the Metazoan Life Cycle. Academic Press, London - New York. KAESTNER, A. (1964): Lehrbuch der Speziellen Zoologie. Bd. I: Wirbellose. 1. Teil. Gustav Fischer Verlag, Stuttgart. KARLING, T. G. (1974): On the anatomy and affinities of the turbellarian orders. Pp. 1 — 16 in RISER, N. W. & MORSE, M. P. (eds.): Biology of the Turbellaria. McGraw-Hill Co, New York. : KASTING, J. F. HOLLAND, H. D. & Kump, L. R. (1992): At- mospheric Evolution: the Rise of Oxygen. Pp. 159- 163 in: SCHOPF, J. W. & KLEIN, C. (eds.): The Protero- zoic Biosphere. A multidisciplinary Study. Cambridge University Press, Cambridge, England. LANDE, R. (1982): Rapid origin of sexual isolation in char- acter convergence in a cline. Evolution 36: 213-223. LASSERRE, P. & RENAUD-MORNANT, J. (1973): Resistance and respiratory physiology of intertidal meiofauna to oxygen-deficiency. Netherlands Journal of Sea Re- search 7: 290-302. MASTERMAN, A. T. (1898): On the theory of archimeric segmentation and its bearing upon the phyletic classi- fication of the coelomata. Proceedings of the Royal Society 22: 270-310. MAYNARD SMITH, J. (1991): Theories of sexual selection. Trends in Ecology and Evolution 6: 146-214. MAYR, E. (1970): Evolution und Verhalten. Verhandlungen der deutschen zoologischen Gesellschaft 64: 322-336. NAGEL, E. (1961): The structure of science. Harcourt, Brace and World, New York. NICHOLAS, W. L. (1984): The Biology of Free-Living Nematodes. 2". ed., Oxford University Press, Oxford. NICHOLAS, W. L. (1991): Interstital Meiofauna. Pp. 129— 145 in: BRYANT, C. (ed.): Metazoan life without oxy- gen. Chapman and Hall, London etc. NYHOLM, K. G. (1943): Zur Entwicklung und Entwick- lungsbiologie der Ceriantharien und Aktinien. Zoolo- giska Bidrag fran Uppsala 22: 85-248. OSCHE, G. (1962): Das Praeadaptionsphánomen und seine Bedeutung fiir die Evolution. Zoologischer Anzeiger 162: 14-49. OSCHE, G. (1973): Das Homologisieren als eine grundle- gende Methode der Phylogenetik. Aufsätze und Reden der Senckenbergischen naturforschenden Gesellschaft 24: 155-165. OSCHE, G. (1983): Die Sonderstellung des Menschen in evolutionsökologischer Sicht. Nova acta Leopoldina NF 55: Nr. 253, 57-72. ÖSCHE, G. (2002): Gerd von Wahlert and the German Evo- lutionary Biology from 1950 — 1970 — a Companion’s Klaus Peter SAUER & Harald KULLMANN: Analyse der biologisch-ökologischen Ursachen der Evolution 163 Perspective. Bonner zoologische Monografien 50: 7 23. Ort, J. AND SCHIEMER, F. (1973): Respiration and an- aerobiosis of free-living nematodes from marine and limnic sediments. Netherlands Journal of Sea Research 7: 233-243. PARKER, G. A. (1970): Sperm competition and its evolu- tionary consequences in the insects. Biological Re- views 45: 525-567. PIANKA, E. R. (1970): On r- and K-selection. American Na- turalist 104: 592-597, REGENFUSS, H. (1975): Die Antennen-Putzeinrichtung der Adephaga (Coleoptera), parallele evolutive Vervoll- kommnung einer komplexen Struktur. Zeitschrift fúr zoologische Systematik und Evolutionsforsehung 15: 278-299. REISE, K. & AX, P. (1979): A Meiofaunal “Thiobios” Limi- ted to the Anaerobic Sulfid System of Marine Sand Does Not Exist. Marine Biology 54: 225-237. REMANE, A. (1936): Gastrotricha und Kinorhyncha. In: Bronns Klassen und Ordnungen des Tierreichs. 4. Bd., II. Abt., 1. Buch, 2. Teil, Akademische Verlagsgesell- schaft, Leipzig. REMANE, A. (1950): Die Entstehung der Metamerie der Wirbellosen. Verhandlungen der deutschen zoologi- schen Gesellschaft Mainz 1949: 16-23. RENSCH, B. (1972): Neuere Probleme der Abstammungs- lehre. Die transspezifische Evolution. 3. erw. Aufl., Ferdinand Enke Verlag, Stuttgart. RIEDL, R. (1966): Biologie der Meereshöhlen. Verlag Paul Parey. Hamburg - Berlin. RIEGER, R. M., HASZPRUNAR, G. & SCHUCHERT, P. (1991): On the origin of the Bilateria: traditional views and re- cent alternative concepts. Pp. 107-112 in SIMONETTA, A. M. & CONWAY MORRIS, S. (eds.): The early evolu- tion of Metazoa and the significance of problematic taxa. Cambridge University Press, Cambridge - New York. SAUER, K. P. (1996): Sexuelle Selektion und ókologische Differenzierung. Journal of zoological Systematics and evolutionary Research 34: 235-249, SAUER, K. P. & HOCH, M. (2002): Evolution des Nerven- systems. Anásthesiologie, Intensivmedizin, Notfall- medizin, Schmerztherapic 37: 305-313. SIEWING, R. (1969): Lehrbuch der vergleichenden Ent wicklungsgeschichte der Tiere. Verlag Paul Parey. Hamburg - Berlin. SIEWING, R. (1980): Das Archicoelomatenkonzept. Zoolo- gische Jahrbücher Anatomie 103: 439-482. SIMPSON, G. G. (1953): The major features of Evolution, Columbia University Press, New York SOUTHWOOD, T. R. E. (1980): Bionomische Strategien und Populationsparameter. Pp. 25-45 in: MAY, R. M. (ed.): Theoretische Okologie. Verlag Chemie, Weinheim - Basel. SZALAY, F. S. & Bock, W. J. (1991): Evolutionary theory and systematics: relationship between process and pat- tern. Zeitschrift für zoologische Systematik und Evolu- tionsforschung 29: 1-39. TEUCHERT, G. (1968): Zur Fortpflanzung und Entwicklung der Macrodasyoidea (Gastrotricha). Zeitschrift fiir Morphologie der Tiere 63: 343-418. WIESER, W., OTT, J., SCHIEMER, F. & GNAIGER, E. (1974): An ecophysiological study of some meiofauna species inhabiting a sandy beach at Bermuda. Marine Biology 26: 235-248. ZAHAVI, A. (1975): Mate selection a selection for a handicap. Journal of Theoretical Biology 53: 205-214. Anschrift der Autoren: Klaus Peter SAUER (korres- pondierender Autor, kpsauer@uni-bonn.de) € Harald KULLMANN, Institut fiir Evolutionsbiologie und Okolo- gie, Rheinisch-Westfälische Friedrich-Wilhelms-Uni- versität Bonn, An der Immenburg 1, D-53121 Bonn, Germany ae nj ” at Bonner zoologische Beitráge Band 53 (2004) Heft 1/2 Seiten 165-167 —_ u 4 3onn, Juni 2005 Peridea clasnaumanni spec. nov. (Lepidoptera: Notodontidae) aus China’ Alexander SCHINTLMEISTER, Dresden (Germany) Abstract. Peridea clasnaumanni is described as a new species from southeast China. It represents the southern sister- species of P. aliena (Staudinger, 1892). Zusammenfassung. Peridea clasnaumanni wird als für die Wissenschaft neue Art aus Südost-China beschrieben. Es handelt sich um die südliche nächstverwandte Art von P. aliena (Staudinger, 1892). BESCHREIBUNG Holotype: $, China, NE. Jiangxi, Wuyi Shan, 1600m, 50km SE Yingtan, 27°56’N, 117°25’E, April 2002, leg. V. Sinjaev and native collectors in coll. A. Schintlmeis- ter, Dresden. Paratypen: (1342 2 YY): 733,1 Y, NE. Jiangxi, Wuyi Shan, 1600 m, 50 km SE Yingtan, 27°56’N, 117°25’E, Mai 2002; 2 dd, ibid, Juni 2002, 4 dd, Jiangxi, Wuyi Shan, Xipaihe, 27°54’N, 117° 20”E, Juli 2003, 1500 m (GU 79-98); 1 Q, Hunan, Cili, 3.1x.1988; (alle in coll. A. Schintl- meister, Dresden, ein Y in coll. Zoologisches For- schungsinstitut und Museum Alexander Koenig, Bonn - ZFMK) Diagnose. Vorderfliigellange (rechter Vorderflúgel von Basis bis zum Apex gemessen) © 26-30 mm, im Mit- tel 28.5 mm, Y Y 34 mm. Die neue Art ähnelt habituell P. aliena (Staudinger, 1892), beschrieben aus dem A- murgebiet und P. hoenei Kiriakoff, 1963, beschrieben aus Yunnan. Sie ist deutlich kleiner als P. aliena (Vor- derfliigellange im Mittel ca. 23 mm bis maximal 25 mm bei den Ö und 27 mm bei den Y Y). Die Antennen der 34 bei P. clasnaumanni kurz ge- kámmt, bei den YY filiform. Thorax schwarzbraun, Abdomen rétlichbraun gefärbt. Die Vorderflügel schwarzbraun mit kontrastierender, orangebrauner Zeichnung. Auffallend ist das postbasale schwarz ver- dunkelte, von orangebraunen Binden eingefaßte Mittel- feld, daß im Gegensatz zu P. aliena und insbesondere zu P. hoenei die Costa der Vorderflúgel nicht erreicht. Ein gutes Erkennungsmerkmal ist ferner der orange- braune, hellbraun eingefaßte und auffallende Discoi- dalfleck. Die Vorderfliigel zeigen weiterhin eine undeutliche, auf den Hinterflügeln fortgesetzte Postme- dianbinde sowie eine gleichartige Submarginalbinde, 1 Clas Michael Naumann zu Königsbrück (26.06.1939 15.02. 2004) zum Gedenken die sich ebenfalls auf den Hinterflügeln fortsetzt. Die Hinterflügel sind schwarzgrau und deutlich dunkler ge- färbt als P. aliena oder P. honei. Charakteristisch sind die oben erwähnten Postmedian- und Submarginalbin- den der Hinterflügel, die zum Außenrand zu hell abge- setzt sind. Der Sexualdimorphismus ist habituell gering entwickelt. Die männlichen Genitalien sind denen von P. aliena ähnlich. Der Uncus von P. clasnaumanni ist weniger stark eingebuchtet und weniger ausladend, die Socci sind im Vergleich zu P. aliena dicker und auch zuge- spitzt. Die Valve ist hat keinen, wie bei P. aliena abge- setzten Apex, der dorsale Valvenfortsatz ın Nähe des Apex ist dicker und es fehlen die für P. aliena charakte- ristischen zwei kleineren Fortsätze in Nähe der Basis der Valve. Der Aedoeagus ist sehr charakteristisch durch den hakenförmigen Fortsatz (größer als bei P. a- liena) und ein ausgedehntes (größer als bei P.aliena) Gebiet mit zahlreichen Cornuti; P. hoenei hat, wie viele andere Arten der Gattung Peridea keine Cornuti. Der 8. Sternit mit dem für die Arten der Gattung typi- schen Anhängen, jedoch der Mittelteil nicht eingebuch- tet wie etwa bei P. aliena. Verbreitung. Die Typenserie von P. clasnaumanni wurde in Südost-China gefunden (Provinzen Jiangxi, Hunan). Aus SW Yunnan, Xishuangbanna, 60 km N Jinghong, 1000 m liegen mir zwei Männchen vor, die zur neuen Art gehören, jedoch deutlich heller gezeichnet sind. Möglicherweise handelt es sich um eine eigene Unterart, weshalb diese beiden Exemplare nicht in die Typenserie aufgenommen werden. clasnaumanni ist vor allem ähnlichen Genitalstrukturen (Ae- doeagus, Uncus, Socii) offenbar die Schwesterart von P. aliena. Nächste Verwandte. P. wegen der äußerst Etymologie. Dem Andenken von Clas NAUMANN ge- widmet. 166 Bonner zoologische Beiträge 53 (2004) LITERATUR SCHINTLMEISTER, A. (1989): Zoogeographie der palaeark- tischen Notodontidae (Lepidoptera). Neue Entomolo- gische Nachrichten 25: 1-117. SCHINTLMEISTER, A. [1992]: Die Zahnspinner Chinas (Le- pidoptera, Notodontidae). Nachrichten entomologi- scher Verein Apollo, Frankfurt am Main, Suppl. 11 (1991): 1-343. SCHINTLMEISTER, A & FANG, Ch. L. (2001): New and less known Notodontidae from mainland China (Insecta, Lepidoptera, Notodontidae). Neue Entomologische Nachrichten 50: 1-143. Anschrift des Autors: Dr. Alexander SCHINTLMEISTER, Calberlastr. 3, D-01326 Dresden, Germany; e-mail: schintIm@aol.com Alexander SCHINTLMEISTER: Peridea clasnaumanni spec. nov. (Lepidoptera: Notodontidae) aus China 167 6 | Abb. 1-4: Imagines Peridea. (1) Peridea clasnaumanni — 3, China, NE. Jiangxi, Wuyi Shan, 1600 m, 50 km SE Yingtan, 27°56’N, 117°25’E, April 2002, Holotype. (2) Peridea clasnaumanni — Y, China, Hunan, Cili, 3.ix.1988, Paratype. (3) Peridea aliena — 3, Russia, Primorye, 30 km NE Wladiwostok, 24.vii.1963. (4) Peridea hoenei — Y, China, Sichuan, Daxue Shan, 40 km W Mianning, 2750 m, 7.-8.v11.1998. Abb. 5-6: Genitalapparate Peridea. (5) Peridea clasnaumanni — &, China, Jiangxi, Wuyi Shan, Xipaihe, 27°54’N, 117° 20°E, Ju- | ly 2003, 1500 m (GU 79-98), Paratype. (6) Peridea aliena — 3, N. Korea, Myohang san, Myohang-Shon-valley, 24.-30.vi.1985 | (GU 24-73). Be I oy Bonner zoologische Beiträge Band 53 (2004) | Heft 1/2 | Seiten 169-185 3onn, Juni 2005 The Variance of Variation: Geographic Patterns of Coat Colouration in Anomalurops and R . . 1 Anomalurus (Mammalia, Rodentia, Anomaluridae) / Anja C. SCHUNKE & Rainer HUTTERER Zoologisches Forschungsmuseum Alexander Koenig, Bonn, Germany Abstract. Variation of coat colouration in Anomalurops beecrofti, Anomalurus derbianus, A. pelii, and A. pusillus was studied. Character states of the dorsal and ventral colouration were defined for each species and specimens assigned to classes accordingly. Geographic patterns were analysed after excluding sex, age, and collection month as possible causes for different colouration. Considerable differences were found among the first three species, while no differences in colouration were found in A. pusillus. Anomalurops beecrofti varies in the distribution area mainly in the frequencies of different colour morphs, while in Anomalurus derbianus several character states are clearly restricted to defined ar- eas. In A. pelii the correlation between colouration and locality is very strong, although its distribution area is much smaller than in the other species. Geographic barriers, especially rivers, are discussed as possible causes for the ob- served variation. Key words. Scaly-tailed flying squirrels, morphology, colouration, Africa, biogeography. 1. INTRODUCTION' The history of the African rain forest has attracted the in- terest of numerous researchers for a long time (e.g. BRSTRUP 1935; FJELDSA & LOVETT 1997; GRUBB 1978, 1990; HAMILTON & TAYLOR 1991; LONNBERG 1929). Be- sides more or less direct methods like the analysis of fossil pollen records (e.g. BRENAC 1988; FREDOUX & TASTET 1988; LIVINGSTONE 1966; MALEY 1983, 1991), the recent distribution of species and subspecies of animals was used to reconstruct this history. The anomalurid species investi- gated here are all able to perform a gliding flight and de- pend on the occurrence of large trees as starting and land- ing points. They are strictly arboreal and behave clumsily on the ground, and therefore their current distribution pat- terns may reveal information on the history of their habitat, the African rain forest. Considerable differences are found in the variation of coat colouration in the species of Anomaluridae. Anomalurus pusillus Thomas, 1887, Idiurus macrotis Miller, 1898, /. zenkeri Matschie, 1894 and Zenkerella insignis Matschie, 1898 show a more or less uniform colouration throughout their distribution range. Anomalurops beecrofti (Fraser, 1853) varies in the amount of golden brown colour on the back and reddish colour on the ventral side. The most pro- nounced colour variation is found in Anomalurus derbi- anus (Gray, 1842), with a range from more or less uni- formly brownish individuals to colourful individuals with several defined markings. Anomalurus pelii (Schlegel & 1 In commemoration of Clas Michael Naumann zu Königsbrück (26.06.1939 — 15.02.2004) Miiller, 1845) shows only black and white in a specific pat- tern, but the relative portions of these colours vary. This study describes the variation of the coat colouration in one species of Anomalurops and in three species of Anomalurus on the basis of 966 museum specimens. This sample represents the majority of skins of these species kept in research collections worldwide and is regarded as a reliable basis for a study of geographic variation. 2. MATERIAL AND METHODS 2.1. Data basis Specimens used for this analysis were studied in 15 col- lections of the following institutes: American Museum of Natural History (AMNH, New York). The Natural History Museum (BMNH, London), Field Museum of Natural History (FMNH, Chicago), Liverpool Museum (LIVCM, Liverpool), Museum National d' Histoire Naturelle (MNHN, Paris), Musée Royal d’Afrique Cen- trale (MRAC, Tervuren), Naturhistorisches Museum Basel (NHMB, Basel), Naturhistorisches Museum Wien (NMW), Naturhistoriska Riksmuseet (NRM, Stock- holm), Naturalis/Nationaal Natuurhistorisch Museum (RMNH, Leiden), Naturmuseum Senckenberg (SMF, Frankfurt/Main), National Museum of Natural History (USNM, Washington), Zoologisches Forschungsinstitut und Museum Alexander Koenig (ZFMK, Bonn). Zoölo- gisch Museum Amsterdam (ZMA), and Museum fiir Naturkunde (ZMB, Berlin). The colour and colour pat- terns were analysed from colour slides taken together with a colour reference. Photographs were taken of the dorsal side of all specimens and as many as possible (depending on the preparation) of the ventral side of 4. 170 Bonner zoologische Beitráge 53 (2004) beecrofti and of several specimens of 4. derbianus and A. pelii. Sex was either determined from the skins or la- bels, from which also the date of collection was re- corded. The relative age was estimated from tooth wear if the skull was available (details to be published else- where). In addition, slides were taken of 128 specimens of A. pusillus and compared but not used in the final - analysis because it was not possible to define any dif- ferences in colouration. The data basis for the analyses is given in Table 1. The maps given in Figures 3 to 5 are based exclusively on skins with coordinates of the col- lection locality. 2.2. Statistics Relationships between the various colouration charac- ters and sex, age, and month of collection were checked with chi-square tests performed in SPSS 10.0. In the re- ported results the number of specimens used for the and P values are given (n.s. = not significant). Additionally, the percentage of cells with an expected frequency less h SRGSE8 nun BEEERE Giese A Bann than five (CEF B E @ DD Dorsal colduration 4 > E Dorsal colouration 5 Ventral colouration 1 (0 E Ventral colouration 2 A [MM] Ventral colouration 3 e@ BB Ventral colouration 4 Fig. 3: Geographical distribution of coat colouration in Anomalurops beecrofti (see Fig. 6 tor border lines). 174 colouration exists between S-Democratic Republic of Congo and E-Angola. As there is only one ventral colouration type (VC 1) in Angola, no statistical calcu- lation for ventral colour frequencies was possible (the same technical problem applies to the dorsal colouration frequencies on Bioko and in a small area north of the Congo River; see Fig. 3). a A yl FF bn paras f Bonner zoologische Beitráge 53 (2004) 3.2. Anomalurus derbianus This species shows the highest variability in colouration (Plate II), ranging from more or less uniformly brownish individuals to specimens with a reddish back, dark grey patagia, black ears and a silvery stripe on the nose. The ventral colouration is much less variable. g h Plate II: Typical representatives of the dorsal and ventral colouration classes in Anomalurus derbianus (see text for details). a: dorsal colouration DC 1 (ZFMK 64.491), b: DC 2 (ZFMK 69.148), c: DC 3 (ZFMK 73.363), d: DC 4 (ZFMK 64.817), e: ventral colouration VC 1 (FMNH 88207), f: VC 2 (ZFMK 64.493), g: VC 3 (ZFMK 69.148), h: VC 4 (BMNH 90.6.8.18). Colouration codes and frequencies Dorsal colouration (DC): 1. more or less uniformly brown (70%, n=329). 2. back more reddish brown than the rather greyish patagia, no sharp border (19%, n=90). 3. back reddish and patagia dark grey, with sharp bor- der (10%, n=45). 4. silvery grey (1%, n=5). Ears (E): 1. considerably darker than rest of the head (66%, n=295). 2. more or less the same colouration as the rest of the head (34%, n=154). Shoulders (S): 1. considerably lighter than neck (42%, n=192). 2. more or less the same colouration as the neck (48%, =267). Throat (T): 1. dark ring around neck, mainly closed ventrally (63%, n=85). 2. throat partially light, dark ring ventrally open (37%, n=50). Anja C. SCHUNKE & Rainer HUTTERER: Geographic Patterns of Coat Colouration in Anomalurops and Anomaluru 175 Ventral colouration (VC): 1. whitish (52%, n=71). 2. greyish without yellow (31%, n=42). 3. yellowish mixed with grey (11%, n=15). 4. light yellow (7%, n=9). Dorsal and ear colouration Dorsal and ear colouration show a highly significant correlation. When all character states are used in the analysis, 25% of the cells have an expected frequency of less than five and the MEF is 1.71 (n = 449, y" 86.637, d.f. 3, P<0.001). After omitting DC 4, all cells have an expected frequency of more than five and the MEF is 15.20 (n = 444, x? 82.553, d.f. 2, P < 0.001). The corre- lation depends on the dorsal colouration class. About half (53%) of the more or less uniformly brown speci- mens (DC 1) have ears considerably darker than the rest of the head, the others (47%) have ears of the same col- our as the head. Individuals with a back slightly (DC 2) or clearly more reddish than the patagia (DC 3) have almost exclusively dark ears (97% and 100% respec- tively), while most silvery (DC 4) specimens have ears of the same colouration as the head. Dorsal and shoulder colouration The correlation for dorsal and shoulder colouration shows results equal to those for dorsal and ear colour- ation. When all character states are used in the analysis, 25% of the cells have an expected frequency of less than five and the MEF is 2.09 (n = 459, y” 194.976, d.f. 3, P < 0.001). After omitting DC 4, all cells have an ex- pected frequency of more than five and the MEF is 15.20 (n = 454, x 190.767, d.f. 2, P < 0.001). The cor- relation of dorsal and shoulder colouration also depends on the colouration class. Only 22% of the uniformly brown individuals (DC 1) have light shoulders, but 88% of the specimens with a slightly reddish back (DC 2), and 100% of those with a strongly reddish back (DC 3). None of the silvery individuals (DC 4) have light shoul- ders. Ear and shoulder colouration Ear and shoulder colouration are also significantly cor- related (n = 444, y” 68.593, d.f. 1, P < 0.001). Some 88% of the specimens with light shoulders have dark ears. In the specimens with shoulders of the same colouration as the neck, both types of ear colouration occur equally. Dorsal and ventral colouration Correlation between dorsal and ventral colouration ap- pears to be weak. When all four character states for both are used in the analysis, the result is not significant (n = 136, x 14.845, d.f. 9, P = n.s.), but 56% of the cells have an expected frequency less than five and the MEF is 0.13. When DC 4 is omitted from the analysis, 42% of the cells have an expected frequency less than five, the MEF is 0.74 and the result shows a slightly signifi- cant correlation between dorsal and ventral colouration (n = 134, y” 14.286, d.f. 6, P < 0.05). The most reliable results were obtained after ventral colouration 4 was omitted as well. Then the percentage of CEF BB - A & = % | unusen > MT [I one SEE 00m 1 r F b ayy E A 3 asus ha y data set. This caused problems for a reliable calculation of statistically significant differences. However, in Tan- zanja specimens from the Usambara Mountains show mainly yellow and yellow mixed with grey underparts, while those from further south are whitish or greyish without yellow. / 3.3. Anomalurus pelii This species is generally black with white margins of the patagia, a white tail and a ventral colouration rang- ing from white to blackish grey. White markings can also be completely absent. The variation in A. pelii is mainly defined by the relative portions of black and white (Plate III). rs nu... mmm Cc d (9, -—TRBRO ore : 48 gk. Wr AN wur BERNER ku. g Plate III: Typical representatives of the dorsal and ventral colouration classes in Anomalurus pelii (see text for details). a: dorsal colouration DC 1 (ZMA 21.400), b: DC 2 (ZMA 21.262), c: DC 3 (ZMA 21.277), d: DC 4 (ZMA 21.282), e: ventral colouration VC 1 (ZMA 21.400), f: VC 2 (ZMA 21.266), g: VC 3 (ZMA 21.277). ¡93 Colouration codes and frequencies white margins less than one-half of pleuropatagia, : = connected with uropatagia margins (23%. n=31). Dorsal colouration (DC): Poe 5 N JE 2. white margins less than one-third of pleuropatagia and separated from uropatagia margins (40%, n=55). 4. white margins more than one-half of pleuropatagia, completely black (7%, n=9). shoulder frequently also white (31%. n=43). 180 Bonner zoologische Beitráge 53 (2004) Ventral colouration (VC): greyish with blackish central stripe (45%, n=26). white with blackish central stripe (45%, n=26). WN — completely white (10%, n=6). Dorsal and ventral colouration A highly significant correlation between dorsal and ven- tral colouration in A. pelii was shown after omitting DC l and VC 1 (n= 49, y? 13.177, d.f. 2, P < 0.001) as well as in all previous steps of the analysis. In A. pelii a correlation between dorsal and ventral colouration is evident, as individuals with a dark dorsal surface tend to be dark on the ventral side, too. The two darker dorsal colouration classes are linked to the dark- est ventral colouration in 90% of all specimens, while the relatively rare completely white ventral colouration is restricted to the two dorsal colouration classes with extensive white markings. However, the latter can be combined with every possible ventral colouration, al- though the darkest colouration is found in less than 12% of the specimens only. Colouration and sex A total of 35 males (56%) and 27 females (44%) were available. The data showed no significant correlation between dorsal or ventral colouration and sex. Colouration and age Some 117 animals were grouped into age class | (5%, n=6), 2 (19%, n=22), 3 (41%, n=48), and 4 (35%, n=41). The data showed no significant association be- tween colouration and age. Colouration and collection area Localities and diagrams of colouration class frequencies in the respective areas are shown in Figure 5, border lines between the areas in Figure 6. In spite of the ex- tremely small area inhabited by 4. pelii, a strong geo- graphic variation exists in the extension of the white markings. West of the Sassandra River only entirely black individuals are found. Specimens with large white markings of the colour classes 3 and 4 are mainly re- stricted to a relatively small area between the Sassandra and Bandama rivers. In the triangle between the Ban- dama and Nzi rivers three specimens of the colour classes 2 and 3 were found, thus forming a transition to the area east of the Bandama and Nzi rivers to the Volta River, where exclusively individuals with small white markings of the colour class 2 occur. 3.4. Combined analysis of geographic boundaries Based on the results shown in Figures 3-5, an attempt was made to identify general borders in the distribution patterns of the characters analysed. Seventeen border lines of different significance for the respective species and colouration characters were recognized (Fig. 6). The Sassandra (Fig. 6, line 1) and Bandama and Nzi (Fig. 6, line 2) rivers obviously are effective barriers for A. pelii, but not for A. beecrofti and A. derbianus. The Volta (Fig. 6, line 3) marks the eastern boundary of the distribution of A. pelii and also seems to play a role for A. beecrofti. Unfortunately, only very few skins of A. beecrofti and none of A. derbianus were available from the small area between the Volta and the Dahomey Gap, thus statements concerning this region have to be treated with caution. The Dahomey Gap (Fig. 6, line 4) is a sig- nificant barrier for colouration frequencies in A. bee- crofti, but only the shoulder colouration of A. derbianus displays a change there, while dorsal and ear colour- ation are not affected. An important border line (Fig. 6, line 5) that significantly separates frequencies for al- most all investigated characters, except for the shoulder colouration in A. derbianus, exists between Nigeria and the highlands of western Cameroon. This highland area differs also from the adjacent plains (Fig. 6, line 6) in the frequencies of dorsal and shoulder colouration of A. derbianus and dorsal colouration of A. beecrofti. From S-Cameroon to Gabon and SW-Central African Repub- lic, the colouration frequencies are homogenous for A. derbianus and quite so for A. beecrofti, with the excep- tion of uniformly reddish specimens from a restricted area in the coastal plains of Equatorial Guinea and N- Gabon. Unfortunately very few individuals were col- lected in Congo Brazzaville, therefore it is not possible to give statements concerning this country. Specimens of A. derbianus from the southern tip of South Ogooue (NW of the Congo River, Fig. 6, line 7; see GAUTIER- HION et al., 1999 for definition of the area) show no dif- ferences in colouration frequencies to those from S- Cameroon to Gabon. The small sample of A. beecrofti from the same area uniformly shows the relatively rare dorsal colouration 3. Bioko is a special case (Fig. 6, line 8). Specimens of A. beecrofti have a uniform dorsal colouration which other- wise is found mainly in W-Cameroon highlands and in Angola. Specimens from Nigeria have completely dif- ferent colourations. Individuals of A. derbianus have identical dorsal colouration frequencies on Bioko and in West Africa (Liberia to Nigeria), but frequencies differ significantly from those of the region between Camer- oon and Congo. The frequencies of the ear colouration on Bioko are intermediate to the frequencies in Nigeria and the Cameroon to Congo area. Light shoulders as typical for West Africa occur on Bioko in a frequency intermediate to that of West Africa and Central and East Africa. The border effect of the lower Congo (Fig. 6, line 9) and Ubangi (Fig. 6, line 10) is also difficult to define. They form clear boundaries for dorsal and shoulder coloura- tion in A. derbianus, but not for ear colouration. In Anja C. SCHUNKE & Rainer HUTTERER: Geographic Patterns of Coat Colouration in Inomalurops and Anomaluru 18] PAN Dorsal colouration 1 Dorsal colourátion 3 | e [N Dorsal colouration 2 = O Dorsal colouration 4 @ BB Ventral colouration 1 O [M ventrai colouration 2 © [L] Ventral colouration 3 Fig. 5: Geographical distribution of coat colouration in Anomalurus pelii (see Fig. 6 for border lines), 182 Bonner zoologische Beitráge 53 (2004) Fig. 6: Border lines of greater (bold) and lesser (dotted) importance for colouration changes in Anomalurops beecrofti, Anomalu- rus derbianus and A. pelii (see text for details). A. beecrofti this river system forms a significant barrier for ventral colouration, but only the Ubangi part signifi- cantly separates neighbouring populations in dorsal colouration. Results for the area between the Congo River and the Ubangi, Uélle, and Kibali river system are also contra- dictory. For A. beecrofti, a significant boundary in dor- sal colouration frequencies between the northwestern and southeastern part of the range (Fig. 6, line 13) cor- responds to the Aruwimi and Ituri river system, but is not matched by the ventral colouration. Dorsal and shoulder colouration of A. derbianus are uniform in this area, but there are differences in ear colouration be- tween specimens from the northwest and souteast (Fig. 6, line 12). However, this boundary lies further west than in 4. beecrofti. The north bow of the Congo River (Fig. 6, line 11) seems to have no effect on the dorsal, ear and shoulder colouration frequencies in 4. derbi- anus. The same applies to the northwestern population of A. beecrofti, while southeastern specimens are sig- nificantly different from those south of the Congo (Fig. 6, line 14). Specimens of A. beecrofti from W-Angola seem to be different from individuals caught in the Congo Basin (Fig. 6, line 15), but the sample size is small. For A. derbianus the situation is more complex, because this species occurs also in E-Angola, Zambia, and Tanzania (Fig. 6, line 16). There are only slight dif- ferences between specimens from the Congo basin and W-Angola. Light epaulettes are missing in all specimens from Angola, S-Democratic Republic of Congo, Zambia and Tanzania. In the same area (except Angola) the rare uniformly silvery specimens occur, and the highest amount of ears displaying the same colouration as the rest of the head is found here too. In the Usambara Mountains the ventral colouration of 4. derbianus dif- fers from that of more southern individuals (Fig. 6, line 17): 4. DISCUSSION Our study has revealed a remarkable mosaic of differ- ences as well as similarities between the respective spe- cies. Anomalurus pusillus, a species studied but results not shown in detail here, shows a homogeneous brownish colouration throughout its range from Camer- oon to E-Democratic Republic of Congo. Anomalurops beecrofti varies significantly in the frequencies of the colouration forms but the majority (ca. 70%) of speci- mens belong to only one of the two colouration forms that occur throughout the distribution area of the spe- Anja C. SCHUNKE & Rainer HUTTERER: Geographic Patterns of Coat Colouration in Anomalurops and Anomalurus 83 cies. In Anomalurus derbianus the correlation between locality and colouration is more pronounced, although dorsal, ear and shoulder colouration follow slightly dif- ferent geographic patterns. In Anomalurus pelii the dor- sal colouration follows clear-cut lines, a pattern not found in any other species. Despite the particular differences between the species, some general geographic patterns can be extracted. Some boundaries seem to have a major impact on the distribution and frequency of the various colour morphs in the studied species (Fig. 6): The Sassandra, the Ban- dama-Nzi river system, the Volta River and the Dahomey Gap in West Africa, the highlands of W- Cameroon, the lower Congo and Ubangi river system, and a border line running from Katanga to NE-Tan- zania, which separates the southwestern parts of the dis- tribution area of 4. derbianus from the rest. The Congo River seems to have a low impact on the geographic colouration pattern in A. beecrofti and A. derbianus, es- pecially in its middle part. Some of these boundaries are found in other mammalian species as well, but others not. Primates are a well- studied group suited for comparison. Subspecific changes occur frequently in the Ivory Coast, often in the vicinity of the Sassandra River (BOOTH 1958; DAN- DELOT 1965; GRUBB 1990; LERNOULD 1988; OATES 1988). The Bandama River forms a border between sub- species of Procolobus badius (BOOTH 1958; OATES 1988). However, both rivers do not seem to represent barriers for guenons as clear as for A. pelii. The Da- homey Gap is a well known border for primates species (BOOTH 1958; GRUBB 1990; LERNOULD 1988; OATES 1988). The highlands of W-Cameroon are a centre of endemism for mammals, and also a border for some guenon subspecies (LERNOULD 1988). Further south, the Sanaga River forms a barrier for many primates (LER- NOULD 1988). A further subdivision of the area between the Sanaga and Congo and Ubangi rivers, as found in mitochondrial haplotypes of gorillas (CLIFFORD et al. 2004), could not be observed in anomalurids. The lower Congo and Ubangi rivers are common barriers for nu- merous primates (DANDELOT 1965; GAUTIER-HION et al. 1999; GRUBB 1990; LERNOULD 1988). In the area be- tween the Congo and Ubangi rivers, no river forms an obvious barrier. However, hybridisation zones or changes of subspecies of primates are frequently found north of the Congo River between the lower Ubangi in the west and the Ituri River and the Kivu region in the east (COLYN 1987, 1988; COLYN et al. 1991; DANDELOT 1965; GAUTIER-HION et al. 1999). The Uélle River, which marks the northern border of the distribution area of the anomalurids, has the same significance for some primates but not for others (COLYN 1987; GAUTIER- HION et al. 1999; LERNOULD 1988). The Congo River, which delimits the inner Congo Basin to the north and forms a significant barrier for primates (COLYN 1987, 1988; COLYN & DELEPORTE 2002; DANDELOT 1965; GRUBB 1990; LERNOULD 1988) and other mammals, seems to have little importance for the distribution of colour patterns in A. beecrofti and A. derbianus. The Rift Valley is also a significant barrier for guenons (LERNOULD 1988), and although it seems to have no in- fluence on A. derbianus, it is the western border for three other species of anomalurids (A. pusillus, Idiurus macrotis, and I. zenkeri). Anomalurus derbianus appar- ently has a high potential of dispersal as the species not only crosses the Rift Valley but also extends far south into Zambia and Tanzania. Generally the distribution of coat colouration patterns in anomalurids is strongly correlated with the occurrence of larger rivers. This seems remarkable for animals with an ability for gliding flight. Anomalurus pelii has been observed to glide for 50 m (DEKEYSER 1954). Distances of 15 to 20 m (DELANY 1975) and even up to 100 m (KINGDON 1974) were reported for A. derbianus, and flights over a distance of 250 m are assumed as possible (KINGDON 1974; MACKAY & CUNNIGHAM VAN SOMEREN 1980). However, larger rivers seem to form barriers that are not regularly crossed. Unfortunately not much is known about the behaviour of anomalurids, but their common way of locomotion is gliding flight from one tree to another, starting from and landing on trunks or branches, and then climbing up the trunk for the next start (ADAMS 1894; KINGDON 1974; RAHM 1969). What do the observed patterns tell us about the evolu- tionary history of these anomalurids? First, the patterns are more complex than expected. Anomalurus pelii is confined to a small area in West Africa where it estab- lished three distinct populations separated by the Sas- sandra and Bandama rivers (Fig. 5). Although genetic data are not yet available, we assume that reduced gene flow exists between the three populations (SCHUNKE & HUTTERER in press). The situation in 4. derbianus and Anomalurops bee- crofti is more complex (Figs. 3, 4). Some of the patterns agree with biogeographic units identified by COLYN & DELEPORTE (2002) in their analysis of forest guenons. Particularly the West Central faunal area (NW of Congo River to Cameroon Mts) is reflected by the distribution of the anomalurids. COLYN & DELEPORTE (2002) found several subunits in the area, apparently a result of fluc- tuating savanna and forest vegetation in this area in the Quaternary. The distribution of the shoulder colouration in A. derbianus (Fig. 4, bottom) fits this picture. During deteriorating conditions, animals with dark shoulders may have retreated into the Congo Basin and the Cam- eroon Mts refuge, and animals with pale shoulders into a West African refuge. In times of ameliorating condi- tions, the West African population dispersed into the 184 former savanna corridor and filled this gap with pale- shouldered animals. The true picture was certainly more complex, the de- tails, however, must still be filled in. Cladogenesis and secondary hybridization at contact zones have probably obscured the original patterns. The periodic model of cladogenesis in African mammals (GRUBB 1999) seems to be well-suited to explain the current patterns. Genetic data are needed to know to which extent cladogenesis has occurred, and to solve the phylogeography of the group. Acknowledgements. This publication benefited from the kind help of curators and staff members of the following institutions: Bob Randall, Pat Brunauer, and Nancy B. Simmons (AMNH), Paula D. Jenkins and Daphne Hills (BMNH), Bruce D. Patterson (FMNH), Clem Fisher (LIVCM), Michel Tranier, Jacques Cuisin and Jean- Francois Ducroz (MNHN), Wim Van Neer and Wim Wendelen (MRAC, Tervuren), Raffael Winkler and Urs Rahm (NHMB), Friederike Spitzenberger and Barbara Herzig (NMW), Olavi Grónwall (NRM), Chris Smeenk (RMNH), Dieter Kock (SMF), Richard W. Thorington, Jr., Michael D. Carleton, Linda K. Gordon, and Helen L. Kafka (USNM), Miguel Vences and Adrian G. Rol (ZMA), and Manfred Ade, Irene Thomas, and Detlef Willborn (ZMB).We thank Jakob Fahr, Gustav Peters, and Bradley Sinclair for constructive criticism of the manuscript. Fi- nancial support was provided by the Deutsche Forschungs- gemeinschaft (DFG, HU 430/1-1 and 430/1-2). Visits to BMNH and MNHN were funded by the TMR Programme of the European Commission (Bioresource London and Parsyst Paris). REFERENCES ADAMS, W. H. (1894): On the habits of the flying-squirrels of the genus Anomalurus. Proceedings of the Zoologi- cal Society of London: 243-246. Booth, A. H. (1958): The zoogeography of West African primates: A review. Bulletin de l'Institut Francais d'Afrique Noire (A) 20: 587-622. BRAESTRUP, F. W. (1935): Remarks on climatic change and faunal evolution in Africa. Zoogeographica 2: 484- 494. BRENAC, P. (1988): Evolution de la végétation et du climat dans l'Ouest Cameroun entre 25 000 et 11 000 ans BP. Actes Xeme Symposium Ass. Palynologues Langue Francaise, Trav. Sect. Sci. Tech. Inst. Francais Pondi- chéry 25: 91-103. CLIFFORD, S. L., ANTHONY, N. M., BAWE-JOHNSON, M., ABERNETHY, K. A., TUTIN, C. E. G., WHITE, L. J. T., BERMEJO, M., GOLDSMITH, M. L., MCFARLAND, K., JEFFREY, K. J., BRUFORD, M. W. & WICKINGS, E. J. (2004): Mitochondrial DNA phylogeography of wes- tern lowland gorillas (Gorilla gorilla gorilla). Molecu- lar Ecology 13: 1551-1565. COLYN, M. (1987): Les primates des foréts ombrophiles de la cuvette du Zaire: interprétations zoogéographiques de modeles de distribution. Revue de Zoologie africai- nes 101: 181-196. Bonner zoologische Beitráge 53 (2004) COLYN, M. (1988): Distribution of guenons in the Zaire- Lualaba-Lomami river system. Pp. 104-124 in: GAU- TIER-HION, A., BOURLIERE, F., GAUTIER, J.-P. & KING- DON, J. (eds.) A primate radiation: evolutionary biol- ogy of the African guenons. Cambridge University Press, Cambridge. COLYN, M. & DELEPORTE, P. (2002): Biogeographic analy- sis of Central African forest guenons. Pp. 61-78 in: GLENN, M. E. & Corps, M. (eds.) The guenons: Diver- sity and adaptation in African monkeys. Kluwer Aca- demic/Plenum Publishers, New York. COLYN, M., GAUTIER-HION, A. & VERHEYEN, W. (1991): A re-appraisal of palaeoenvironmental history in Central Africa: evidence for a major fluvial refuge in the Zaire Basin. Journal of Biogeography 18: 403-407. DANDELOT, P. (1965): Distribution de quelques especes de Cercopithecidae en relation avec les zones de végéta- tion de l'Afrique. Zoologica africana 1:167-176. DEKEYSER, P. L. (1954): A propos des ecureuils volants. Notes africaines 64:121-124. DELANY, M. J. (1975): The Rodents of Uganda. Trustees of the British Museum (Natural History), London. FJELDSA, J. & LOVETT, J. C. (1997): Geographical patterns of old and young species in African forest biota: the significance of specific montane areas as evolutionary centres. Biodiversity and Conservation 6: 325-346. FREDOUX, A. & TASTET, J. P. (1988): Stratigraphie pollini- que et paléoclimatologie de la marge septentrionale du Golfe de Guinée depuis 200.000 ans. Institut Francais Pondichery, Travaux Section Science et Technique 25:175-183. GAUTIER-HION, A., COLYN , M. & GAUTIER, J.-P. (1999): Histoire naturelle des primates d'Afrique Centrale. ECOFAC, Libreville. GRUBB, P. (1978): Patterns of speciation in African mam- mals. Bulletin of Carnegie Museum of Natural History 6:152-167. GRUBB, P. (1990): Primate geography in the Afro-tropical forest biome. Pp. 187-214 in: PETERS, G., & HUT- TERER, R. (eds.) Vertebrates in the tropics. Proceedings of the International Symposium on Vertebrate Bio- geography and Systematics in the Tropics, Bonn, June 5-8, 1989. Museum Alexander Koenig, Bonn. GRUBB, P. (1999): Evolutionary processes implicit in dis- tribution patterns of modern African mammals. Pp. 150-164 in: BROMAGE, T. G., & SCHRENK, F. (eds.) Af- rican biogeography, climate change, & human evolu- tion. Oxford University Press, New York & Oxford. HAMILTON, A. C. & TAYLOR, D. (1991): History of climate and forests in tropical Africa during the last 8 Million years. Climatic Change 19: 65-78. JAHNS, S., HULS, M. & SARNTHEIM, M. (1998): Vegetation and climate history of West Equatorial Africa based on a marine pollen record off Liberia (site GIK 16775) covering the last 400,000 years. Revue Palaeobotanie et Palynologie 102: 277-288. KINGDON, J. (1974): East African Mammals. An Atlas of Evolution in Africa. Volume II. Part B (Hares and Ro- dents). Academic Press, London, New York. LERNOULD, J.-M.(1988): Classification and geographical distribution of guenons: a review. Pp. 54-78 in: GAU- TIER-HION, A., BOURLIERE, F., GAUTIER, J.-P. & KING- DON, J. (eds.) A primate radiation: evolutionary biol- ogy of the African guenons. Cambridge University Press, Cambridge. Anja C. SCHUNKE & Rainer HUTTERER: Geographic Patterns of Coat Colouration in Anomalurops and Anomalurus 8 LIVINGSTONE, D. A. (1967): Postglacial vegetation of the Ruwenzori Mountains in Equatorial Africa. Ecological Monographs 37: 25-52. LONNBERG, E. (1929): The Development and distribution of the African Fauna in connection with and depend- ing upon Climatic Changes. Arkiv foer Zoologi 21A (4): 1-33. MACKAY, A. & CUNNINGHAM VAN SOMEREN, G. R. (1980): Some observations on anomalures near Chemisia, north Nandi, Kenya. East African Natural History So- ciety Bulletin (May-June): 42-43. MALEY, J. (1983): Histoire de la végétation et du climat de l'Afrique nord-tropicale au Quaternaire recent. Bothalia 14 (3-4): 377-389. MALEY, J. (1991): The African rain forest vegetation and paleoenvironments during Late Quaternary. Climatic Change 19: 79-98. OATES, J. F. (1988): The distribution of Cercopithecus monkeys in West African forests. Pp. 79-103 in: GAU- TIER-HION, A., BOURLIERE, F., GAUTIER, J.-P. & KING- DON, J. (eds.) A primate radiation: evolutionary biol- ogy of the African. guenons. Cambridge University Press, Cambridge. RAHM, U. (1969): Dokumente über Anomalurus und /diu rus des östlichen Kongo. Zeitschrift für Säugetierkun- de 34: 75-84. SCHUNKE, A. C. & HUTTERER, R. (in press): Geographic variation in the West African scaly-tailed squirrtl Anomalurus pelii (Schlegel and Müller, 1845) and de- scription of a new subspecies (Rodentia: Anomaluri- dae). Pp 321-328 in: HUBER, B. A., SINCLAIR, B. J. & LAMPE, K.-H. (eds.) African Biodiversity: Molecules, Organisms, Ecosystems. Proceedings of the Sth Inter- national Symposium on Tropical Biology, Museum Koenig, Bonn. Springer Verlag, Berlin. Authors’ address: Anja C. SCHUNKE (corresponding author) and Rainer HUTTERER, Zoologisches Forschungs- museum Alexander Koenig, Adenauerallee 160, D- 53113 Bonn, Germany, e-mail: a.schunke.zfmk@uni- bonn.de Table 1: Characters and respective numbers of specimens analysed in this study. A. pusillus was not analysed in detail because no appreciable variation exists Character/Species A. beecrofti A. derbianus A. pelii A. pusillus Dorsal colouration 227 469 142 128 Ventral colouration 169 137 58 48 Ear colouration not used 449 not used not used Shoulder colouration not used 459 not used not used Throat colouration not used 135 not used not used Sex 156 342 64 not used Age class 160 365 117 not used Month of collection 177 359 109 not used Locality coordinates 208 421 118 113 Pa Mase E E . a { ‘ E E vn + > , Bonner zoologische Beitrage Band 53 (2004) | Heft 1/2 | Seiten 187-191 3onn, Juni 2005 Issues in Delimiting Genera in Invertebrates: an Example from the Lepidoptera (Macariini: Geometridae: Ennominae)' J Malcolm J. SCOBLE Department of Entomology, The Natural History Museum, London, UK Abstract. Deciding on how inclusive to make genera in Lepidoptera and other invertebrate taxa is scientifically trivial compared with identifying monophyletic groups and building classifications. But the issue is important when making our classifications usable. In this paper, the results of a recent taxonomic review at the genus level of the Geometridae moth tribe Macariini are discussed to highlight some general issues. 1. INTRODUCTION Professor Clas NAUMANN was a prominent figure, both as the Director of the Museum Alexander Koenig, and as a lepidopterist. This paper is about the importance of taking a global approach to a taxonomic problem that is widely en- countered by taxonomists. It was with Clas NAUMANN’s strongly international approach to his work in mind that I elected to discuss this general topic. His special interest in Lepidoptera encouraged me to illustrate the generalities by a group of Geometridae moths. R. A. CROWSON bemoaned the loss of the broad Lin- naean concept of the genus (CROWSON 1970). In charac- teristically waspish mode, he suggested that had sys- tematists admitted intermediate categories between the genus and species they would have shown “a degree of public responsibility and foresight which experience shows is unrealistic to expect of ordinary human beings”. A similar approach has been adopted by other authors, who have tried to be more synthetic in their approach (e.g., DAVIS & HEYWOOD 1963, for angiosperms). With the massive increase in the discovery of species since the time of LINNAEUS, it was inevitable that species would be grouped into ever more clusters. And it was understand- able that these clusters would be given generic names, thus splitting the minimal number of genera accepted by LIN- NAEUS, his contemporaries and slightly later followers. LINNAEUS and his students described species from many parts of the world, not just their native Scandinavia, with many of the specimens collected at trading posts associated with the old shipping lines or the hinterlands of these ports. Nevertheless, their system was founded largely on species encountered in their European surroundings. Division of the genera occurred rather later: for Lepidoptera the proc- ess started notably with HUBNER (1816-[1825]) (EMMET 1991; SCOBLE 1991; HAUSMANN 2001). l In commemoration of Clas Michael Naumann zu Königsbrück (26.06.1939 - 15.02.2004) Those species of Lepidoptera described by LINNAEUS were assigned to just three genera, Papilio, Sphinx and Phalaena. With the subdivisions of Phalaena included, the total number of divisions (call them ‘genera’) was nine. Today, these divisions are treated mainly as super- families (e.g., Geometra — Geometroidea; Noctua Noctuoidea). But as large numbers of new species were discovered from outside Europe, those classifications based significantly on European species became unsatis- factory with the consequence that genera became split. New genera were sometimes described for new species with little reference to taxonomic context, particularly where taxonomists were isolated from major European collections. This problem was encountered in a study of the Macariini (SCOBLE & KRUGER 2002), a tribe of en- nomine Geometridae moths distributed worldwide but described originally from Europe, where differences were emphasized at the expense of similarities. 2. THE PROBLEM The practical problem, which is faced widely by tax- onomists working on species-rich groups, is one of how to apply the rank of genus. It is agreed widely among the taxonomic community that each genus should be monophyletic as far as that can be determined. As a sci- entific problem, the issue of applying rank to the com- ponents of a classification is trivial; the more profound question is one of identifying monophyletic groups. But in practical terms applying the rank of genus is not triv- ial, because the results of taxonomic revision are used not only by taxonomists specialising in areas close to the target taxon, but also by taxonomists working on less closely related taxa, and by a much wider user community with identification and information needs. Taxonomy should be viewed as an information science (GODFRAY 2002), and providing a usable. as well as a formalised, system is a social responsibility. As MABBERLEY (1997) wrote: “... the Gestalt of a fig is 188 Bonner zoologische Beitráge 53 (2004) Table 1: Major genus-group concepts of the Marcariini GUENEE PROUT JANSE (1932) WEHRLI FORBES HOLLOWAY SCOBLE & [1858] (1915-16) South Africa (1939-54) (1948) [1994] KRUGER (2002) Palaearctic Palaearctic New York & Borneo Global Neighbou- ring States Macaria Macaria Semiothisa Semiothisa Eumacaria Lampadopteryx Platypepla Halia Narraga s.l. s.l. Enconista Iridoplecta Milocera Isturgia (Synonyms (Subgenera: Itame Hypephyra Plateoplia Bichroma Macaria, Go- Semiothisa Semiothisa Oxymacaria Narraga Fidonia donela, Teph- Godonela Macaria Macaria Heliomata (sensu rina (in part), Thyridesia Isturgia Godonela Isturgia Treitschke) Osteodes, Neomacaria Mellilla [tame Itame Discalma, Macaria Trigrammia Chiasmia Peridela) Ligdiformia Mellilla Diastictis Chiasmia Dissomorphia Osteodes Plateoplia Diastictis Paramelora Platypepla Asmate) Eumacaria Hyostomodes Gnopharmia Milocera Bichroma Oxymacaria Narraga Semiothisa Isturgia Macaria Itame Boarmioides Tephrina Parosteodes Gnopharmia Chiasmia Malgassothisa Lampadopteryx Digrammia Hypephyra usually unmistakable but to split the genus Ficus into several on the basis of characters only revealed by lenses seems academic self-indulgence.” . . . \ 4000 The problem has arisen through a combination of two frequently observed characteristics of taxonomy. First, taxonomy is the product of a long history with a scat- tered literature. Second, because many taxonomists lack access to comparative material (specimens and litera- ture), a lack of context is often apparent. For about 100 years after LINNAEUS (1758) published the tenth edition of Systema Naturae, a relatively small number of spe- cies were described (Fig. 1) and by few taxonomists. With the explosion of exploration from the mid 19" Century there was a corresponding increase in collec- tion of specimens and in the description of species. The 3000 2000 Number of species 1000 0 : oe a E } rate of species description fell around the mid 20" Century en ieee 1850 1909 . Decade and taxonomy entered a more synthetic phase where which should encourage taxonomists to integrate further their efforts and to be prepared to synthesise. 1950 2000 species were synonymised or recombined with new gen- era. The emphasis on accepting only monophyletic taxa as valid and worthy of naming, gave an apparent intellectual basis for splitting taxa until all recognised groups were monophyletic. Currently there is a welcome emphasis on the value of taxonomy to a wider user community, Fig. 1: Graph of number of species of Geometridae described by decade since 1758 (simplified from GASTON et al. 1995). I use the results of a genus-level study (SCOBLE & KRUGER 2002) for the geometrid moth tribe Macariini to exemplify some issues faced by taxonomists classify- Malcolm J. SCOBLE: Issues in Delimiting Genera in Invertebrates: an Example from the Lepidoptera 189 ing taxa that are widely distributed. From this basis some general messages are presented. 3. THE TRIBE MACARIINI There are about 700 species of Macariini, a tribe belonging to the subfamily Ennominae of the Geometridae. The group includes medium-sized geometrid moths and is represented in all biogeographical regions. The larval foodplants of many species belong to Leguminosae (Fabaceae). The moths are encountered frequently in light traps. The tribe was described (as ‘Macaridae’) by GUENEE [1858] for four genera, two of which are now excluded from the tribe. Six prior regional works had a significant influence on the global generic classification proposed by SCOBLE & KRUGER (2002), the genera accepted in each publication being listed in Table 1. Neither PROUT (1915- 16), JANSE (1932), nor WEHRLI (1939-54) united their genera under a tribal division, but they indicated that the genera were associated. JANSE (on the South African spe- cies) included all the large macariine genera under Semio- thisa (including Macaria, which has priority), starting a tradition that has been followed to the present day by many lepidopterists understandably confused by the plethora of names and unclear generic concepts. WEHRLI (Palaearctic fauna) recognised more taxonomic structure than did JANSE, treated JANSE’s synonyms as subgenera of Semio- thisa, and added several other subgenera. FORBES (1948), in the few genera treated in his restricted area of New York and neighbouring States, included all genera with the characteristic uncus ‘horns’ in Semiothisa, noting that the genus was enormous and of world-wide distri- bution. Unfortunately both Janse and Forbes were un- aware that the type species of Semiothisa lacks ‘horns’ and has genitalia that are distinctive, the genus being exclu- sively neotropical and having 18 other members. Thus Semiothisa in their sense is equivalent to Macaria plus Chiasmia in the sense of SCOBLE & KRUGER (2002). The genera accepted by SCOBLE & KRUGER (2002) are also listed in Table 1. These authors concluded that the Macariini are not certainly defined as a monophyletic group — the boundaries are unclear. There appear to be two large, arguably monophyletic genera (Macaria and Chiasmia), and one large genus (/sturgia) that is less convincingly monophyletic. Within these genera occur several well-defined subgroups. Within the other Macariini there are a number of small and probably mo- nophyletic genera. 4. RESULTS OF THE GLOBAL GENERIC REVIEW Taxonomic treatments of the genera are found in the re- view (SCOBLE & KRUGER 2002), but for the purposes of this account a few key results are highlighted. Macariini are not defined perfectly as a tribe, but the spe- cies can be recognised by their expression of one or more of three characters: a divided valva; stout setae (“horns’) on the uncus in the male genitalia; and chaetosemata extended across the head. In many species the eighth abdominal sternum is cleft, excavated or emarginated. A further pos- sible character is the presence of a fovea, a raised cuticular patch at the base of the forewing in males largely denuded of scales, but the taxonomic value of this structure is not established. The core genera of the Macariini are properly associated, but the association of certain genera within the Tribe remains questionable. We treated many generic names as synonyms. Of the three largest genera Chiasmia has 12 synomyms, Macaria 16 and /sturgia three. These were among the earliest to be de- scribed for the tribe, and were erected for European spe- cies. Discoveries of material from other continents led to the description of new genera without detailed compari- sons being made with type species of the European genera. There are, among Macariini, species with similar wing pattern and colour that fall into quite different genera when genital morphology is compared. This is unsur- prising given that the early genera were described on ‘facies’ alone. Consequently, macariine taxonomy suf- fered both from a multiplicity of inadequately defined genera and from the erroneous generic assignment of many of the species. As a result of the confusion, many lepidopterists simply, and understandably, used one ge- nus, Macaria (or, incorrectly, Semiothisa), in which to place most macariine species. The revised classification proposed was intended to re- flect best the considerable amount of taxonomic structure apparent within the Macariini. Most species fall into one of three large genera - Chiasmia, Macaria or Isturgia. Species are included that look quite different in external appearance within the same genus, but morphological similarities suggest their association. Within these gen- era several species groupings were noted. There are a number of well-defined smaller genera of which the genus Semiothisa is one. Thus the revised classification is a compromise between placing most species in a massive single genus *Semiothisa” (in the incorrect sense of many authors) and creating numerous genera for small clusters of species or single species. The principle, however. of try- ing to recognise only monophyletic groups remains. 5. MESSAGES Taxonomists will be familiar with the issues we encoun- tered in revising the Macariini. The general points that arose are as follows. 1. Much taxonomy is focused on the fauna of a particu- lar region. While it is perfectly reasonable to undertake faunal studies, taxonomists bear a responsibility to exe- 190 Bonner zoologische Beitráge 53 (2004) cute their work in a broad context. An excellent con- temporary example of how a faunal review can be writ- ten within a strongly contextual taxonomic framework is the multi-volume treatment of the Moths of Borneo by J. D. HOLLOWAY (e.g., HOLLOWAY [1994], which in- cludes Macariini). Holloway treated the Macrolepidop- tera moths of the region to the level of species, but made extensive comparative studies across S E Asia and Aus- tralasia and, using the rich collections of the Natural History Museum, London, provided a global context. A practical problem arises in that many taxonomists work remotely from major comparative reference collections and libraries. If those of us based in such collections expect such a comparative approach from others, we shall need to pro- vide better access to the comparative information by creat- ing cross-institutional infrastructures. The Internet will be the medium through which this infrastructure is delivered. 2. It is preferable not to describe new genera for single species or small groups of species unless they fall de- monstrably outside the generic system currently in use. Classifications and names do and should change, within reason, when research shows existing systems are not optimal. However, taxonomists should be sensitive to those who use their results. We must be ever more con- scious of that wide community that uses our results and ask ourselves: “will the changes we make to classifica- tions help or hinder those who use the classifications?” With careful thought there should be no reason why we should compromise on scholarship. Where uncertainty exists as to the generic association of a species, placing inverted commas around the genus name adopted can be a helpful convention to indicate that the species should probably be excluded. This pro- tocol avoids the need to create new genera, but indicates that further study is required. For example, “Semio- thisa” nigroalbana Cassino implies that the generic placement of nigroalbana is uncertain. This simple modification avoids the complexity of discarding the binomial system as suggested, for example, by MICH- ENER (1964), yet protects the integrity of monophyletic groups. Like MICHENER, many taxonomists (myself included) have probably felt at times that we might be better off cutting the requisite nomenclatural link between genus and species. Such a break would enable species to be described without them having to be assigned to a genus when their associations remained unknown. The advantage of the proposal would obviate the need either to associate each species with an existing genus or to describe a new genus for every unassociated species. However, the cost of abandoning the binomial system of nomenclature would be heavy. Loss of a standard that has served taxon- omy so well for 250 years is likely to lead to confusion. Moreover, with increasing computerisation of taxo- nomic information the established link between genus and species is one that is embedded in many databases. 3. Taxonomists frequently encounter the situation where a large genus includes a number of demonstrably mono- phyletic subgroups and also many single species unasso- ciated with any others. The use of informal species-groups within a genus can be helpful to indicate taxonomic structure within such a genus without the need to describe a new genus for each subgroup or for each unassociated species. In the macariine exemplar, several subgroups, often closely related species from the same area, are evident in the three large genera, Macaria, Chiasmia and Isturgia, alongside numerous unassociated single species. 4. In nearly all higher taxa we find genera for which the monophyly is convincingly well founded and many genera where it is not. An argument for splitting a genus that is not demonstratively monophyletic into better-established monophyletic genera, and as many further genera as there are unassociated species, is that the situation might best reflect the natural pattern. However, taxonomists would do well to follow this course of action only where the original genus is demonstrably not monophyletic. 5. The taxonomic history of a group helps one under- stand problems of classification. Bacterial (prokaryote) taxonomy was re-based in 1980 (in effect, a new Sys- tema Naturae was created for bacteria) because so many type cultures had been lost that few names could be ref- erenced to specimens (SNEATH 1986). But such is not the case for Lepidoptera taxonomy, where types dating from the time of LINNAEUS still exist. Taxon concepts often have a long history, and there seems little to be gained and much to be lost by discarding the insights and information associated with those concepts (e.g., PULLAN et al. 2000; YTOW et al. 2001; BERENDSOHN 2003). Therefore, any review of genera (or any other taxonomic rank) in Lepidoptera (and most other eu- karyote organisms) requires that we examine the differ- ent systems through time. Proposing new classifications without understanding how previous taxonomists came to their conclusions leads to poor taxonomy. 6. CONCLUSION The description of all species of organisms is a worthy goal, but an immense undertaking, particularly when microorganisms are brought into consideration. Indeed description is not a single process, but requires both recog- nition that a putatively new species is actually undescribed and also its assignment to the appropriate genus - the generic name being a part of the name of the species. To say that a species should not be described in a taxonomic vacuum may sound like stating the obvious, yet the number of new species described within an inadequate generic framework is significant. Material advances in our Malcolm J. SCOBLE: Issues in Delimiting Genera in Invertebrates: an Example from the Lepidoptera 19] understanding of Lepidoptera biodiversity can be made more rapidly by focusing on generic reviews on a global or large regional basis, as has been done over the last decade on Geometridae (e.g., PITKIN 1996, 2002; SCOBLE 1995) including the work on the Macariini discussed in this paper. An advantage of this approach is that it accelerates coverage of Lepidoptera biodiversity while at the same time improving the quality of the taxonomic framework. Such studies will better equip those taxonomists with access to collections that are regional in scope to under- take work on species within their geographical domain. A further reason for good quality taxonomy of higher taxa was expressed by WILLIAMS & GASTON (1994) who discussed the use of higher taxa as surrogates for biodiver- sity assessment (for summary see http://www.nhm.ac.uk/ science/projects/worldmap/refs/key.htm/fkey4). They con- cluded that “With careful choice of higher-taxon rank, it may be possible to re-deploy effort from taxonomically intensive to taxonomically extensive surveys, in order to estimate the global distribution of a much larger propor- tion of overall biodiversity at the same cost”. Acknowledgements. I thank Linda Pitkin, Natural History Museum, London, for reading a draft of this manuscript and for her collaboration on Geometridae taxonomy over the years. REFERENCES BERENDSOHN, W. G. (Ed.) (2003): MoReTax: handling factual information linked to taxonomic concepts in biology. Federal Agency for Nature Conservation: Bonn. Crowson, R. A. (1970): Classification and Biology. Heinemann, London. Davis, P. H. & HEYWOOD, V. H. (1963): Principles of angio- sperm taxonomy. Oliver and Boyd, Edinburgh; London. EMMET, A. M. (1991): The scientific names of the British Lepidoptera. Harley Books, Colchester. FORBES, W. T. M. (1948): The Lepidoptera of New York and Neighboring States 2. Memoirs of the Cornell University agricultural Experimental Station, No. 274. GASTON, K. J., SCOBLE, M. J. & CROOK, A. (1995): Pat- terns in species description: a case study using the Geometridae (Lepidoptera). Biological Journal of the Linnean Society 55: 225-237, GODFRAY, H. C. J. (2002): Challenges for taxonomy. Na- ture 417: 17-19. GUENEE, A. (1858 [dated 1857]): Uranides et Phalénites 1. Pp. [i]-Ivi, 1-514 in: BOISDUVAL, M & GUENEE, A. (eds). Histoire Naturelle des Insectes, Species Général des Lépidopteres 9. HAUSMANN, A. (2001). The Geometrid Moths of Europe, 1. 282 pp., Apollo Books, Stenstrup. HOLLOWAY, J. D. ([1994], dated 1993). The Moths of Borneo family Geometridae, subfamily Ennominae; part 11. Ma- layan Nature Journal 47: 1-309, 593 figs, 19 col. pls HUBNER, J. (1816-[1825]): Verzeichniss bekannter Schmett- Linge [sic]. Augsburg. JANSE, A. J. T. (1932): Sematuridae and Geometridae. The Moths of South Africa 1. Pretoria. ! LINNAEUS, C. (1758): Systema Naturae, 10" ed. 824 pp., Laurentii Salvi1, Holmiae. MABBERLEY, D. J. (1997): The Plant-Book. 2" ed. Cam- bridge University Press, Cambridge, UK. MICHENER, C. (1964): The possible use of uninominal no- menclature to increase the stability of names in biol- ogy. Systematic Zoology 13: 182-190, PITKIN, L. M. (1996): Neotropical Emerald moths: a review of the genera (Lepidoptera: Geometridae, Geometrinae). Zoological Journal of the Linnean Society 118: 309-440. PITKIN, L. M. (2002): Neotropical ennomine moths: a re- view of the genera (Lepidoptera: Geometridae). Zoo- logical Journal of the Linnean Society 135: 121-401. PULLAN, M. R., WATSON, M. F., KENNEDY, J. B., RAGUE- NAUD, C. & HYAM, R. (2000): The Prometheus Taxo- nomic Model: a practical approach to representing multiple classifications. Taxon 49: 55-75. PROUT (1915-16): Geometrinae. Pp. 303-413 in: SEITZ, A. (ed.), pp. 303-413, pls. 14-25. The Macrolepidoptera of the World 4, The Palaearctic Geometrae. Alfred Kernen, Stuttgart. SCOBLE, M. J. (1991): Classification of the Lepidoptera. Pp. 11-45 in: EMMET, A. M. & HEATH, J. (eds) The Moths and Butterflies of Great Britain and Ireland. 7(2). Harley Books, Colchester. SCOBLE, M. J. (1995): A review of the moth tribe Palyadini with the description of a new genus (Geometridae: Ennominae). Systematic Entomology 20: 35-58. SCOBLE, M. J. & KRUGER, M. (2002): A review of the gen- era of Macaruni with a revised classification of the tribe (Geometridae: Ennominae). Zoological Journal of the Linnean Society 134: 257-315. SNEATH, P. H. A. (1986): Nomenclature of bacteria. Pp. 36-47 in: RIDE, W. D. L. & YOUNES, T. (eds) Biologi- cal Nomenclature Today. IRL Press: Oxford. WEHRLI, E. (1939-54). 6. Die Spanner des Palaearktischen Faunengebietes. Subfamilie: Geometrinae, in: SEITZ. A. (ed). Die Gross-Schmetterlinge der Erde 4 (Suppl). 254-766, pls 1-53, Alfred Kernen, Stuttgart. WILLIAMS, P. & GASTON, K. J. (1994). Measuring more of biodiversity: can higher-taxon richness predict wholesale species richness? Biological Conservation 67: 211-217. YTOW, N., Morse, D. R. & ROBERTS, D. McL. (2001): No- mencurator: a nomenclatural history model to handle mul- tiple taxonomic views. Biological Journal of the Linnean Society 73: 81-98. (DOI:10.1006/bjls.200 1.0527) Author’s address: Dr Malcolm J. SCOBLE, Department of Entomology, The Natural History Museum; Cromwell Road. London SW7 5BD, UK. E-mail: m.scoble@nhm.ac.uk ur f 1 1 " . y o e ni 7 u 7 4 f 7 : wo je . de ‘ vn 7 > v it ‘ : ra 7 y E a . m . \ N PN) 4 3 7 N ' | | y i i 4 5 A Ñ a ' i . : : ‘ 7 te t ‘ , : y whe od : 7 ñ : an ‘ + ’ . . u ‘ \ y \ , ‘ 4 ad ‘ ll I ' i e * ‘ 7 \ iv e i. 1 5 “ i - y 7 \ u sen 7 A , y ‘ \ " MN + A , te Bonner zoologische Beiträge Band 53 (2004) Heft 1/2 Seiten 193-209 Bonn, Juni 2005 Revision of the Trichoclinocera dasyscutellum Group from East Asia (Diptera: Empididae: Clinocerinae) A Bradley J. SINCLAIR” & Toyohei SAIGUSA” “ Zoologisches Forschungsmuseum Alexander Koenig, Bonn, Germany “Fukuoka, Japan Abstract. The Trichoclinocera dasyscutellum group is revised and includes the following eight species: 7. asiatica sp. nov., 7. dasyscutellum (Saigusa), T. feuerborni (Engel) comb. nov., 7. fluviatilis (Brunetti), 7. naumanni sp. nov., 7 nepalensis sp. nov., T. taiwanensis sp. nov., and 7. yunnana sp. nov. This group is restricted to East Asia, ranging from Nepal to Japan and as far south as Java. Male genitalia are illustrated and a key to species is provided. Clinocera ve- lutina (Engel), originally published together with 7. fewerborni, is also redescribed. Lectotypes are designated for both species. Key words: Taxonomy, aquatic, new species, morphology, genitalia 1. INTRODUCTION The genus Trichoclinocera Collin is among the most commonly collected clinocerines in North America and Japan. They are normally encountered on emergent rocks in large streams and rivers, usually where the ri- parian vegetation is open overhead (SINCLAIR 1994). Adults fly close to the water surface in search of prey and consequently are difficult to collect by sweep net and rarely taken in malaise traps. They are most readily collected by use of an aspirator, sucking up individual specimens that are found sitting on emergent rocks in the stream. There are currently 24 described species found in the Nearctic, Palearctic and Oriental Regions (SINCLAIR 1994, 1995), plus a recently described species from In- dia (WAGNER et al. 2004). The genus was thoroughly redescribed and divided into six species-groups by SIN- CLAIR (1994). The 7. dasyscutellum group, the focus of this paper, is considered the sister clade to the western North American 7. comata group on the basis of setulae on the scutellum. The recently described species, 7. ser- rata Wagner & Leese, 2004 lacks this feature and can not be assigned to the 7. dasyscutellum group, and is most closely related to 7. cyanescens Vaillant, 1960. Numerous congeneric species from East Asia await de- scription and this paper is the first of a series of studies of this genus from this region. This study 1s dedicated in the memory of our colleague and fellow entomologist. Dr. Clas M. NAUMANN. 1 In commemoration of Clas Michael Naumann zu Königsbrück (26.06.1939 — 15.02.2004) 2. MATERIALS AND METHODS This study is based on some 1600 specimens borrowed from or deposited in the following institutions: Biosys- tematics Laboratory, Kyushu University, Fukuoka, Ja- pan (BLKU); The Natural History Museum, London. England (BMNH); Bernice P. Bishop Museum, Hono- lulu, USA (BPBM); Canadian National Insect Collec- tion, Ottawa, Canada (CNC); Hungarian Natural History Museum, Budapest (HNHM); Faculty of Agriculture, Hokkaido University, Sapporo, Japan (HUS): Staat- liches Museum fur Naturkunde, Stuttgart, Germany (SMNS); Zoologisches Forschungsinstitut und Museum Alexander Koenig, Bonn, Germany (ZFMK); Zoo- logical Survey of India, New Alipore, Calcutta, India (ZSD. Terms and methods used in specimen preparation fol- low that of SINCLAIR (1994, 1995). The use of a very mild acid (e.g., lactic acid) for clearing and studying the genitalia of male Trichoclincera is very important. Strong acids such as KOH often destroy the delicate membranous tip of the phallus. The following abbrevia- tions are used in the description: acr - acrostichal setu- lae; de - dorsocentral bristles; dm - discal medial: npl - notopleural bristles; pal - postalar bristles: ph - post- humeral bristle: pprn - postpronotal bristles: presut spal - presutural supar-alar bristles: psut spal - postsutural supra-alar bristles; R - radial vein; sctl - scutellar bris- tles; T - tergite. Also the following abbreviations are used for the most frequent collectors’ names: J. Emoto - JE; R. Matsumoto - RM; H. Shima - HS: B.J. Sinclair - BJS; T. Saigusa - TS. 194 Bonner zoologische Beitráge 53 (2004) 3. TAXONOMY 3.1. Trichoclinocera dasyscutellum group This species group is distinguished from other species of Trichoclinocera by the following suite of characters: long, slender legs; pronounced rows of stout ventral bristles on the fore femur absent; normally only spine- like preapical bristles present on femora (Figs. 22-27); lacking anterodorsal and posterodorsal leg bristles; scutellum with setulae on disc; face generally flat, with small apical notch (Fig. 19); cell dm produced apically, not truncate; surstylus normally with scolex-like apex; phallus with short, elaborate membranous apical lobes. 3.2. Key to species of the Trichoclinocera dasyscutellum group 1 Dorsocentral bristles longer than ocellar bristles or two-thirds length of antennal arista-like style........... 2 — Dorsocentral bristles shorter than ocellar bristles or one-half length of antennal arista-like style.............. 3 [59] Hind tibia with posterodorsal spur-like bristle one- third from base; one slender lower notopleural bris- tle; clasping cercus tapered and narrow apically (Fig. 2 T. dasyscutellum (Saigusa) — Hind tibia without spur-like bristle; 2-3 stout lower notopleural bristles; clasping cercus rounded apically (Fig. 1) (South Korea, Russia: Primorsk Territory) ......... e reece eons eee T. asiatica sp. nov. 3 Pleura brown, lacking distinct blue pruinescence....... 4 — Pleura with distinct blue pruinescence..............: 6 4 Radial vein with 3-5 erect bristles proximal to origin of Rs, ca. one-half length of basal costal bristle, in addition to setulae (Java) ...... T. feuerborni (Engel) comb. nov. — Radial vein with setulae only, lacking long erect BASES a inca 5 5 Male fore femur with two black, spur-like anteroven- tral subapical bristles and posteroventral row of nu- merous stout bristles, those on basal half nearly sub- equal to width of femur; clasping cercus strongly hooked and tapered; surstylus with scolex-like apex (Fig. 6); female fore femur with 3-4 distinct spur-like subapical bristles (China).......... T. naumanni sp. nov. — Male fore femur with one black, spur-like anteroven- tral subapical bristle and a sparse posteroventral row of pale hairs, those on basal half less than width of femur (Fig. 25); clasping cercus finger-like, slightly curved; surstylus with subapical pointed projection and stout apical bristle; female fore femur with one distinct spur-like subapical bristle (Fig. 8) (Nepal, WICHMANN). cdcmescnudecsensoveatedeandvates T. nepalensis sp. nov. 6 Base of stalk of halter brown, not brightly coloured: anterior apex of phallus with long, slender pointed membranous sac (Fig. 10) (Taiwan)... sakes Se datgout re T. taiwanensis sp. nov. — Base of stalk of halter bright orange-brown; anterior apex of phallus with short, broad membranous sac (females difficult to distinguish beyond this couplet) 7 Clasping cercus straight beyond base, tapered only at extreme apex; length of surstylus less than half length of cercus with narrow angular basal prolonga- tion (Fig. 5) (India, Nepal, China: Yunnan)................ Bie aorta setae Sancta ee T. fluviatilis (Brunetti) — Clasping cercus arched throughout length, apical fourth tapered to slender apex; length of surstylus more than half length of cercus with rounded (Fig. 11) or broad angular basal prolongation (China, o wesen T. yunnana sp. nov. 3.3. Trichoclinocera asiatica sp. nov. (Figs. 1, 7, 22) Type Material. Holotype male labelled: “USSR: Pri- mor’ye/ 17km SSW of/ Krounovka, 140m/ 29.vii.1990/ T. Saigusa leg.” (BLKU). Paratypes: RUSSIA: 141 4, 299 9, 28-29.vii.1991, same data as holotype (BLKU); 23 $, 58 Y, Primorskij Kraj [Primorsk Terr.], SW Krounovka, nr. Mt Med- vezh’ja, 8-9.vii.1993, T. Yasunaga (BLKU); 2 ¢, 9 9, Primorskij Kraj, SW Krounovka, nr. Mt Medvezh’ja, 4- 6.v11.1993, T. Nakamura (BLKU). SOUTH KOREA: 1 S, Samjeong, Hwagae-Myeon, 600-800 m, 8.vi.1991, TS (BLKU). Several specimens deposited in ZFMK. Recognition. This species is distinguished by its yel- lowish coxae, long dorsocentral bristles and 2-3 stout, lower notopleural bristles. Description. Wing length 3.6-4.2 mm Male. Brown in ground colour. Head rounded; com- pound eye large, rounded. Face narrowest at middle, ca. width of antennal sockets, with bright blue pruines- cence, lower margin of face nearly straight (clypeus) lacking pronounced carina or notch; width of gena one- sixth height of eye. Anterior margin of ocellar triangle, lateral postocciput and gena with blue pruinescence; vertex and upper part of postocciput dark when viewed dorsally. Ocellar bristles longest among cranial bristles; occiput with 3 pairs of bristles posterior to ocellar trian- gle, ca. one-half length of ocellar bristles; postocular with row of short bristles along margin of eye, upper 3- 4 bristles darker and more stout than lower bristles. Arista-like stylus short and slender, slightly tapered api- cally. Palpus slender, length less than one-half height of eye. Bradley J. SINCLAIR & Toyohei SAIGUSA: Trichoclinocera dasyscutellum Group from East Asia 195 cl cerc Figs. 1-6: Terminalia of Trichoclinocera dasyscutellum group. (1) T. asiatica sp. nov., male; sp. nov., male. Scale bar = 0.1 mm. Abbreviations: cere - cercus: cl cere - clasping cercus; ej apod - ejaculatory epandrium; hypd - hypandrium; ph - phallus; sur - surstylus; T - tergite. (2) 7. dasyscutellum (Saigusa), male; (3) 7. dasyscutellum (Saigusa), female; (4) 7. feuerborni (Engel), male: (5) T. Aluviatilis (Brunetti), male: (6) 7. naumanmn apodeme: epand - 196 Bonner zoologische Beitráge 53 (2004) Pleura and lateral margin of scutum, including post- pronotal lobe and notopleuron entirely clothed in blue pruinescence; posterolateral corner of scutum near postalar ridge somewhat paler. Thoracic bristles long, some longer than width of eye; several acr anterior to first de; 5 pairs de; | pprn; | presut spal; 1 upper, 2-3 stout lower npl; 1 psut spal; 1 pal; 2 pairs sctl and nu- merous pale setulae on disc; several pale bristles on notopleural depression. Laterotergite with patch of pale bristles; prothoracic episternum, katepisternum, and hind margins of metepisternum with long, pale hairs. Wing narrow, infuscate; narrow, faint, parallel-sided stigma present. One long, basal costal bristle, subequal to length of posterior de. Row of setulae from base of R to tip of Ry; cell dm strongly produced apically; Ry and Rs divergent apically. Halter dark with pale base. Legs with femora pale brown beneath and basally, be- coming darker toward apex. Coxae with long pale, mar- ginal hairs; fore coxa yellowish with blue pruinescence on basal half; mid and hind coxae somewhat pale with blue pruinescence. Fore femur with row of 6-7 slender anterior bristles on apical third (Fig. 22), dorsal to spine-like bristles; antero- and posteroventral row of pale, slender bristles along entire length, posteroventral setae subequal to width of femur, twice length of an- teroventral bristles; 3 black, very short, peg-like an- teroventral bristles on apical third; anterodorsal region with row of 5-6 stout, black bristles on basal third. Fore tibia with biserial row of short, stout bristles beneath. Mid femur with 3 pairs of black, spine-like antero- and posteroventral preapical bristles, nearly as long as one- half width of femur; anterior margin with long, slender bristles along entire length, longer than width of femur; lacking erect antero- and posterodorsal bristles. Hind femur with 2-3 pairs of antero- and posteroventral preapical bristles, less stout than preapical bristles of mid femur; lacking antero- and posterodorsal bristles. Hind tibia with 3-4 erect, spine-like dorsal and ventral bristles on apical third, longer than width of tibia. Tarsi longer than corresponding tibiae. Empodium pulvilli- form, subequal in length to claw, less than one-half length of tarsomere. Hypandrium (Fig. 1) long and cylindrical, longer than epandrium. Phallus long, distinctly bent sub-basally; apex with slender, curved anterior process and pair of straight, short, rod-like posterior processes; ejaculatory apodeme strongly recurved anteriorly. Surstylus very slender, somewhat twisted and arched medially, with membranous oval scolex-like apex. Clasping cercus slender, finger-like, parallel-sided, with rounded apex; row of stout, peg-like bristles on inner dorsal margin. Female. Similar to male except as follows: stout an- teroventral bristles on apical third of fore femur longer and stouter. Terminalia very similar to 7. dasyscutellum: sclerites of segment 8 narrowly separated laterally; T8 with posterior row of long and short bristles, length of dorsal bristles subequal to length of sclerite; T10 with stout, curved bristles on posterior half, with 3-4 apical bristles more stout than remaining bristles; anteroventral corner of T10 not narrowed and prolonged. Geographical Distribution and Seasonal Occurrence. This species occurs in the far east of Asia, specifically the Primorsk Territory of Russia and the Korean Penin- sula (Fig. 7). Remarks. See under 7. dasyscutellum. Etymology. The specific name refers to the distribution of this species along the far eastern region of mainland Asia. 3.4. Trichoclinocera dasyscutellum (Saigusa) (Figs. 2, 3, 7, 13, 18-20, 23) Acanthoclinocera dasyscutellum Saigusa, 1965: 54. Trichoclinocera dasyscutellum: SINCLAIR 1994: 1015. Type Material. Holotype, male labelled: “[HONSHU]/ Kanayama/ YAMANASHI/ 19.VIII.1962/ T. SAI- GUSA” (BLKU). Paratypes: 2 4, 1 Y, same data as holotype (CNC); 21 3,17 Y, same data as holotype (BLKU). Additional Material. (375 specimens) JAPAN: Akita: Kuroyu, Hachimantai, 8.v11.1962 (BLKU). Nyntosan, Hachimantai, 9.vii.1962 (BLKU). Aomori: Oirase- keikoku nr. Tawadoko, 14.v11.1962 (BLKU). Gifu: Futatsuya, Kawai-mura, 11.v1.1977 (BLKU). Iwate: Hayachinesan, 800-1000 m, 24.v11.1973 (BLKU); Hata- tine, 4.vili.1964 (HUS). Nagano: Inakoyu, 14.v111.1975 (BLKU); Inakoyu, Kitayatsugatake, 14.v111.1975 (BLKU); Iwanadome, Shimashima, 13.vii.1963 (BLKU); Tobi- rakósen, Utsugushigaharn, 18.v11.1963 (BLKU). Ya- manashi: Hirogawara, Kitadake, 18-19.v111.1978 (BLKU); Kanayama, Kinpusan, 1.v11.1963 (BLKU); Kanayama, Sudama, 10-11.viii.1961, 6.v11.1962, 30.vi., 24.viii.1963, 12-13.viil.1973, 20,30.vi., 2-31.vii., 1,16.1x.1975 (BLKU); Kanayama, Kanayamotöge, 2.vii.1963 (BLKU); Motodanigawa, Kanayama, 9.vii.1975 (BLKU); Tokusatóge, Sudama, 23.v1.1975 (BLKU). Several specimens deposited in BMNH and ZFMK. Recognition. This species is readily distinguished from other species of the 7. dasyscutellum group by the red- dish-brown coloured legs, long dorsocentral bristles and spur-like bristle on the hind tibia. Description. Wing length 3.6-4.2 mm. Male. Dark brown to black in ground colour. Head rounded; compound eye large, rounded. Face narrowest at middle, ca. width of antennal sockets, with pale blue Bradley J. SINCLAIR & Toyohei SAIGUSA: Trichoclinocera dasyscutellum Group from East Asia 197 pruinescence, lower margin of face nearly straight, lack- ing pronounced carina or notch (Fig. 19); width of gena one-sixth height of eye. Ocellar bristles longest among cranial bristles; occiput with 3 pairs of bristles posterior to ocellar triangle; postocular with row of short bristles along margin of eye, upper 3-4 bristles darker and more stout than lower bristles. Arista-like stylus long and slender, tapered apically. Palpus slender, length less than one-half height of eye. Pleura entirely clothed in blue pruinescence; poster- olateral corner of scutum near postalar ridge pale brown. Thoracic bristles long, some longer than width of eye; several acr anterior to first de; 5-6 pairs de; | pprn; | presut spal; 2 npl; 2 psut spal; | pal; 2 pairs sctl and numerous pale setulae on disc; several pale bristles on notopleural depression. Laterotergite with patch of pale bristles; prothoracic episternum, katepisternum, and hind margins of metepisternum with long, pale hairs. Wing (Fig. 13) narrow, infuscate; narrow, faint, paral- lel-sided stigma present. One long, basal costal bristle, subequal to length of posterior de; often 1 proximal cos- tal bristle, shorter than one-half length of basal bristle. Row of setulae from base of R to tip of Ry; cell dm strongly produced apically; Ry and Rs slightly divergent. Halter brown with pale base. Legs reddish-brown, darker toward apex. Coxae with long pale, marginal hairs; mid and hind coxae with tinge of pale pruinescence. Fore femur with 4-5 slender ante- rior bristles on apical third, dorsal to spine-like bristles (Fig. 23); antero- and posteroventral row of long, pale slender bristles along entire length, slightly longer than width of femur; 4-5 black, spine-like anteroventral bris- tles on apical fourth; anterodorsal region with row of short, stout, black bristles on basal third. Fore tibia with anteroventral row of short, stout bristles and posteroven- tral row of slender bristles. Mid femur with pair of black, spine-like antero- and posteroventral preapical bristles, longer and more slender than corresponding bristles of fore femur; lacking erect antero- and postero- dorsal bristles. Hind femur with 1-2 stout anteroventral preapical bristles, similar to mid femur; lacking antero- and posterodorsal bristles. Hind tibia with posterodorsal spur-like bristle one-third from base, slightly longer than width of tibia; 2-3 posterodorsal preapical spine- like bristles, longer than width of tibia. Tarsi longer than corresponding tibiae. Empodium pulvilliform, sub- equal in length to claw, less than one-half length of tar- somere. Hypandrium (Fig. 2) long and cylindrical, longer than epandrium. Phallus long, distinctly bent sub-basally: apex with cup-shaped process; ejaculatory apodeme gently recurved anteriorly. Surstylus very slender, somewhat twisted and arched medially, with membra- nous oval scolex-like apex. Clasping cercus slender, finger-like, bearing stout bristles on inner dorsal margin Female. Similar to male except as follows: fore tibia lacking modified bristles beneath. Terminalia: sclerites of segment 8 narrowly separated laterally; T8 with pos- terior row of long bristles, subequal to length of sclerite (Fig. 3); TIO with stout, curved bristles along inner and posterior margins, with 3-4 apical bristles more stout than remaining bristles; anteroventral corner of TIO not strongly narrowed and prolonged. Geographical Distribution and Seasonal Occurrence. This species is restricted to the Japanese island of Hon- shu (Fig. 7), with flight records from mid-June to mid- September. Remarks. This species was originally assigned to the genus Acanthoclinocera Saigusa, but was transferred to Trichoclinocera when the former was considered to be congeneric (SINCLAIR 1994). Trichoclincera dasyscutel- lum is most closely related phylogenetically to 7. asiat- ica on the basis of the long thoracic bristles and simi- larities of the clasping cercus and apex of the phallus. 3.5. Trichoclinocera feuerborni (Engel) comb. nov. (Fig. 4) Clinocera (s.s) feuerborni Engel, 1931: 487. Type Material. Lectotype male (here designated) la- belled: “FS 2a. 6/XII”; “Ngatiloga-Fall/ b. Sarangan. M. Java./ 6.X11.28. (Feuerborn.) [hand-written]”: “[red la- bel, no writing]”; “Y Clin./ Feuerborni n.sp./ det. Engel” (SMNS). A lectotype label “LECTOTYPE/ of Cli- nocera/ feuerborni Engel/ des. B.J. Sinclair 2004 [red label]” has been attached to this specimen. Paralectotype: Same data as lectotype, with a yellow paralectotype label attached (1 SMNS) [abdomen glued to thorax]. ENGEL (1931) writes that the two type specimens were stored in alcohol, but it must have been dried by him subsequently. ENGEL (1931) also described the species Kowarzia velutina (also based on two specimens) in the same publication (see Appendix), and given that there were four specimens all bearing red labels (without writing) stored together under the label C/inocera feuerborni, it appears that the type material of these two species were subsequently mixed. Consequently lecto- types are designated for both species at this time to clearly distinguish the type material of these two spe- cies. Recognition. This species is distinguished from other species of the 7. dasyscutellum group by its brown ground colour and long bristles on the radial vein. Description. Wing length 4.0-4.1 mm. 198 Bonner zoologische Beitráge 53 (2004) Fig. 7: Known distribution of species of Trichoclinocera dasyscutellum group. T. asiatica sp. nov. (MW), 7. dasyscutellum (Sai- gusa) (9), 7. taiwanensis sp. nov. (0). Male. Dark brown in ground colour. Head rounded; compound eyes large, rounded. Face narrowest at mid- dle, ca. width of antennal sockets, with pale blue pru- inescence on lower half, lower margin of face with slightly pronounced carina or notch; width of gena one- fifth height of eye. Ocellar bristles longest among cra- nial bristles; occiput with 3 pairs of bristles posterior to ocellar triangle; postocular with row of short bristles along margin of eye, upper 3-4 bristles darker and more stout than lower bristles. Arista-like stylus long and slender, only slightly tapered apically. Palpus slender, length less than one-half height of eye. Pleura, postpronotal lobe and notopleural depression en- tirely clothed in pale blue pruinescence. Thoracic bris- tles well developed, all shorter than width of eye; acr absent; 5 pairs de; 1 pprn; 1 presut spal; 2 npl; 2 psut spal; 1 pal; 2 pairs sctl and numerous setulae on disc; several pale bristles on notopleural depression. Latero- tergite with patch of pale bristles; prothoracic epister- num, katepisternum, and hind margins of metepisternum lacking long pale hairs. Wing narrow, infuscate; narrow, faint, parallel-sided stigma present. One long, basal costal bristle, subequal to length of sctl. Row of setulae from base of R to tip of R,, with 3-5 outstanding bristles proximal to origin of Rs ca. one-half length of basal costal bristle; cell dm long, strongly produced apically; Ry and Rs very slightly divergent. Halter brown with pale base. Legs pale brown, darker toward apex. Coxae with long pale, marginal hairs; mid and hind coxae with tinge of pale pruinescence. Fore femur lacking anterior bristles and spine-like bristles; antero- and posteroventral mar- gins with row of long, pale slender bristles along entire Bradley J. SINCLAIR & Toyohei SAIGUSA: Trichoclinocera dasyscutellum Group from East Asia 199 length, shorter than width of femur; posteroventral base with | long bristle, ca. twice width of femur. Fore tibia with antero- and posteroventral row of short stout bris- tles. Mid femur lacking spine-like preapical bristles, all outstanding erect bristles lacking, except several slender preapical anteroventral bristles. Hind femur with 1-2 slender anteroventral preapical bristles, similar to mid femur; lacking antero- and posterodorsal bristles. Hind tibia lacking posterodorsal spur-like bristle; 2-3 erect anteroventral preapical bristles, longer than width of tibia. Tarsi longer than corresponding tibiae. Empodium pulvilliform, subequal in length to claw, less than one- half length of tarsomere. Hypandrium (Fig. 4) long and cylindrical, longer than epandrium. Phallus long, distinctly bent sub-basally; apex with narrow lateral membranous ridges; ejacula- tory apodeme gently recurved anteriorly. Surstylus ta- pered, with narrow membranous scolex-like apex. Clasping cercus strongly arched at base, finger-like, ta- pered apically, lacking stout bristles on inner margin. Female. Similar to male except as follows: ventral bris- tles of fore femur not as numerous; apical fourth with 6- 8 spine-like anteroventral spine-like bristles; fore tibia lacking modified bristles beneath. Terminalia (not dis- sected): sclerites of segment 8 narrowly separated later- ally; T8 with posterior row of long bristles. T10 with stout curved bristles along inner and posterior margins. Geographical Distribution and Seasonal Occurrence. This is the first known occurrence of this genus from Indonesia. The type specimens were collected at a wa- terfalls in middle Java (ENGEL 1931). Remarks: This species was listed under Clinocera Meigen (unplaced to species-group) by SINCLAIR (1995), but syntypes had not been examined. 3.6. Trichoclinocera fluviatilis (Brunetti) (Figs. 5, 12, 14, 24) Clinocera fluviatilis Brunetti, 1913: 34. Trichoclinocera fluviatilis: SINCLAIR 1994: 1016. T. fluviatilis: WAGNER et al. 2004: 25. Type Material Examined. Holotype, male labelled: “Clinocera/ fluviatilis’ Brun. Typ ©” [hand written]: “[India; Uttar Pradesh] NEAR BHOWALI./ KUMAON: 5,700 FT/ A.D. IMMS 1910/ 12th June. On stones in stream”; “Clinocera fluviatilis/ m ©” [hand written]; “TYPE”; “1422/ HI” [terminalia partially damaged, only midlegs remain] (ZSI). Additional Material. (101 specimens) CHINA: Yun- nan: Dali, Daboping, 1900-2200 m, 7kmW Xiaguan, 16.vi11.1995 (BLKU); Jingping, Tatyangzhai, 1500- 1600 m, 27.v.1996 (BLKU); Pingbian, Xinxinghe, 780 m, S Pingbian, 3.11.1995 (BLKU). INDIA: Himachal Pradesh: Punj-pul Nullah, Dalhousie, 6500', 8.v.1927 (BMNH). NEPAL: Chiliwa, 1350 m, 27° 29'N, 87 47'E, Shibku, 2100m, 27° 30'N, 87° 41'E, 7.vi.1972 (BLKU); Handhurung, 800 m, 27° 23'N, 87° 40'E, Linba, 1200 m, 27° 24'N, 87° 38'E, 27.vi.1972 (BLKU): Jilikinpthi, 1850 m, 27° 09'N, 87° 30'E, Pontak, 1800 m. 27° 13'N, 87° 33'E, 13.v.1972 (BLKU); Kathmandu, Godavari, 6000', 22.vii.1967 (CNC); Lelep, 1770 m, 27 31'N, 87° 47'E, 1-4.vi.1972 (BLKU): Lelep, 1770 m, 27 31'N, 87° 47'E, Chiliwa, 1350 m, 27° 29'N, 87° 47'E, 7.v1.1972 (BLKU); Methirum, 1000 m, 27° 24'N, 87 42'E, Tiwa, 1400 m, 27° 29'N, 87° 46'E, 29.v.1972 (BLKU); Naudhara - Bhadaue, nr. Pokhara, 1400-1600 m, 9.xi.1971 (BLKU); Shibku, 2100 m, 27° 30'N, 87° 41'E, Shawa, 2300 m, 27° 29'N, 87° 39'E, 8.v1.1972 (BLKU); Siwapuri, 1640 m, 22.v.1981 (BLKU). Sev- eral specimens deposited in ZFMK. Recognition. This species is distinguished from other species of the 7. dasyscutellum group by its blue pru- inescence and relatively straight clasping cercus. Description. Wing length 3.6-4.3 mm. Male. Dark brown with bright blue pruinescence. Head rounded; compound eye large, rounded. Face narrowest at middle, ca. width of antennal sockets, with bright blue pruinescence, lower margin of face nearly straight, lacking pronounced carina or notch; width of gena one- sixth height of eye. Anterior margin of ocellar triangle pale brown, lower postocciput and gena with blue pru- inescence; vertex and upper part of postocciput dark when viewed dorsally. Ocellar bristles longest among cranial bristles; occiput with 3 pairs of bristles posterior to ocellar triangle; postocular with row of short bristles along margin of eye, upper 3-4 bristles dark, stouter than lower bristles. Arista-like stylus long and slender. slightly tapered apically. Palpus somewhat clavate, length less than one-half height of eye. Pleura, postpronotal lobe, notopleuron entirely clothed in bright blue pruinescence; posterolateral corner of scu- tum including postalar ridge pale brown. Thoracic bris- tles short, shorter than one-half width of eye: acr lack- ing; 5 pairs slender de; 1 slender pprn: | presut spal: 2 stout npl; 2 psut spal; I very short, slender pal: 2 pairs sctl and several dark marginal setulae; several pale setu- lae on notopleural depression. Laterotergite with patch of pale bristles; prothoracic episternum with pale hairs: katepisternum and hind margins of metepisternum lack- ing hairs. Wing (Fig. 14) narrow, infuscate: narrow, faint. ellipti- cal, stigma present. One basal costal bristle. nearly twice length of posterior de. Row of setulae from base of R to tip of R;: cell dm strongly produced apically; Ry 200 Bonner zoologische Beitráge 53 (2004) 11 Figs. 8-11: Terminalia of Trichoclinocera dasyscutellum group. (8) T. nepalensis sp. nov., male; (9) T. taiwanensis sp. nov., fe- male; (10) 7. taiwanensis sp. nov., male with phallus partially and totally extended; (11) 7. yunnana sp. nov., male with inner apical rod retracted. Scale bar = 0.1 mm. and R; somewhat divergent apically. Halter knob dark with pale shaft. Legs entirely dark; coxae with bright blue pruinescence. Coxae with long golden marginal hairs. Fore femur with | black, spur-like anteroventral bristle on apical fifth (Fig. 24); lacking anterior bristles on apical third, dorsal to spine-like bristles; anteroventral row of short, pale bristles, shorter than one-third width of femur; pos- teroventral row of slender, pale bristles longer than an- teroventral bristles, with 1 basal bristle longer than width of femur; anterodorsal region lacking row of stout bristles on basal third. Fore tibia with biserial row of short stout bristles beneath. Mid femur with 1 black, spur-like anteroventral preapical bristle, slightly shorter than width of femur, similar in size to anteroventral bristle of fore femur; lacking erect antero- and postero- dorsal bristles. Hind femur with | anteroventral preapi- Bradley J. SINCLAIR & Toyohei SAIGUSA: Trichoclinocera dasyscutellum Group from East Asia 201 cal bristle, subequal to width of femur, much more slender than preapical bristle of mid femur; lacking an- tero- and posterodorsal bristles. Hind tibia lacking pos- terodorsal spur-like bristle near base; 2 ventral preapical bristles, longer than width of tibia. Tarsı than corresponding tibiae. Empodium pulvilliform, sub- equal in length to claw, less than one-half length of tar- somere. Hypandrium (Fig. 5) cylindrical, broad, longer than epandrium. Phallus short, strongly curved sub-basally; anterior margin of apex slightly curved, lacking mem- branous sac; posterior margin of apex with rounded, membranous lobes; ejaculatory apodeme sharply re- curved anteriorly. Surstylus sharply produced posteri- orly, gun-shaped, with membranous oval scolex-like apex. Clasping cercus straight, projected obliquely, fin- ger-like, strongly tapered to narrow apex, bearing stout anteroventral bristles on outer face; inner margin with long bristles, lacking peg-like bristles. Female. Similar to male except as follows: fore femur with 2-3 spur-like bristles, apical 2 bristles subequal to width of femur; biserial row of bristles beneath not as dense and distinctive as male; fore tibia lacking modi- fied bristles beneath. Terminalia very similar to 7. fai- wanensis: sclerites of segment 8 overlap laterally; TS with sparse posterior row of long and short bristles, dor- sal bristles nearly subequal to length of sclerite; T10 with stout, curved bristles on apical two-thirds, with apical bristles more stout than remaining bristles; an- teroventral corner of T10 narrowed and prolonged obliquely. Geographical Distribution and Seasonal Occurrence. This species ranges from Northern India and Nepal, eastward to Yunnan (Fig. 12). In Nepal it ranges from 1000-2000 m, sympatric with 7. nepalensis. 3.7. Trichoclinocera naumanni sp. nov. (Figs. 6, 12) Type Material. Holotype, male labelled: “[CHINA: Si- chuan]/ Maziba, 1100-/ 1300 m, Emeishan/ Aug. 16, 1998/ T. SAIGUSA col.” (BLKU). Paratypes: CHINA: Sichuan: 4 $, 8 Y, same data as holotype (BLKU, ZFMK). Recognition. Distinguished by its brownish integument, especially the pale brown coxae and arched and strongly tapered clasping cercus. Description. Wing length 3.6-4.0 mm. Male. Dark brown with brownish-orange markings and faint blue pruinescence. Head rounded; compound eye large, rounded. Face narrowest at middle, ca. width of antennal sockets, with pale whitish pruinescence on lower third; lower margin of face with slight cleft with longer small carina or swelling; width of gena one-sixth height of eye. Vertex and ocellar triangle dark brown, lower postocciput and gena slightly pruinescent. Ocellar bris- tles longest among cranial bristles; occiput with 3 pairs of bristles posterior to ocellar triangle; postocular with row of short bristles along margin of eye, upper 34 bristles dark, stouter than lower bristles. Arista-like sty- lus long and slender, not tapered apically. Palpus some- what clavate, length less than one-third height of eye. Pleura brown with pale areas along sutures, with tinge of pale blue pruinescence when viewed from above: postpronotal lobe, notopleuron, posterior half of ane- pimeron pale brown and posterolateral corner of scutum including postalar ridge pale brown. Thoracic bristles short, less than one-half width of eye; acr lacking: 5 pairs slender dc; 1 slender short pprn, one-third to one- half length of de; 1 presut spal; 2 stout npl; 2 psut spal: 1 very short, slender pal; 2 pairs sctl and several pale setulae laterally; several pale setulae on notopleural de- pression. Laterotergite with small patch of dark bristles: prothoracic episternum with pale hairs; katepisternum and hind margins of metepisternum lacking hairs or very scarce. Wing narrow, infuscate; narrow, faint, elliptical, stigma present. One basal costal bristle, nearly twice length of posterior de. Row of setulae from base of R to tip of Ry: cell dm produced apically; Ry and R; divergent. Halter knob dark with pale shaft. Legs dark brown, except inner base of mid femur and base of hind femur pale; coxae brownish orange, darker on anterior margin of fore coxa; lacking blue pruines- cence. Coxae with long dark marginal hairs. Fore femur with 2 black, spur-like anteroventral subapical bristles: lacking anterior bristles on apical third, dorsal to spine- like bristles; anteroventral row of short, pale bristles, ca. one-half width of femur; posteroventral row of slender. pale bristles with basal bristles nearly subequal to width of femur, with I basal bristle twice width of femur. Fore tibia with biserial row of short, stout bristles beneath. Mid femur with | black, spur-like anteroventral preapi- cal bristle, subequal to width of femur, similar in size to anteroventral bristles of fore femur; lacking erect an- tero- and posterodorsal bristles. Hind femur with | slen- der anteroventral preapical bristle, subequal to width of femur, much more slender than preapical bristle of mid femur; lacking antero- and posterodorsal bristles. Hind tibia lacking posterodorsal spur-like bristle near base; 2 ventral erect bristles, longer than width of tibia on apical fifth; 1-2 erect, preapical dorsal bristles. Tarsi longer than corresponding tibiae. Empodium pulvilliform, sub- equal in length to claw, ca. one-half length of tarsomere. Hypandrium (Fig. 6) cylindrical, distinctly longer than epandrium; anterior margin produced into broad apo- deme. Phallus not extending beyond clasping cercus: 202 Bonner zoologische Beiträge 53 (2004) Fig. 12: Known distribution of species of Trichoclinocera dasyscutellum group. 7. fluviatilis (Brunetti) (O), 7. naumanni sp. nov. (E). somewhat curved sub-basally; anterior margin of apex curved, with short membranous sac; posterior margin of apex with rounded pair of lobes, clothed in microtrichia; long, straight rod with membranous sheath projected apically, nearly one-third length of phallic shaft, often partially or completed retracted within shaft of phallus; ejaculatory apodeme arched strongly anteriorly. Sursty- lus slender, subequal in length to clasping cercus with angular base, partially arched subapically, with mem- branous cylindrical scolex-like apex. Clasping cercus strongly curved, claw-like, strongly tapered to narrow apex, bearing short bristles on outer face, concentrated near apex; inner margin with short slender bristles. Female. Similar to male except as follows: fore femur with 3-4 preapical, anteroventral spur-like bristles; bise- rial row of bristles beneath not as dense and distinctive as male; fore tibia lacking modified bristles beneath. Terminalia similar to 7. faiwanensis. Geographical Distribution and Seasonal Occurrence. This species 1s currently known only from the Sichuan Province of China (Fig. 12). Remarks. This species is most closely related to 7. yunnana, on the basis of their very similarly shaped clasping cercus and configuration of the membranous apex of the phallus. Etymology. The specific name honours the late Dr. Clas M. NAUMANN, in recognition of his dedication to ento- mology, systematics and Museum Koenig. 3.8. Trichoclinocera nepalensis sp. nov. (gs. 8, 15 17,25) Type Material. Holotype, male labelled: “NEPAL, Ktmd./ Godavari 6000'/ 18 July 1967/ Can. Nepal Ex- ped.” (CNC). Paratypes: NEPAL: 10 Í, 5 Y, same data as holotype (CNC); 8 Í, 14 Y, 22.vii.1967, same locality as holo- type (CNC); 5 6, 17 Y, Lelep, 1770 m, 27° 31'N, 87° ATE, 1-4.vi.1972, JE/ HS (BLKU); 1 9, Pulchauk, 6600', 21.vii.1967, Can. Nepal Exped. (CNC); 2 6, Takwa, 1500 m, 27° 31'N, 87° 48'E, Lelep, 1770 m, 27° 31'N, 87° 47'E, 1.vi.1972, HS (BLKU). VIETNAM: 12 3,22 Y, Lao Cai Prov., Sa Pa, 1600 m, 27.v.1997, RM (BLKU). Several specimens deposited in ZFMK. Recognition. Distinguished from other Nepalese spe- cies by lacking distinctive blue pruinescence, straight Bradley J. SINCLAIR & Toyohei SAIGUSA: Trichoclinocera dasyscutellum Group from East Asia 203 e A + CA oF 2% 15 16 Figs. 13-16: Wings of Trichoclinocera dasyscutellum group. (13) T. dasyscutellum (Saigusa); (14) 7. fluviatilis (Brunetti): (15) 7 nepalensis sp. nov.; (16) 7. yunnana sp. nov. Scale bar = 1.0 mm. Fig. 17: Known distribution of species of Trichoclinocera dasyscutellum group. T. nepalensis sp. nov. (0). 7. vunnana sp. nov. (m). clasping cercus and surstylus bearing a stout, apical antennal sockets, with pale whitish pruinescence on bristle. lower half; lower margin of face with slight cleft with Dbeseription Wine length 3.6-4.3-tani: smell un % swelling; w idth of gena one-sixth height of eye. Vertex and ocellar triangle dark brown, lower Male. Dark brown with brownish-orange markings and postocciput and gena slightly pruinescent. Ocellar bris- faint blue pruinescence. Head rounded; compound eye _ tles longest among cranial bristles: occiput with 3 pairs large, rounded. Face narrowest at middle, ca. width of of bristles posterior to ocellar triangle; postocular with 204 row of short bristles along margin of eye, upper 3-4 bristles dark, stouter than lower bristles. Arista-like style long and slender, slightly tapered apically. Palpus somewhat clavate, length less than one-third height of eye. Pleura brown with tinge of pale blue pruinescence; postpronotal lobe, notopleuron, posterior half of ane- pimeron pale brown and posterolateral corner of scutum including postalar ridge pale brown. Thoracic bristles short, less than one-half width of eye; acr lacking; 5 pairs slender de; 1 slender short pprn, one-fourth length of de; 1 presut spal; 2 stout npl; 2 psut spal; 1 very short, slender pal; 2 pairs sctl and several pale setulae laterally; several pale setulae on notopleural depression. Laterotergite with patch of dark bristles; prothoracic episternum with pale hairs; katepisternum and hind margins of metepisternum lacking hairs or very scarce. Wing (Fig. 15) narrow, infuscate; narrow, faint, ellipti- cal, stigma present. One basal costal bristle, nearly twice length of posterior dc. Row of setulae from base of R to tip of R,; cell dm produced apically; Ry and Rs; somewhat divergent apically. Halter knob dark with pale shaft. Legs dark brown, except inner base of hind femur pale; coxae brownish orange, darker on anterior margin, es- pecially fore coxa; lacking blue pruinescence. Coxae with long dark marginal hairs. Fore femur with | black, spur-like anteroventral subapical bristle (Fig. 25); lack- ing anterior bristles on apical third, dorsal to spine-like bristles; anteroventral row of short, pale bristles, ca. one-half width of femur, with 1 basal bristle nearly twice width of femur; posteroventral row of slender, pale bristles similar to anteroventral bristles, with | basal bristle twice width of femur; anterodorsal region with row of very short, reduced bristles on basal third. Fore tibia with biserial row of short stout bristles be- neath. Mid femur with | black, spur-like anteroventral preapical bristle, subequal to width of femur, similar in size to anteroventral bristle of fore femur; lacking erect antero- and posterodorsal bristles. Hind femur with 1 slender anteroventral preapical bristle, shorter than width of femur, much more slender than preapical bris- tle of mid femur; lacking antero- and posterodorsal bris- tles. Hind tibia lacking posterodorsal spur-like bristle near base; 2 ventral erect bristles, longer than width of tibia on apical fifth; 1 erect, preapical dorsal bristle. Tarsi longer than corresponding tibiae. Empodium pulvilliform, subequal in length to claw, ca. one-half length of tarsomere. Hypandrium (Fig. 8) oval, more broad anteriorly, slightly longer than epandrium. Phallus short, shallowly curved sub-basally; anterior margin of. apex curved, with short membranous lobe clothed in microtrichia; posterior margin of apex with rounded membranous Bonner zoologische Beitráge 53 (2004) lobes; ejaculatory apodeme gradually recurved anteri- orly. Surstylus ca. one-half length of clasping cercus, sharply produced anteroapically, with membranous pos- terior and apical margin clothed with short microtrichia; apex bearing long bristle mounted on cylindrical proc- ess. Clasping cercus straight, finger-like, gradually ta- pered to rounded apex, bearing slender anteroventral bristles on outer face; inner margin with short, stout, peg-like bristles. Female. Similar to male except as follows: fore femur with 0-3 stout anteroventral bristles basal to preapical spine-like bristle; biserial row of bristles beneath not as dense and distinctive as male; fore tibia lacking modi- fied bristles beneath. Terminalia very similar to 7. tai- wanensis: sclerites of segment 8 partially overlap later- ally; T8 with sparse posterior row of long and short bristles, dorsal bristles nearly subequal to length of sclerite; T10 with stout, curved bristles on apical two- thirds, with apical bristles more stout than remaining bristles; anteroventral corner of T10 narrowed and pro- longed obliquely. Geographical Distribution and Seasonal Occurrence. This species is known from Nepal and northern Vietnam (Fig. 17) and is sympatric with 7. fluviatilis in Nepal and 7. yunnana in Vietnam. Remarks. The clasping cercus of the Vietnam popula- tion is more straight compared to the Nepalese popula- tion which is slightly arched apically. Etymology. The specific name refers to the type local- ity and where the majority of the specimens were col- lected. 3.9. Trichoclinocera taiwanensis sp. nov. (Figs, 7,9, 10, 21,26) Type Material. Holotype male labelled: “(Tatwan)/ Wulai/ Taipei Hsien”; “17.1v.1965/ T. Saigusa” (BLKU). Paratypes: TAIWAN: Chiayi-hs.: 2 Í, Chiaoliping, 14.1v.1965, TS (BLKU). Nantou-hs.: 3 Y, Tungyenchi, 950 m, Meichi, Jenai-hsiang, 19.x1.1997, BJS (CNC). Taichung-hs.: 32 $, 37 Y, Suchilanchi, 1600 m, Huan- shan, nr. Lishan, 24-25.x1.1997, river, BJS (ZFMK). Taipei-hs.: 26 Ö, 16 Y, same data as holotype (BLKU); 1 3,1 9, Wulai, 31.111.1984, K. Ohara (BLKU); 16 3, 32 Y, Fushan, Wulai-hsiang, 400-600 m, 28.x1.1997, river, BJS (CNC, ZFMK); 1 4, 2 Y, Han Hsien, Fushan, LTER Site, small river No. 8, 27.11.2003, L. Papp (HNHM). Recognition. This species is readily distinguished by the slender pointed anteroapical lobe of the phallus and dull coloured stalks of the halteres. Description. Wing length 4.5-5 mm. Bradley J. SINCLAIR & Toyohei SAIGUSA: Trichoclinocera dasyscutellum Group from East Asia () 188 rn ER ay RR Z aD Figs. 18-21: Scanning electron photographs of Trichoclinocera. spp. (18) T. dasyscutellum (Saigusa), antennae: (19) 7. dasys- cutellum (Saigusa), compound eye; (20) 7. dasyscutellum (Saigusa), compound eye and ommatrichia; (21) 7. raiwanensis sp. nov., male terminalia, lateral view. Male. Dark brown with distinct blue pruinescence. Head rounded; compound eye large, rounded. Face nar- rowest at middle, ca. width of antennal sockets, with distinct blue pruinescence, lower margin of face nearly straight, lacking pronounced carina or notch; width of gena one-sixth height of eye. Ocellar bristles longest 7 among cranial bristles; occiput with 3 pairs of bristles posterior to ocellar triangle; postocular with row of short bristles along margin of eye, bristles of upper half to third of eye dark, stouter than lower bristles. Arista- like stylus long and slender, slightly tapered apically. Palpus somewhat clavate, length less than one-half height of eye. Pleura, postpronotal lobe and notopleuron entirely clothed in bright blue pruinescence; posterolateral cor- ner of scutum including postalar ridge pale blue. Tho- racic bristles short, nearly one-half width of eye: acr lacking; 5 pairs de; I pprn; 1 presut spal; 2 npl; 2 psut spal; I very short, slender pal; 2 pairs sctl and numerous dark setulae on disc; several pale bristles on notopleural depression. Laterotergite with patch of pale bristles; I prothoracic episternum, katepisternum, and hind mar- gins of metepisternum with pale hairs. Wing narrow, infuscate; narrow, faint, parallel-sided stigma present. One short, basal costal bristle, subequal to length of posterior dc. Row of setulae from base of R to tip of Rj; cell dm strongly produced apically: Ry and Rs parallel-sided. Halter dark with dull base. not brightly coloured. Legs dark; coxae with pale blue pruinescence. Coxae with long golden marginal hairs. Fore femur with 3-4 black, spur-like anteroventral bristles on apical fourth (Fig. 26); lacking anterior bristles on apical third, dorsal to spine-like bristles; anteroventral biserial row of short bristles, shorter than half width of femur: posteroventral row of slender bristles subequal to one-half width of femur, 2-3 basal bristles subequal to width of femur: an- terodorsal region lacking row of stout bristles on basal third. Fore tibia with biserial row of short, stout bristles beneath. Mid femur with | black, spine-like anteroven- tral preapical bristle, shorter than width of femur, simi- 206 lar to fore femur; lacking erect antero- and posterodorsal bristles. Hind femur with | anteroventral preapical bris- tle, subequal to width of femur, more slender than preapical bristle of mid femur; lacking antero- and pos- terodorsal bristles. Hind tibia lacking posterodorsal spur-like bristle near base; 2 ventral and 2 dorsal preapical bristles, longer than width of tibia. Tarsı longer than corresponding tibiae. Empodium pulvilli- form, subequal in length to claw, less than one-half length of tarsomere. Hypandrium (Figs. 10, 21) cylindrical, only slightly longer than epandrium. Phallus short, curved sub- basally; anterior margin of apex curved, produced into curved, blade-like, membranous tapered sac; posterior margin of apex with rounded pair of roughened lobes, and long, straight rod with membranous sheath pro- jected apically, nearly one-third length of phallic shaft; ejaculatory apodeme straight anteriorly. Surstylus slen- der, distinctly bent, twisted and arched medially, with membranous oval scolex-like apex. Clasping cercus curved, finger-like, strongly tapered to narrow apex, bearing rows of short bristles on outer face; inner mar- gin with long silky bristles, subequal to width of lobe. Female. Similar to male except as follows: fore femur with 5-6 spur-like bristles, increasing in length apically, with stout, short bristles intermixed in anteroventral row; fore tibia lacking modified bristles beneath. Ter- minalia (Fig. 9): sclerites of segment 8 overlap laterally; TS with sparse posterior row of long and short bristles, dorsal bristles subequal to length of sclerite; T10 with stout, curved bristles on apical two-thirds, with apical bristles more stout than remaining bristles; anteroventral corner of T10 narrowed and prolonged obliquely. Geographical Distribution and Seasonal Occurrence. This species is restricted to Taiwan (Fig. 7). Remarks. The apex of the phallus was often found re- tracted (Fig. 10), where the long apical rod is withdrawn in the shaft of the phallus and the blade-like sac is also not fully inflated. This retractable rod is present in 7. fluviatilis, T. naumanni, T, yunnana and possibly also 7. feuerborni. This structure is unique to this subgroup and supports there close phylogenetic relationship. Etymology. The specific name refers to its geographic distribution. 3.10. Trichoclinocera yunnana sp. nov. (Figs. 11, 16, 17, 27) Type Material. Holotype, male labelled: “[CHINA: YUNNAN]/ Huanglianshan/ 1800m, nr. Sanmeng/ Lu- chun, 9 Mar.1995/ T. SAIGUSA col.” (BLKU). Paratypes: CHINA: Shaanxi: 1 @, 1 Y, Fuping-x, Lian- fengya, 2000-2100 m, 12 kmWNW, Longcaoping, Bonner zoologische Beitráge 53 (2004) 24.vi.1997, TS (BLKU); 6 3, 14 9, Fuping-x, nr. Long- cao, 1310-1400 m, 8 kmN Longcaoping, 27.vi.1997, TS (BLKU); 8 3, 3 Y, Fuping-x, Xigou, 1340-1400 m, 5 kmNW Longcaoping, 27.vi.1997, TS (BLKU); 1 Ú, 1 7, Zhouzhi-x, Shuimoping, 1500 m, SSW Banfangzi, 3.vii.1997, TS (BLKU). Sichuan: 5 Í, 10 9, Jingshui, 700-750 m, Emeishan, 7.viii.1998, TS (BLKU); 1 9, Maziba, 1100-1300 m, Emeishan, 16.viii.1998, TS (BLKU). 1 Í, 4 Y, Wuxiangang, 700 m, Emeishan, 14.viii.1998, TS (BLKU). Yunnan: 36 4, 56 9, same data as holotype (BLKU); 19 4, 32 Y, Dali, Daboping, 1900-2200 m, 7 kmW Xianguan, 16.vili.1996, TS (BLKU); 2 3, Hekou, 3 km from Laofanzha, 200 m, 1.1ii.1995, TS (BLKU); 5 d, 1 2, Hekou, Nanxi- Quiatou, 200 m, 37 km from Hekon, 28.11.1995, TS (BLKU); 2 $, 1 Y, Nanxi-Quiatou, 150 m, 27 km from Hekou, 26.11.1995, TS (BLKU); 42 Í, 52 Y, Jingcheng, 28 km from Jiangcheng, 1125 m, 12.111.1995, TS (BLKU); 1 9, Jingping, Fenshuiling Nat. Res. 1850 m, 25.v.1996, TS (BLKU); 2 Í, 2 Y, Pingbian, Xinxinghe, 780 m, S. Pingbian, 3.111.1995, TS (BLKU); 1 d,5 9, Yongping, Meihuapu, 2300 m, 83 km from Xiaguan, 4.vi.1996, TS (BLKU). VIETNAM: 1 4, 1 9, Lao Cai Prov., Sa Pa, 1700 m, 26.v.1997, RM (BLKU); 5 $, 12 Q, Lao Cai Prov., Sa Pa, "11600" m 274119972 RM (BLKU). Several specimens deposited in ZFMK. Recognition. This species is distinguished by the or- ange-brown coloured stalks of the halteres and the strongly arched and tapered clasping cercus. Description. Wing length 4.3-5.4 mm. Male. Dark brown with distinct blue pruinescence. Head rounded; compound eye large, rounded. Face nar- rowest at middle, ca. width of antennal sockets, with distinct blue pruinescence, lower margin of face nearly straight with slight notch; width of gena one-fifth height of eye. Ocellar bristles longest among cranial bristles; occiput with 3 pairs of bristles posterior to ocellar trian- gle; postocular with row of short bristles along margin of eye, bristles of upper third of eye dark, stouter than lower bristles. Arista-like stylus long and slender, ta- pered apically. Palpus somewhat clavate, length ca. one- third half height of eye. Pleura, postpronotal lobe and notopleuron entirely clothed in bright blue pruinescence; posterolateral cor- ner of scutum including postalar ridge pale brown. Tho- racic bristles short, less than one-half width of eye; acr lacking; 5 pairs de; 1 pprn; | presut spal; 2 npl; 2 psut spal; | very short, slender pal; 2 pairs sctl and numerous pale setulae on disc and margin; several pale bristles on notopleural depression. Laterotergite with patch of long, dark bristles; prothoracic episternum with pale hairs; katepisternum and hind margins of metepisternum lack- ing hairs. Bradley J. SINCLAIR & Toyohei SAIGUSA: Trichoclinocera dasyscutellum Group from East Asia () 188 rn Figs. 22-27: Scanning electron photographs of male forelegs of Trichoclinocera spp., lateral view. (22) 7. asiatica sp. nov.: (23 Wing (Fig. 16) narrow, infuscate; narrow, faint, paral- lel-sided stigma present. One short, basal costal bristle, longer than length of posterior de. Row of setulae from base of R to tip of Rj; cell dm strongly produced api- cally; Ry and Rs parallel-sided, except at extreme apex. Halter dark with pale base. Legs dark, except inner base of hind femur paler; coxae brownish orange, darker on anterior margin, especially fore coxa; some specimens with blue pruinescence. Coxae with long, golden marginal hairs. Fore femur with 1 black, spur-like anteroventral preapical bristle (Fig. 27); lacking anterior bristles on apical third, dorsal to spine-like bristles; anteroventral row of slender bris- tles, one-half to one-third width of femur; posteroventral row of slender bristles similar to anteroventral row, syscutellum (Saigusa); (24) T. fluviatilis (Brunetti); (25) 7. nepalensis sp. nov.; (26) T. taiwanensis sp. nov.; (27) T. yunnana sp. nov basal bristles only slightly longer than preceding bris- tles; anterodorsal region lacking row of stout bristles on basal third. Fore tibia with biserial row of slender bris- tles beneath, one-half width of tibia. Mid femur with black, spine-like anteroventral preapical bristle, sub- equal to width of femur, similar to fore femur: lacking antero- and posterodorsal bristles. Hind femur with | anteroventral preapical bristle, subequal to width of fe- mur, more slender than preapical bristle of mid femur: lacking antero- and posterodorsal bristles. Hind tibia lacking posterodorsal spur-like bristle near base; 2 ven- tral and 2 dorsal, erect preapical bristles, longer width of tibia. Tarsi longer than corresponding tibiae Empodium pulvilliform, subequal in length to claw, more than one-half length of tarsomere 208 Hypandrium (Fig. 11) cylindrical, slightly longer than epandrium; anterior margin produced and pointed. Phal- lus not extending beyond clasping cercus; somewhat curved sub-basally; anterior margin of apex curved, with short membranous sac; posterior margin of apex with rounded pair of lobes, clothed in microtrichia; long, straight rod with membranous sheath projected apically, nearly one-third length of phallic shaft, often partially or completed retracted within shaft of phallus; ejaculatory apodeme gradually arched anteriorly. Surstylus slender, subequal to length of clasping cercus, partially arched subapically, with membranous oval scolex-like apex. Clasping cercus strongly curved, claw- like, strongly tapered to narrow apex, bearing short bris- tles on outer face, concentrated near apex; inner margin lacking stout bristles. Female. Similar to male except as follows: fore femur with 2-6 spine-like bristles, increasing in length apically with bristles of posteroventral margin increasing in length basally; fore tibia lacking modified bristles be- neath. Terminalia very similar to 7. taiwanensis: sclerites of segment 8 partially overlap laterally; T8 with posterior row of long and short bristles, dorsal bris- tles more dense and nearly subequal to length of sclerite; T10 with stout, curved bristles on apical two- thirds, with apical bristles more stout than remaining bristles; anteroventral corner of T10 narrowed and pro- longed obliquely. Geographical Distribution and Seasonal Occurrence. This species is known from Shaanxi, Yunnan and Sichuan Provinces of China and northern Vietnam (Fig. 17). Remarks. The population from Shaanxi Province bears stouter bristles on the clasping cercus and the base of the surstylus is more distinctly elbowed. Etymology. The specific name refers to the region where the majority of specimens were collected, includ- ing the type locality. 3.11. Trichoclinocera sp. Material examined. THAILAND: 1 2, S. Banna, Nak- hon, 108 m, 5-10.v.1958, T.C. MAA (BPBM). Diagnosis. Coxae and base of hind femur orange- brown, bearing golden bristles. Fore femur with 2 an- teroventral spine-like bristles; single, dorsal spur-like seta at base of hind femur. Radial branch very short; M, and M, arising from cell dm closely approximated. Remarks. The limited material available of this species prevents providing a full description and formal specific name. On the basis of leg colouration, it is possibly re- lated to T. asiatica. Acknowledgements. We are indebted to the following cu- rators and their respective institutions for the loan or per- Bonner zoologische Beitráge 53 (2004) mission to examine material: J. Chainey (BMNH), N.L. Evenhuis (BPBM), J.M. Cumming (CNC), M. Ohara (HUS), M. Földvari, (HNHM), P. Tschorsnig (SMNS) and M. Datta (ZSI). T. Tachi and A. Inoue (BLKU) are thanked for producing the wing photos, assisted with the Japanese regional map and creating the SEMs, I.V. Shamshev (St. Petersburg, Russia) provided helpful interpretations for Russian locality names and B. Huber (ZFMK) kindly gave advice concerning the presentation of the maps. This study was partially funded by a Grant from the Ministry of Edu- cation (Japan). REFERENCES BRUNETTI, E. (1913): New Indian Empidae. Records of the Indian Museum 9: 11-45. ENGEL, E. O. (1931): XV. Empididae (Dipt.) aus Java. Ar- chiv fiir Hydrobiologie, Supplement Band 8: 486-489. SAIGUSA, T. (1965): Two new species of Clinocerinae from Japan (Diptera, Empididae). Kontyú 33(1): 53-57. SINCLAIR, B. J. (1994): Revision of the Nearctic species of Trichoclinocera Collin (Diptera: Empididae: Clinoceri- nae). The Canadian Entomologist 126: 1007-1059. SINCLAIR, B. J. (1995): Generic revision of the Clinocerinae (Empididae), and description and phylogenetic relation- ships of the Trichopezinae, new status (Diptera: Empi- doidea). The Canadian Entomologist 127: 665-752. WAGNER, R., LEESE, F. & PANESAR, A. R. (2004): Aquatic dance flies from a small Himalayan mountain stream (Diptera: Empididae: Hemerodromiinae, Trichopezi- nae and Clinocerinae). Bonner zoologische Beitráge 52(1-2): 3-32. Authors’ addresses: Bradley J. SINCLAIR (correspond- ing author), Zoologisches Forschungsmuseum Alexan- der Koenig, Adenauerallee 160, 53113 Bonn, Germany, E-mail: b.sinclair.zfmk(@uni-bonn.de; Toyohei SAIGUSA, 2-7-1-402, Baikoen, Chucku, Fukuoka, 810-0035, Japan. APPENDIX Clinocera velutina (Engel) Kowarzia velutina Engel, 1931: 488. Clinocera velutina: SINCLAIR 1995: 693 Type Material. Lectotype male (here designated) la- belled: “Fy 2C 9/7”; “Tjiborom-Fall/ Tjobodas. W. Java./ 9.V11.29. (Feuerborn.) [hand-written]”; “[red la- bel, no writing]”; [dissected, micro-slide mounted be- neath] (SMNS). A lectotype label “LECTOTYPE/ of Kowarzia/ velutina Engel/ des. B.J. Sinclair 2004 [red label]” has been attached to this specimen. Paralectotype: “Wasserfall.b./ Sarangan. M. Java./ Prof. Feuerborn./ 6.V11.28 [hand-written]”; “[red label, no writing]” (1 Y, SMNS). A yellow paralectotype label has been attached to this specimen. Redescription. Wing length 3 mm. Bradley J. SINCLAIR & Toyohei SAIGUSA: Trichoclinocera dasyscutellum Group from East Asia 209 Male. Face with pale blue pruinescence, except upper margin of eye; lateral margins with 2-3 fine, pale setu- lae; lower third of face flat; vertex, gena and posterior region of head brown. Arista-like stylus long and slen- der. Postocular bristles dark and stout. Pleura with pale pruinescence present when viewed from several angles. Scutum with pair of indistinct vit- tae, separated by reddish-brown median stripe. Several short, pale acr scattered anterior to first de; 5 de; | pprn very slender and reduced; 2 npl, lower weak and short; I psut spal; I pal; 2 sctl. Laterotergite with patch of long, dark bristles. Wing infuscate; stigma at apex of cell sc; 1 short basal costal seta; costal spines present. R>,3 and Ry,s straight; auxiliary crossvein between Ry and R»,; absent; Ry bell- shaped; anal vein indistinct or lacking; cell dm produced apically. Halter dark. Coxae with pale pruinescence; coxae and femora bright yellowish-brown, tibia and tarsi increasingly becoming darker. Fore coxae with pale hairs. Fore femur with an- tero- and posteroventral row of short stout bristles on basal two-thirds; anterior preapical comb present; dense white ventral pile lacking. Fore tibia with row of stout bristles beneath. Mid and hind femora lacking preapical bristles. Hind tibia with dorsal and ventral erect bristles on apical third. Fifth tarsomere lacking dorsoapical ex- tension; empodium pale brown laterally. / Terminalia slide-mounted, compressed. Hypandrium broadly triangular. Phallus somewhat straight: phallic filament long, flexible and whip-like. Clasping cercus narrow with rounded tip, with dense bristles on inner anterior face. Surstylus with pointed tip. Female. Similar to male except as follows: ventral bristles of foreleg less distinct; legs darker, especially femora. Remarks. The male terminalia is not redrawn as the original illustration of ENGEL (1931) is adequate and it is permanently slide mounted and pinned beneath the specimen. SINCLAIR (1995) assigned this species to the C. lineata-group on the basis of the form of the sursty- lus. The presence of facial setulae is a defining synapo- morphy of Kowarzia, but they are also present in several Asian species, including C. velutina (SINCLAIR 1995). u „un Bonner zoologische Beiträge Band 53 (2004) | Heft 1/2 Seiten 211-220 Bonn, Juni 2005 Aphyosemion (Mesoaphyosemion) etsamense (Cyprinodontiformes: Aplocheiloidei: Nothobranchiidae), a New Species from the Monts de Cristal, Northwestern Gabon’ Rainer SONNENBERG” & Thomas BLUM” . 'Zoologisches Forschungsmuseum Alexander Koenig, Bonn, Germany 2 5 Owingen, Germany Abstract. Aphyosemion (Mesoaphyosemion) etsamense sp. nov., a species belonging to the A. (M.) cameronense group, is described from the Monts de Cristal in northwestern Gabon. It is actually known from a small number of localities in the Monts de Cristal, which is also the westernmost known distribution area of the A. cameronense species group in Ga- bon. The new species differs from the geographically close populations of other species by the male colouration. The distinct status is also supported by mtDNA data. This is the second endemic nothobranchiid species described from the Monts de Cristal. Key words. Aphyosemion cameronense species group, biodiversity, Central Africa 1. INTRODUCTION Gabon has, together with Cameroon, the highest diver- sity of cyprinodontiform fishes in Africa (data from HUBER 2000). The nothobranchiids comprise with al- most 50 described species the largest group, many of them with small distribution areas and endemic to Ga- bon. Several descriptions of new species in recent years demonstrate the limited knowledge of their distribution and diversity (HUBER 1994, 1998a, b, 1999: LEGROS 1999). As large parts of this country are still inade- quately known with regard to freshwater fish, the num- ber will certainly grow in the future, and without doubt it is important for biodiversity and conservation consid- erations to have detailed knowledge on species numbers and distribution. The Nothobranchiidae are the most abundant and spe- cious group of cyprinodontiform fishes in Gabon (HUBER, 2000). From the Ivindo basin, one of the main Ogoue tributaries in the northern inland, up to eight cy- prinodontiform species in syntopy are reported, seven of them nothobranchiids (BROSSET 2003). In the majority of the collection localities a member of the Aphvo- semion (Mesoaphyosemion) cameronense species group is found. They inhabit a large area in the inland plateau of Cameroon, Equatorial Guinea and Gabon and proba- bly also in the neighbouring areas of the Central African Republic and the Republic of Congo (AMIET 1987; DADANIAK et al. 1995; HUBER 2000). This group actu- ally includes the following described species: 4. (M.) 1 In Commemoration of Clas Michael Naumann zu Königsbrück (26.06.1939 — 15.02.2004) obscurum (Ahl, 1924a), A. (M.) amoenum Radda & Pürzl, 1976, A. (M.) haasi Radda & Pürzl, 1976, A. (M.) halleri Radda & Pürzl, 1976, A. (M.) maculatum Radda & Pürzl, 1977, and A. (M.) mimbon Huber, 1977 with mostly small distribution areas in Cameroon and Gabon and the highly polymorphic A. (M.) cameronense (Boulenger, 1903) with a large distribution area in the countries mentioned above. AMIET (1987) was the first who, during his study of the Cameroonian Aphyosemion and Fundulopanchax spe- cies, grouped populations of A. (M.) cameronense ac- cording to their differing male colour patterns into three 'phenotypes' which he suggested might have full species status. Later, some killifish hobbyists joined their knowledge, accumulated through own field trips, aquar- ium observations and several publications, about the 4. (M.) cameronense group in Cameroon and Gabon and classified within 4. (M.) cameronense six additional 'phenotypes' (DADANIAK et al. 1995). From a taxonomic point of view the status of all these populations, which are often restricted to small areas, still remains unclear. This 1s mostly due to a lack of clear cut diagnostic male colour patterns and generally an extremly conservative morphology. The first published evidence of the species described here is given in a report of a collection in Gabon by HUBER (1977, 1980). The author identified this species tentatively as A. sp. aff. obscurum, because the single collected male shows a superficial similarity with 4 (M.) obscurum trom Cameroon (HUBER 1977, fig. 11). Subsequent collections revealed high variability in male colour patterns as described below (see also figs. on pp. 212 Bonner zoologische Beitráge 53 (2004) 29, 416, 421 in DADANIAK et al. 1995 and pp. 66-67 in SEEGERS 1997). Later the species was called A. camer- onense phenotype 6 by DADANIAK et al. (1995), follow- ing the numbering of different phenotypes of the 4. (M.) cameronense group in Cameroon by AMIET (1987). Data presented here are mainly based on a collection from Gabon in 2002 where it was possible to find the new species and several other populations of the 4. (M.) cameronense group, including the formally described species A. (M.) mimbon and A. (M.) maculatum. Based on material from Gabon and Cameroon collected by the authors and additional material from Equatorial Guinea provided by M. JUHL we also tested the taxonomic status of the new species with mtDNA data. 2. MATERIAL AND METHODS The description is based on specimens collected by the authors during fieldtrips to Gabon 1999 (TB) and 2002 and deposited at the Zoologisches Forschungsinstitut und Museum Koenig, Bonn, Germany (ZFMK). A number of specimens of the type series will be sent to the Musée Royale de l'Afrique Centrale, Tervuren, Bel- gium (MRAC) and the Institute de Recherche en Ecolo- gie Tropicale (IRET) in Libreville, Gabon. All fishes were collected with a handnet, sacrified with MS222, preserved in 3,5% Formalin and later trans- ferred to 70% Ethanol for storage. Specimens for DNA studies were stored directly in 96% Ethanol. Measurements and counts follow AMIET (1987) with the addition of the caudal peduncle, measured from the pos- terior end of anal fin base to caudal end of peduncle. All measurements are presented as percentages of standard length (SL) and were made point to point with a digital calliper and corrected to the nearest 0.1 mm. Geo- graphical coordinates of sample points visited by the first author are taken with a Garmin GPS 12, map date was WGS 84. The spelling of locality names was taken from the IGN map (Kango, NA-32-V, scale 1:200 000) of this area. The distribution point map is made with the program DIVA-GIS version 4.2 (HIJMANS et al. 2001), GIS files for rivers, roads and country borders are used from the Central African Regional Program for the Envi- ronment (CARPE) website (http://carpe.umd.edu/). Addi- tional locality data are provided by M. JUHL, Denmark, U. KAMPF, Germany (pers. comm. 2003) and from Killi- Data 2000 (HUBER 2000). Definition and naming of the phenotypes 1-9 follows the definitions of AMIET (1987) and DADANIAK et al. (1995). A Map 1: Known distribution area of A. (M.) mimbon A, A. (M.) etsamense e and A. (M.) cameronense @ in northwestern Gabon and southern Equatorial Guinea. Type localities of A. (M.) mimbon (Akoga), A. (M.) etsamense (Etsam 1) and A. escherichi are shown together with the localities of the DNA samples, which are enclosed in an open circle and numbered according to field lo- cality number (see Table 1). Data from HUBER (2000), own data and M. JUHL, Denmark (pers. comm.). Rainer SONNENBERG & Thomas BLUM: Aphyosemion (Mesoaphyosemion) etsamense Table 1: List of specimen used for DNA sequencing with GenBank accession number, voucher number and geographical coor- dinates of collection localities. Sampleno. Species S lo Country SELL Latitude Longitude locality ; accession age? ae RS 639 A. (M.) cameronense G 02/153 Gabon AY 748279 0°57'38,7"N 11°08'25,3"1 RS 931 A. (M.) cameronense G 02/154 Gabon AY748280 0°58'18,8"N 11°05'13,2%8 RS 638 A. (M.) cameronense G 02/155 Gabon AY 74828] 1200'22,8"N 10°54'13,0"1 RS 262 A. (M.) cameronense CMM 40 Cameroon AY748282 2°48'56,1"N 10°40'35,0"I RS 570 A. (M.) mimbon G 02/158 Gabon AY748283 0°57'29,7"N 10°39'01,2"E RS 571 A. (M.) mimbon G 02/159 Gabon AY748284 0°56'46,0"N 10°38'14,2"E RS 640 A. (M.) mimbon G 02/157 Gabon AY748285 0258'06,3"N 10%41'33,5"! RS 929 A. (M.) mimbon G 02/156 Gabon AY748286 1°00'06,0"N 10%45'50,9"] RS 930 A. (M.) mimbon G 02/159 Gabon AY748287 0°56'46,0"N 10°38'14,2" I RS 974 A. (M.) mimbon EG 03/20 Equatorial Guinea AY748288 1°06'47,2"N 10°46'3 1.1" RS 545 A. (M.) etsamense G 02/160 Gabon AY748289 0°46'34,1"N 10°24'03,0"1 RS 577 A. (M.) etsamense BS 02/12 Gabon AY748290 0°46'34,1"N 10°24'03,0"E RS 604 A. (M.) etsamense G 02/160 Gabon AY748291 0°46'34,1"N 10°24'03,0"E RS 633 A. (M.) etsamense G 02/160 Gabon AY748292 0°46'34,1"N 10°24'03,0"E RS 923 A. (M.) etsamense G 02/160 Gabon AY748293 0°46'34,1"N 10°24'03.0"E RS 924 A. (M.) etsamense G 02/160 Gabon AY748294 0°46'34,1"N 10°24'03,0"E RS 925 A. (M.) etsamense BS 02/13 Gabon AY748295 0°43'36,9"N 10°21'58,1"I RS 1355 A, (M.) etsamense G 02/161 Gabon AY748296 0°42'58,1"N 10°21'37,0"E The specimens used in the DNA analysis are listed in Table 1 and were deposited in the tissue collection of the ZFMK. Map 1 shows the localities of the DNA samples. We sequenced a part of the mitochondrial cy- tochrome b gene, a detailed description of the method will be given in a subsequent paper. The resulting sequences were aligned by eye in BioEdit 5.0.9 (HALL 1999) and uncorrected pair-wise sequence divergence calculated with MEGA 2.1 (KUMAR et al. 2001). The sequences were deposited in GenBank, ac- cession numbers are listed in Table 1. Here we used the molecular data set only to get additional information on the distinctness of the species described here from its closest relatives and not to reconstruct a phylogeny of this complex species group. As indicators for species status we use distinctness of colour pattern with respect to all surrounding popula- tions of other related species and in addition the mono- phyletic cluster of mitochondrial haplotypes. This is seen as indirect evidence for different biospecies sensu MAYR. 3. RESULTS Aphyosemion (Mesoaphyosemion) etsamense, new species (Figs. 1-4) Aphyosemion sp. aff. obscurum Huber, 1977:6 (in part, only locality 55), fig. 11, 1980:38 1996:333, 2000:480 Aphyosemion sp. aff. cameronense Phänotyp 6, Dadaniak et al., 1995:416-424 , figs. pp. 29, 416, 421 Aphyosemion sp. aff. cameronense |Pop. 6], Seegers, 1997:66-67, 3 figs. Fig. 1: Aphyosemion (Mesoaphyosemion) etsamense sp. nov., male, same locality as Type, not preserved. Fig. 2: Aphyosemion (Mesoaphyosemion) etsamense sp. nov.. female, same locality as Type, not preserved. Holotype: ZFMK 39832, male, 32.4 mm SL: Gabon. western slopes of the Monts de Cristal, a small river at the village Etsam |, crossing the road N5 trom Sees to Kougouleu (0%46'34,1" N, 10°24'03" E), collected 2 July 2002 by T. Blum, G. F leck and R. Sonnenberg, col: lection locality G 02/160. 214 Bonner zoologische Beitráge 53 (2004) Fig. 3: Aphyosemion (Mesoaphyosemion) etsamense sp. nov., male from a locality north of Assok (BS 02/13), not preserved Fig. 4: Aphyosemion (Mesoaphyosemion) etsamense sp. nov., male from Assok, not preserved Paratypes: MRAC A4-42-P-1-4, same data as holotype, ZFMK 39833 — 39842, same data as holotype, ZFMK 39843, male, Gabon, north of the village Assok, col- lected 11. September 2002 by T. Blum and P. Sewer, collection locality BS 02/13 ( 0°43'36,9" N, 10°21'58,1" E). ZFMK 39844 - 39846, collected by F. Bitter, T. Blum and P. Sewer in August 1999, same locality data as holotype, preserved after approximately 2 years in aquarium. IRET, Gabon, two specimens, one male and one female, same data as holotype. Additional Material: Colour pictures of live specimens of the above and other populations of the A. camer- onense group are used to compare colour patterns (AMIET 1987; DADANIAK et al. 1995; HUBER 1977; SEEGERS 1997). Diagnosis: Aphyosemion (Mesoaphyosemion) et- samense is distinguished from other species of Aphyo- semion by its unique combination of colouration charac- ters in males. This distinctness is supported by a monophyletic lineage of mtDNA haplotypes. Males of A. (M.) etsamense are distinguished from all other members of this species group by its nearly com- plete yellow dorsal fin with small irregular red dots only on basal and posterior egde of fin versus blue or pale yellow with more red dots or flames (compare A. (M.) cameronense Figs. 10-12). The only exceptions are A. (M.) mimbon (Figs. 5 and 6) and a population of A. (M.) cf. cameronense in the Ivindo basin between Makokou and Ovan (Fig. 16), which are distinguished by their dif- ferent body, caudal, and anal fin colouration. 4. (M.) et- samense 1s also distinguished from all species and phe- notypes with a yellow caudal peduncle (A. (M.) amoenum (Fig. 8), A.(M.) halleri, A.(M.) sp. aff. camer- onense phenotype 3, 4, 5 (Fig. 14) and 9) through its complete blue colouration of the sides in males versus blue or bluegreen with a yellow or orange peduncle. It is distinguished from A. (M.) maculatum (Fig. 9), A. (M.) mimbon (Figs. 5 and 6) and A.(M.) sp. aff. maculatum from Equatorial Guinea by the pattern of the red pig- mentation on the males side. In A. (M.) etsamense the pigmentation is arranged in horizontal rows of dots which can fuse to closed or irregularly interrupted lines and in some specimens with red reticulation on the cau- dal peduncle versus more vertical distribution of red pigment clusters and horizontal rows of red dots only on anterior part of body and less extensive reticulated pat- tern (see Figs. 1, 3, 4, 5, 6, 10 for comparison). It differs from A. (M.) obscurum (Fig. 7) and A. (M.) sp. aff. cam- eronense phenotype | (Fig. 13) and 4 by the variability of the red colour pattern, which exceeds the observed variation in both species and the yellow dorsal and the sometimes yellow marginal bands in the caudal and anal fins in most specimens versus always white bands in the other three. Fig. 5: Aphyosemion (Mesoaphyosemion) mimbon from locali- ty BBS 99/23, same as G 02/157 Fig. 6: Aphyosemion (Mesoaphyosemion) mimbon from locali- ty GEB 94/25, same as G 02/158 Rainer SONNENBERG & Thomas BLUM: Aphyosemion (Mesoaphyosemion) It differs from A. (M.) sp. aff. cameronense phenotype 8 form fishes, a strong sexual dimorpl (Fig. 15) by a different colour pattern on the fins and more colourful and larger than females. Dor more red markings on the sides, where phenotype 8 has fin in males slightly pointed, caudal fin trapezoid less markings and more red dots in the dorsal and finer — slightly rounded posterior end, no filamentou red markings in the caudal fin. sions like in the A. calliurum group for example. | BE : Re females in general smaller and rounded. Dorsal fin on Description: See Figs. | — 4 for general appearance and ; = A : gin behind origin of anal fin (D/A ) and Table 2 for morphometric data of the type series. A ; : 5 hind mid length of body. Dorsal fin with 1] slender and elongate medium sized Mesoaphvosemion : E : 4 e eres anal fin with 15 16 rays, scales on mid-longitudina species. Snout is rounded, mouth directed upwards, pos- : Be ae ee : : series 31 O2 3 on caudal fin base. Traı terior end of mouth at same level as center of the eye. = : er rows of scales above pelvic fin 8 10, circu Dorsal profile straight. Shows, like most cyprinodonti : : peduncular scale row 13 - 14. Table 2: Morphometrics of Aphyosemion etsamense. All measurements in percents of standard length (SL), ¢ | length in mm. TL= total length, PD= predorsal fin distance, HL= length of head, pPD= prepelvic fin distance, pAD= preanal fıı distance, HB= greatest body height, HC= height of caudal peduncle, CL= length of caudal peduncle, BD= base of dor fi BA= base of anal fin, D= dorsal fin rays, A= anal fin rays, D/A= dorsal / anal fin position, E= eye diameter, [= interorbital widt LLS= lateral line scales, TS= transverse row of scales, CS= scales around caudal peduncle, SD= standard deviation TL SL PD HL pPD pAD HB HC CL BD_BA_D A D/A E I _LLS IS CS CU Holotype 125,4 32,4 65,4 28,1 43:7, 58,5 20,3 12,1 23,6 14,4 19,5 12.0 16,0 Paratypes mean (/) 124,7 30,4 67,9 26,2 45,8 60,5 18,4 10,4 22 | 0), | | | 8.0 Paratypes mean (©) 125,7 29,2 68,6 27,3 47,1 59,6 18,8 10,9 2 14,5 1,2 | 15, alltypes mean 125,3 29,8 68,2 26,9 46,4 59,9 18,7 10,8 22,4 14,2 0) 3 15.6 all types SD 2,6 5,4 2,6 1,4 3,0 2,2 1,1 06 11 09 1 Wi back; sides light blue, with brilliant hue, but not as me- tallic as in other species: on sides posterior to operct lum rows of red dots, forming up to four more or less continuous rows; in some specimen additional short rows, one starting below dorsal fin to caudal and one ventral behind pectoral fin which can fuse with ventral red band which is seen in most A. (M.) cameronense group fishes. The red colouration can also be irregularly interupted or fuse to red bands or blotches on caudal body sides, where it often forms a reticulated pattern on the caudal peduncle. Fig. 7: Aphyosemion (Mesoaphyosemion) obscurum, near Ma- tomb, Cameroon. Dorsal fin mainly yellow, with blue only on basal and posterior part, with red dots or flames basal and poste- rior distal. Anal fin colour ranging from light blue to almost completely yellow with small basal blue several specimens with red dots or flames on basal part: submarginal red band of varying size, marginal band white, blueish or yellow, in some specimens white with partially yellow. Caudal fin colour pattern also variable: center light blue with horizontal red flames or dots: red ee on anterior, posterior and submarginal parts often fused which encloses the blue central part: upper and lower narginal band white, bluish grey or yellow, as in anal in often yellow and white combined. Pectoral fins 2 transparent with marginal blue, pelvic fins blue, with Fig. 8: Aphyosemion (Mesoaphvosemion) amoenum, Cameroon. : . submarginal red band and white or blue marginal sim : a 2 > arto anal. Coloration: Living males (Figs. 1, 3, 4): snout yellow, dorsal part of body from mouth to dorsal fin base brown Living females (Fig. 2): body brown, dorsally darke to yellow, beneath eye a red sub-ocular line, on opercu- han ventrally; red dot at end of scales, fusing to a re lum three oblique bands of fused red dots, the upper in iculated pattern on caudal peduncle, traces of red sub- some specimens overlayed by yellow colouration of the ocular line and three oblique red lines on operculum 216 Bonner zoologische Beitráge 53 (2004) Fig. 9: Aphyosemion (Mesoaphyosemion) maculatum, near Fig. 13: Aphyosemion (Mesoaphyosemion) sp. aff. camero- Matora, collection loc. G 02/125, Gabon nense phenotype 1, near Mvilé, Cameroon Fig. 10: 4Aphvosemion (Mesoaphyosemion) cameronense, col- Fig. 14: Aphvosemion (Mesoaphyosemion) sp. aff. camer- lected in Gabon at Olong II near the border to Equatorial Gui- — onense phenotype 5, near Koumaméyong, Gabon nea, G 02/149 Fig. 11: Aphyosemion (Mesoaphyosemion) cameronense from Fig. 15: Aphyosemion (Mesoaphyosemion) sp. aff. camero- Cameroon, between Akom II and Ebolowa, collecting loc. nense phenotype 8, West Mitzic, Gabon CMM 40 Fig. 12: Aphyosemion (Mesoaphyosemion) cameronense from Fig. 16: Aphyosemion (Mesoaphyosemion) cf. cameronense the IRET Research Center near Makokou, G 02/126 BSW 99/11, between Makokou and Ovan at Minkwala. Rainer SONNENBERG & Thomas BLUM: Aphyosemion (Mesoaphyosemion) elsamens Fig. 17: A. escherichi, aquarium raised male from northern Gabon Fig. 18: 4. striatum, aquarium raised male from Cap Esterias, Gabon Fig. 19: Type locality of Aphyosemion (Mesoaphyosemion) etsamense at Etsam I, Gabon All unpaired fins and pelvic fins transparent with inter- radial red flames; anal and pelvic fins with traces of a small blueish marginal band, pectoral fins transparent with small whitish marginal band. Preserved in Ethanol: Males: body brown, upper part darker than lower, belly light brown, dark pigment on scales forming a reticulated pattern posterior to opercu- lum; fins pale grey, dorsal distal and posterior part with dark brown interradial flames, on caudal and anal fin whitish marginal bands, submarginal dark brown bands, posterior end of caudal dark brown, interradial dark brown flames; pelvic and pectoral fins pale grey, with submarginal dark brown and small whitish marginal band. Females: body brown, upper and dorsal part dar) brown, belly light brown, dark pigment on scales form ing a reticulated pattern posterior to operculum; all fin pale grey with dark brown interradial flames on all ex- cept pectorals. / DNA data: The alignment contains 800bp for a set of 18 specimens in total, 8 A. (M.) etsamense from 3 dif- ferent localities, 6 A. (M.) mimbon from 5 localities and 4 A. (M.) cameronense from 4 localities. 115 positions are variable, 89 might be phylogenetically informative. There are ten unambiguous potential syn- apomorphic positions in the sequences of A. (M.) er- samense. 14 positions are ambiguous in having whithin one A. (M.) etsamense (specimen RS 925) a character state also found in the outgroup (9) or one outgroup specimen having a state like in A. (M.) etsamense (5). The amount of uncorrected pair-wise sequence diver- gence between A. (M.) etsamense and A. (M.) mimbon is 4,25 - 6,63 % and between A. (M.) efsamense and the other specimens tentatively identified as A. (M.) camer- onense 6,25 - 8,37 %. The divergence between A. (M.) mimbon and A. (M.) cameronense is 5,62 - 6,88 %. Within A. (M.) etsamense sequence divergence is up to 2,25 % and, in comparison, within A. (M.) mimbon up to 3,13 %. We want to state that we use the distinctiveness of haplotypes with their potential apomorphic character states and not the genetic distance value as additional indicator for species status together with the colouration characters which unite the groups of populations from the different species. Etymology: This species is named after the village Et- sam I, Gabon, at the type locality (Fig. 19). Distribution and habitat: Only known from some populations found in rivulets along the road N5 between Medouneu and Kougouleu: localities near the villages Etsam I, Assok and Song (Map 1). The closest known locality of A. (M.) mimbon is the village Ntom (DADANIAK et al. 1995), ca. 2 km south of Akoga, which is the type locality of this species (HUBER 1977) and approximately 15 km north of Etsam I. According to the official map (Kango, NA-32-V, scale 1:200 000) all localities of A. (M.) mimbon in Gabon are found in tributaries of the Komo and the Mbe, those of 4. (ML) etsamense except at Song seem to belong to the Bin- guilé which is also a tributary of the Mbe below the Barrage de Tchimbele. The rivulets around the village Song empty into the Song river, a tributary of the Noya. The closest localities east of Medouneu are populated by fishes identified as A. (M.) cameronense s.s. (DADANIAK et al. 1995) and are found in tributaries ot the Mvo and Abanga rivers. Habitats are small rainforest creeks and rivers with slow flowing water. The type locality (Fig.19) is a larger, 218 slow flowing river, about 3-4 m wide and up to 0,8 m deep. This is a rather unusual habitat for species of the A. (M.) cameronense group which are usually found in rivulets not that deep. The fishes are all found between the terrestrial vegetation which hangs into the water, in addition they are also found in smaller rivulets emptying into the main river. The other known localities are small rainforest creeks between 0,5 and 2,0 m wide and usu- ally not deeper than 0,4-0,5 m. Remarks: Because A. (M.) etsamense could be found near Song (HUBER 1977, locality JH 55) within rivulets emptying into the Song river, which itself is a tributary to the Noya river (Nduya river according to WILDEKAMP 1993 and SEEGERS 1987, 1988, Nga river according to AHL 1924b) it is necessary to discriminate between A. (M.) etsamense and A. escherichi (AHL 1924b) as the type locality of the latter is at ‘Attogon- dema, Nga-Zuflüsse, Kamerun’ (AHL 1924b). In this area the two species A. escherichi and A. stria- tum (Boulenger, 1911) are often found in syntopy and both species have a very similar colour pattern (Figs 13 and 14). Bonner zoologische Beitráge 53 (2004) Whereas SEEGERS (1988) and WILDEKAMP (1993) list A. escherichi as a valid species with the junior syno- nyms A. microphtalmum Lambert & Gery, 1967 and A. simulans Radda & Huber, 1976, HUBER (1998c, 2000) is of the opinion that it represents a junior synonym of A. striatum (Boulenger, 1911). As A. (M.) etsamense occurs in the same river system like the former two spe- cies, we have to show that the new species is not con- specific with 4. escherichi. HUBER (1998c) based his opinion on small morphologi- cal differences, mainly dorsal and anal fin ray and ver- tebrae counts, although the species have a nearly com- plete overlap in these morphological values (Table 3). The vertebrae counts given by HUBER (1998c) for A. striatum and A. microphtalmum (mean 27,50 and 27,70 respectively) are closer to each other than to the type se- ries of A. escherichi (mean 29,55), so if they had a dis- criminatory value, then 4. striatum and A. microphtal- mum were closer together than to A. escherichi and the latter might represent a distinct species. Because of the large overlap, the previously listed morphological fea- tures have no discriminatory value in this case, and for a statistical significant distinction the number of speci- mens and samples had to be considerably larger. Table 3: Comparison of morphometric data from A. escherichi, A. microphtalmum, A. striatum (from HUBER, 1998c) and A. (M.) elsamense A. striatum (n = 8) (n= 3) Dorsal fin rays 10-11 10-11 Mean 10.75 10.33 Anal fin rays 14-16 16-17 Mean 14.88 16.33 D/A deviation 49-47 +7 - +8 Mean 6.00 7.67 A. microphtalmum A. escherichi A, (M.) etsamense (n=11) (n=21) 11-12 11-13 11.18 12.29 13-15 15-16 13.91 15.57 +5 -+6 +7 -+8 5.82 7.90 On the other hand, according to SEEGERS (1988) and WILDEKAMP (1993) and own observations, the major differences between 4. striatum versus A. escherichi and 4. (M.) etsamense are the obviously different colour patterns of the fins, especially the dorsal fin (Fig. 1, 3, 4, 17, 18). This character seems to be stable throughout the distribution area of the species. In addition the different relation between caudal peduncle length and height is a diagnostic character. These characters are also men- tioned in the first descriptions by AHL (1924b) and HOLLY (1930), who restudied AHL’S type series, for A. escherichi (caudal peduncle nearly 2 x height in length) and by BOULENGER (1911) for A. striatum (caudal pe- duncle about 1,33 x height in length). Despite the absence of significant differences in the caudal peduncle between 4. escherichi and A. (M.) et- samense (2,0 - 2,2 x times height in length, Table 2), the male colouration allows the identification of both species (see Figs. 1, 3, 4, 17) and also the discrimination between A. (M.) etsamense and A. striatum (Figs. 1, 3, 4, 18 ). The dorsal fin in A. (M.) etsamense shows little blue or red pigmentation on the posterior or basal part versus little yellow with lots of red pigmentation in 4. escherichi (dots and flames). A. striatum shows two parallel horizontal bands on blue or yellow background, the caudal peduncle height about 1,33 times in length. The often very regular horizontal rows of red dots on body sides in males in 4. escherichi and A. striatum are clearly an unrealiable discriminatory character to the ir- regular red pigmentation in A. (M.) etsamense, because on one hand we found several 4. (M.) etsamense males which also show a more or less regular pattern of hori- zontal red dots, on the other hand we found especially in the area of the Monts de Cristal populations of A. stria- tum with a very irregular pigmentation similar to the former species. Rainer SONNENBERG & Thomas BLUM: Aphyosemion (Mesoaphyosemion) etsamense 4. DISCUSSION In general it is difficult to find diagnostic characters to discriminate other species against the heterogenous as- semblage of populations which is summed up in the taxon A. (M.) cameronense. However, the discrimi- nation between A. (M.) etsamense and its genetically and geographically closest relative, 4. (M.) mimbon is easy due to the different colour patterns (see Figs. 5, 6). Also the distinction to the geographically close popula- tions we collected between Sam and Medouneu in the tributaries of the Mvo and Abanga rivers and most other populations, which are identified as A. cameronense (Figs 10-12) (DADANIAK et al. 1995; HUBER 2000), is possible because of their differing phenotype with little red pigmentation on the sides of males. Mesoaphyosemion: The taxon Mesoaphyosemion was erected by RADDA (1977) as a subgenus of Aphyo- semion. As type species he chose 4. cameronense (BOULENGER, 1903). However, RADDA also included in his newly erected subgenus all those species groups which show a superficial similarity in morphology and therefore turned this group into a taxonomic 'wastebas- ket'. Actual research by MURPHY & COLLIER (1999) based on DNA sequences confirm that Mesoaphyo- semion is not a monophyletic group. In this paper we suggest to restrict the taxon name Mesoaphyosemion Radda, 1977 to the A. (M.) cameronense species group and exclude all other species which then makes Mesoaphyosemion a monophyletic or natural group. This group now contains the following described spe- cies: A. (M.) cameronense (Boulenger, 1903), 4. (M.) obscurum (Ahl, 1924a), A.(M.) amoenum Radda & Púrzl, 1976, A. (M.) haasi, Radda & Púrzl, 1976, A. (M.) halleri Radda & Piirzl, 1976, 4. (M.) maculatum Radda & Piirzl, 1977, A. (M.) mimbon Huber, 1977 and A. (M.) etsamense. It includes also the undescribed, but recognizable groups of populations defined by AMIET (1987) and DADANIAK et al. (1995) with their different male colour patterns, which might, like A. (M.) et- samense, be regarded as distinct species. This is the second described endemic species of notho- branchiid fishes in the Monts de Cristal besides A. (M.) mimbon. For the nothobranchiids it is a species poor area, as only one other species group of Aphyosemion and very rarely a species of Episemion could be found in syntopy with a Mesoaphvosemion species (HUBER 1977, 2000; own data). For comparison, whithin the Ivindo basin the highest number is seven species of dil- ferent species groups (Brosset 2003) and for the coastal plain up to five or six species of different groups in syn- topy (own observations). Further collections might re- veal a higher diversity, as only a small part of this area is known with regard to cyprinodontiform fishes. 219 Résumé. Aphosemion (Mesoaphosemion) etsamense sp nov., une espece du groupe A. (M.) cameronense, est décrite des Monts de Cristal du nord-est du Gabon Cette espece n'est connue que d'un petit nombre de lo- calités situées dans les Monts de Cristal ce qui en fait l’espece du groupe cameronense a la distribution la plus occidentale. Cette espece nouvelle se différencie des populations des autres especes géographiquement pro- che par la coloration du male. Son statut d’espece a part entiere est également supporte par l’analyse d’ADN mi- tochondrial. Elle représente la deuxieme espece endé- mique de nothobranchiid des Monts de Cristal. Acknowledgements. The authors thank the Ministry of Scientific and Technical Research (MINREST), Yaounde, Cameroon, the Institut de Recherche Agricole pour le Développement (IRAD), Yaounde, the [RAD Fisheries & Oceanography Research Station, Batoke, Limbe, Camer- oon, for research permisson and support in Cameroon, the Centre National de la Recherche Scientifique et Tech- nologique (CENAREST) and the Institut de Recherche en Ecologie Tropicale (IRET), especially Prof. Dr. P. Posso, for research permisson and support in Gabon . We would also like to thank Dr. G. Fleck, Bonn for his support during the time in Gabon and the translation of the french abstract, F. Herder, PD Dr. B. Y. Misof, A. C. Schunke for their comments on earlier drafts of the manuscript and U. Kampf for providing excellent slides for publication. We also want to thank C. Akum and family and C. E. Gabche and family, Limbe, Cameroon for their generous help and support, and Dr. A. Kamdem Toham, WWF, Libreville, Gabon for his help. We like to thank F. Bitter, W. Eigel- shofen, Dr. J. H. Huber, M. Juhl, the Kampf family, Dr. H. Kullmann, K.-H. Lúke, Dr. L. Seegers, A. Tránkner, R.H. Wildekamp, all colleagues at the molecular lab of the ZFMK and a lot of other people too much to be listed here for various support. Both authors would like to thank especially all those friendly people in Cameroon and Gabon who helped us during fieldwork. A part of this work was financed by the DFG Project MI- 649/2-1, a travel grant for Cameroon was provided to RS by the Alexander-Koenig-Stiftung, Bonn. Rainer Sonnenberg wishes to dedicate this paper to the late Professor Dr. C. M. Naumann, in memory of his continu- ous support and encouragment during all my years at the ZFMK in Bonn. His comments, critics and help will be missed. REFERENCES AHL, E. (1924a): Zur Systematik der altweltlichen Zahnkarpten der Unterfamilie Fundulinae. Zoologi- scher Anzeiger 60: 49-55. AHL, E. (1924b): Uber neue afrikanische Zahnkarpfen der Gattung Panchax. Zoologischer Anzeiger 60: 303-313. AMIET, J. L. (1987): Le genre Aphyvosemion Myers (Pisces, Teleostei, Cyprinodontiformes). Faune du Cameroun, 220 Bonner zoologische Beitráge 53 (2004) Vol. 2. Sciences Nat, Compiegne, France: 1-262, plates 1-68. BOULENGER, G. A. (1903): Description of new Freshwater Fishes from Southern Cameroon. Annals and Maga- zine of Natural History ser. 7, 12: 435-441. BOULENGER, G. A. (1911): Descriptions of new African Cyprinodont Fishes. Annals and Magazine of Natural History, ser. 8, 8: 260-268. BROSSET, A. (2003): Convergent and divergent evolution in rain-forest populations and communities of cypri- nodontiform fishes (Aphyosemion and Rivulus) in Af- rica and South America. Canadian Journal of Zoology 81: 1484-1493. DADANIAK, N., LUTJE, R. & W. EBERL (1995): Faszination Killifische: Die “Aphyosemion cameronense”-Gruppe. 480 pages, figs., published by the authors. HALL, T. A. (1999): BioEdit: a user-friendly biological se- quence alignment editor and analysis program for Windows 95/98/NT. Nucl. Acids. Symp. Ser. 41: 95- 98. HIJMANS, R. J., GUARINO, L., CRUZ, M. & ROJAS, E. (2001): Computer tools for spatial analysis of plant genetic re- sources data: |. DIVA-GIS. Plant Genetic Resources Newsletter 127: 15-19. HOLLy, M. (1930): Synopsis der Süßwasserfische Kame- runs. Sitzungsberichte der Akademie der Wissenschaf- ten Wien (Mathematisch-naturwissenschaftliche Klas- se) 139, Abteilung I: 195-281. HUBER, J. H. (1977): Une chaine de deux Aphvosemion sympatriques dans les monts de Cristal, Gabon, avec description d'une espece nouvelle: A. (M.) mimbon n. sp.. Revue frangaise d'Aquariologie 4: 3-10. HUBER, J. H. (1980): Rapport sur la deuxieme expédition au Gabon (aoút 79). Etude des Cyprinodontidés récol- tés. Revue francaise d'Aquariologie 7: 37-42. HUBER, J. H. (1994): Aphyosemion passaroi, Espece iné- dite du Gabon Sud-oriental, au Patron de Colouration unique et Description complementaire de Aphyose- mion decorsei Pellegrin. Revue frangaise d'Aquariolo- gie Herpétologie 20: 77-79. HUBER, J. H. (1996): Killi-Data 1996. 2™ edition. Société Francaise d'Ichthyologie, Paris. HUBER, J. H (1998a): Aphyosemion alpha, n.sp. eme neue Art der Untergattung Chromaphyosemion mit einem ausgeprágtem Fárbungsmuster und einer besonders südlichen Verbreitung. Das Aquarium 32(350): 15-23. HUBER, J. H. (1998b): A new Cyprinodont species with a uniquely-coloured female, Aphyosemion hera n.sp. (Cyprinodontiformes, Pisces), from northwestern Ga- bon. Revue Suisse Zoologie 105: 331-338. HUBER, J. H. (1998c): Miscellaneous Notes on Some Sys- tematic Difficulties Regarding Old World Cyprino- donts. Journal of the American Killifish Association 31(1): 3-17, 28-32. HUBER, J. H. (1999): A new Species of Cyprinodont Fish, Aphyosemion tirbaki n. sp. (Cyprinodontiformes: Ap- locheilidae) from Gabon, with further Evidence of the Frontier Species Concept. Freshwater and Marine Aquarium (FAMA) 22(4): 104-118. HUBER, J. H. (2000 ): Killi - Data 2000. Société Francaise d'Ichthyologie, Paris (german edition). KUMAR, S.; TAMURA, K.; JAKOBSEN, I. B.& NEI, M. (2001): MEGA2: Molecular Evolutionary Genetics Analysis software, Arizona State University, Tempe, Arizona, USA. LAMBERT, J. G. & GERY, J. (1967): Poissons du Bassin de I'Ivindo. IH. Le genre Aphyosemion. Biologia Gaboni- ca 3: 291-315. LEGROS, O. (1999): Aphyosemion kouamense n. sp., ein neues Chromaphyosemion (Teleostei: Aplocheilidae) vom Rand der Monts de Cristal im Nordwesten Ga- buns. DKG-Journal 31: 32-40. MURPHY, W. J. & COLLIER, G. E. (1999): Phylogenetic Re- lationships of African Killifishes in the Genera Aphyo- semion and Fundulopanchax Inferred from Mitochon- drial DNA Sequences. Molecular Phylogenetics and Evolution 11: 351-360. RADDA, A. C. (1977): Vorláufige Beschreibung von vier neuen Subgenera der Gattung Aphyosemion. Aquaria 24: 209-216. RADDA, A. C. & HUBER, J. H. (1976): Cyprinodontiden- Studien in Gabun. I. Allgemeines - Nordwestgabun. Aquaria 23: 179-189. RADDA, A. C. & PURZL, E. (1976):Der Aphyosemion cameronense-Komplex. DKG-Journal 8: 131-144. RADDA, A. C. € PURZL, E. (1977): Cyprinodontiden- Studien in Gabun. II. Nordgabun. Aquaria 24: 21-31. SEEGERS, L. (1987): Was ist Aphyosemion escherichi? Die Aquarien und Terrarien Zeitschrift (DATZ) 40(12): 539-542. SEEGERS, L. (1988): Bemerkungen tiber die Sammlung der Cyprinodontiformes (Pisces: Teleoste1) des Zoologi- schen Museum Berlin. I. Die Gattungen Aphyosemion Myers, 1924 und Fundulosoma Ahl, 1924. Teil 2. Mit- teilungen des Zoologischen Museum Berlin 64(1): 3-70. SEEGERS, L. (1997): Killifishes of the world. Old World Killies I. Aqualog, A.C.S. GmbH, Mörfelden- Walldorf, Germany WILDEKAMP, R. H. (1993): A World of Killies. Atlas of the Oviparous Cyprinodontiform Fishes of the World. The American Killifish Association, Vol. 1, 1-311, figs. Authors’ addresses: Rainer SONNENBERG (correspond- ing author), Zoologisches Forschungsmuseum Alexan- der Koenig, Adenauerallee 160, D-53113 Bonn, Ger- many, e-mail: r.sonnenberg.zfmk@uni-bonn.de; Thomas BLUM, Weidenweg 4, D-88696 Owingen, Germany, e-mail: blum.thomas@t-online.de Bonner zoologische Beiträge Band 53 (2004) Heft 1/2 Seiten 221 (50) N 3onn, Juni 2005 Ambia naumanni sp. n., a New Species of Musotiminae from Yunnan (Lepidoptera, Crambidae)' Wolfgang SPEIDEL & Dieter STÜNING Zoologisches Forschungsmuseum Alexander Koenig, Bonn, Germany Abstract. Ambia naumanni sp. n. is described from Lijiang (Yunnan, China) and differentiated from related species. The following new combinations are proposed: Ambia colonalis (Bremer, 1864) comb. n. (described as Hydrocampa) and Ambia locuples (Butler, 1889) comb. n. (described as Oligostiema). 1. INTRODUCTION In the revision of the Palearctic Acentropinae (SPEIDEL 1984), all the species of the Palearctic part of China recognized to belong to this subfamily have been re- vised. However, one Chinese species — which was found to be undescribed — turned out to be misplaced in the Acentropinae and remained without an identification and name, as it was excluded from the study. It was rec- ognized to belong to another subfamily: Musotiminae. The Musotiminae were formerly included in the Nymphu- linae (= Acentropinae) (e.g. HAMPSON 1897; MUNROE 1972), but separated as an independant subfamily by SPEI- DEL (1981), a treatment which is now generally accepted (MUNROE & SOLIS 1999; SOLIS & MAES 2002). The authors found additional specimens of this musotimine species in the collection of the Alexander Koenig Research Institute and Museum of Zoology (ZFMK) and decided to describe the species as new, a plan which remained unreal- ised for a long time, because of the problem of the associa- tion with a reasonable genus. There is no revision of the subfamily in a world-wide scale, and there are several gen- era recognised in the subfamily, and a few more can still be found misplaced among other subfamilies. It finally be- came clear that the species belongs either to Musotima Meyrick, 1884 or to Ambia Walker, 1859. To our opinion, the present species fits quite well in the latter genus. How- ever, several obviously closely related species from Japan were treated in the genus Musotima (YOSHIYASU 1985) and so there remained some uncertainty about its generic position. Later we found that the figures of the genitalia of the generic type-species Ambia ptolycusalis Walker, 1859 by LANGE (1956) are incorrect. They represent a mis- identified acentropine species, a member of the genus Paracymoriza of the group of P. eromenalis (Snellen, 1880) and parallelalis Sauber, 1902. Therefore, we here provide a correct figure of the holotype of Ambia ptolv- 1 In commemoration of Clas Michael Naumann zu Königsbrück (26.06.1939 — 15.02.2004) cusalis and its genitalia from Sarawak, Borneo (Hope Entomological Collections, University Museum, Ox- ford) (Figs 8, 9). Both figures clearly show that 4. ptolycusalis represents another species and belongs to an entirely different subfamily. The wrong figure given by LANGE is probably the reason why YOSHIYASL (1985) did not use Ambia for any species treated in his revision. To our opinion, Musotima has to be syn- onymized with Ambia or at least a part of the species treated under Musotima by YOSHIYASU have to be trans- ferred to Ambia. We do not propose a formal syn- onymisation of Ambia and Musotima here, as the sys- tematics of the subfamily is presently in a process of development and changes: Very recently, two new gen- era are named and others are new defined in order to es- tablish monophyletic entities (SOLIS et al. 2004; YEN et al. 2004) which will eventually make it necessary to split the genus Ambia as well. Ambia ptolycusalis has a typical spine-like saccular process (see fig. 9) which does not occur in any species of the subfamily Acentropinae and which is not present in LANGE's misidentified figure. It is also found in 4m- bia colonalis (Bremer, 1864) comb. n. (described as Hydrocampa) and specimens which we associate with the description of Ambia locuples (Butler. 1889) comb. n. (described as Oligostigma). This spined sacculus is also found in other crambid subfamilies, e.g. in the Sco- pariinae. Therefore, it seems difficult to use this charac- ter as an autapomorphy of Ambia, especially as appar- ently closely related species lack this process. In the present new species, the saccular spine is absent, but there is a conspicous, angled process to the aedeagus, arising dorsally close to the insertion of the ductus ejaculatorius. We also found this apomorphic character in a species identified as Ambia locuples which is, on the other hand, provided with a saccular spine. The furcated aedeagus is a rather rare character in Ambia and absent in other Musotiminae, but it is weakly indicated also in the type-species A. prolvcusalis. How- ever, generic separation of species with such an unusual 222 Bonner zoologische Beitráge 53 (2004) Figs. 1-4: 1. Ambia naumanni sp. nov., holotype, male (Lijiang, 8. vin. 1934) (wing span 15 mm). 2. Ambia naumanni sp. nov., paratype, male (Lijiang, 13. vi. 1934) (wing span 16 mm). 3. Ambia thyridialis (Lederer, 1855), male (Ghazir) (wing span 15 mm). 4 . Ambia naumanni sp. nov., male paratype, lateral view of head. aedeagus-structure as found in Ambia locuples and the new species seems not advisable. We consider Ambia thyridialis (Lederer, 1855) (fig. 3), a species recorded from the Lebanon and Egypt which also has no saccular tooth as a closely related species. 2. DESCRIPTION OF SPECIES Ambia naumanni Speidel & Stiining, sp. nov. (Fig. 1, 2) Holotype @: “Li-kiang. (China). Provinz Nord- Yuennan. 8. 8. 1934. H. Höne” Paratypes: Males: dto. 1 4 13. 6. 1934; 1 & 19. 7. 1934; 14 2.8. 1935: 1 & 17. 7. 1934; 1 3 7. 8. 1934; 2 4 3.8.1935; 1 & 12.7. 1934; 1 4 7.7. 1934, GS-15036 d-SB; 1 8 17. 7. 1934, GU 137 8 SP12741934: Prep. 15378 SB and FU 160 SP; 27. 6. 1934, 1 Y 24. 6. 1935 with genitalia in glycerol; Females: 1 Y 3. 8. 1935; 1 Y 8. 9. 1935; 2 Y 3. 7. 1934 with FU 159 SP; 1 9 29. 6. 1934, GU 15063; 1 $ 15. 6. 1935; 1 Y “Li-kiang ca. 2000 m, Prov. Nord-Yuennan, 26. 6. 1934. H. Hóne” with genitalia in glycerol. Diagnosis (fig. 1, 2).- Head and body: Antenna com- paratively stout in the males, pubescent. Labial palpi rather long, upcurved, terminal segment almost reaching base of antennae. Frons smoothly scaled, whitish like the palpi. Thorax and abdomen greyish-brown, mixed with white, abdominal tergites with posterior white tranverse bands. Wolfgang SPEIDEL & Dieter STÚNING: Ambia naumanni sp. n., a New Species of Musotiminae from Yunnan 1mm = Fig. 5: Ambia naumanni sp. nov., male genitalia. Wings: Wingspan 15-17 mm, forewing length 7,5-8 mm. Wings with termen sinuous as usual in the genus, apices produced. Ground colour yellowish. Transverse fasciae prominent, white, bordered blackish-brown. The basal fascia strongly oblique, broken, the proximal one slightly curved, the distal one outcurved anteriorly and strongly retracted below middle, reaching the inner margin at about one half. The latter fascia is widened to a quadrangular spot at inner margin. Submarginal fascia conspicuous, widening towards tornus. Medial area with a white disco-cellular spot and another larger white spot in the anterior angle of the distal fascia. Marginal area yellow. Hindwing with similar pattern. Proximal fascia wider than in forewing, submarginal fascia strongly widening towards apex, incised at vein M2. Medial area suffused with brownish scales, like in the forewing, with two large white spots at the costal region and a small one at inner margin. Marginal area yellow. The venation (fig. 7) shows no principle difference to the other mem- bers of the genus. 1mm A DIE 2 026 Fig. 6: Ambia naumanni sp. nov., female genitalia. Male genitalia (Fig. 5): Vinculum large, with u-shaped saccus. Tegumen short, uncus with a broadly triangular base, otherwise slender, distinctly flattened ventrally, curved ventrad, pointed. Gnathos with strong lateral arms, medial process very narrow, slightly curved parallel to the uncus, minutely dentate dorsally, pointed at tip. Juxta large, with two dorsal spines situ- ated laterally at the posterior margin. Valva simple, broadening towards apex, without a saccular spine or other appendages, slightly ciliate. Aedeagus with a slender, sharply bent process, arising just distally to the opening of the ductus ejaculatorius. Shaft of aedeagus strongly incurved beneath this process. A curved ap- pendage originates at the inner side of the sinus. dorso- Female genitalia (Fig. 6): Ovipositor comparatively short, with moderately long apophyses. Ostium wide, with a bilobed sclerotized structure inside. Upper third of the ductus bursae sclerotized, with the ductus semi- nalis originating in the center of this region. Ductus bur- sae slightly broadening towards corpus, which is small and with a trace of a signum only. 224 Bonner zoologische Beitráge 53 (2004) Fig. 7: Ambia naumanni Sp. nov., wing venation. HOPE ENT. COLL. OXF. UNIY. MUS. GENITALIA NO.: 1309-83 HOPE ENT. COLL. OXF. UNIY. MUS. NO.: 1301-93 : rep. J.Minet Petey 54234 542 b HOPE ENT. COLL. OXF. UNIV. MUS. u. OF 1310-93 shee TYPE LEP: 909 mbia HOPE DEPT.OXFORD Fig. 8: Ambia ptolycusalis Walker, 1859. Holotype with labels (Sarawak, Borneo). The new species is very similar to the Western Palearc- tic Ambia thyridialis Lederer, which can easily be dis- tinguished by the more brownish ground colour of the wing surface, with yellowish scales only found in the marginal fasciae. Ambia thyridialis Lederer is also simi- lar in the male genitalia, lacking the saccular process of the valvae as well, but the aedeagus is completely dif- ferent. 0, 1mm 9 Fig. 9: Ambia ptolycusalis Walker, 1859. Male genitalia of holotype. Distribution and bionomics: Hitherto only known from the type-locality in SW. China, Yunnan, Yueling- shan near Lijiang, at elevations between 2800 and 3500 m. Flight-period from mid of June until beginning of September. First instars and foodplant unknown, but probably fern-feeding, as most of the Musotiminae are. Acknowledgments and etymology. The new species is named in honour of Prof. Dr. C. M. NAUMANN who im- proved the situation in the entomological department of our museum considerably during his directorship and facili- tated our work in many respects. We are grateful to Prof. Dr. K. Efetof and V. Saenko for the drawings of the genita- lia and to D. J. Mann (Hope Entomological Collections, Oxford) for the loan of the type slides of Ambia ptoly- cusalis. REFERENCES LANGE, W. H. (1956): A generic revision of the aquatic moths of North America (Lepidoptera: Pyralidae, Nymphulinae). Wasmann Journal of Biology 14(1): 59-144. MUNROE, E. & SOLIS, M. A. (1999): The Pyraloidea. pp. 233-256 in KRISTENSEN, N. P. (ed.): Lepidoptera, Moths and Butterflies. 1. Evolution, Systematics, and Biogeography. In FISCHER, M. (Ed.): Handbuch der Zoologie 4. Arthropoda: Insecta (35). Berlin, New York, 491 pp. Wolfgang SPEIDEL & Dieter STUNING: Ambia naumanni sp. n., a New Species of Musotiminae from Yunnan 225 SOLIS, M. A. & MAES, K. V. N. (2002): Preliminary phy- logenetic analysis of the subfamilies of Crambidae (Pyraloidea Lepidoptera). Belgian Journal of Ento- mology 4: 53-95. SOLIS, M. A., YEN, S.-H. & GOOLSBY, J. H. (2004): Spe- cies of Lygomusotina new genus and Neomusotima Yoshiyasu (Lepidoptera: Crambidae) from Australia and Southeastern Asia feeding on Lygodium micro- phyllum (Schizaeaceae). Annals of the Entomological Society of America 97(1): 64-76. SPEIDEL, W. (1981): Die Abgrenzung der Unterfamilie Acentropinae. Atalanta 12: 117-129, SPEIDEL, W. (1984): Revision der Acentropinae des paläarktischen Faunengebietes (Lepidoptera, Cram- bidae). Neue Entomologische Nachrichten 12, 157 pp.. including 3 colour plates. HAMPSON, G. F. (1897): On the classification of two sub- families of moths of the family Pyralidae: the Hydro- campinae and Scoparianae. Transactions of the Ento- mological Society of London 1897: 127-240. MUNROE, E., in DOMINICK, R. B., et al. (1972-3): The Moths of America North of Mexico 13. 1. Pyraloidea (Pyralidae part.). London (E. W. Classey), 304 pp., pl. 1-13 and A-K. YEN, S.-H., SOLIS, M. A. & GOOLSBY,:J. A. (2004): Aus- tromusotima, a new Musotimine genus (Lepidoptera: Crambidae) feeding on Old World Climbing Fern, /2- godium microphyllum (Schizaeaceae). Annals of the Entomological Society of America 97(3): 397-410. YOSHIYASU, Y. (1985): A systematic study of the Nym- phulinae and the Musotiminae of Japan (Lepidoptera: Pyralidae). Scientific Reports of the Kyoto Prefectural Univiversity (Agriculture) 37: 1-162. Authors’ addresses: Dr. Wolfgang SPEIDEI (w.speidel.zfmk@uni-bonn.de.) & Dr. Dieter STUNING (corresponding author) (d.stuening.zfmk@uni-bonn.de). Alexander Koenig Research Institute and Museum of Zoology, Adenauerallee 160, D-53113 Bonn, Germany. Bonner zoologische Beitráge Band 53 (2004) | Heft 1/2 | Seiten 227-234 | Bonn, Juni 2005 A New Paracymoriza Species from Lombok (Indonesia) (Lepidoptera, Crambidae) ' / Wolfgang SPEIDEL”?, Ulf BUCHSBAUM” & Michael A. MILLER” " Zoologisches Forschungsmuseum Alexander Koenig, Bonn (Germany) 2 Zoologische Staatssammlung, Munich (Germany) Abstract. A new species Paracymoriza naumanniella sp. nov. from Lombok island is described and assigned to the Paracymoriza albifascialis species group. Moreover, data on the egg structure, habitat, and behaviour of the adult moths are given. In addition, DNA sequence information of mitochondrial cytochrome oxidase subunit | are furnished, which constitute the first report of genetic data concerning this gene within the family Crambidae. Key words: Acentropinae, Paracymoriza naumanniella sp. nov., taxonomy, mtDNA, COI 1. INTRODUCTION The genus Paracymoriza Warren, 1890 is a species rich genus within the family Crambidae, belonging to the superfamily Pyraloidea (Pyralid moths). Within the Crambidae, it is a member of the subfamily Acentropi- nae (former Nymphulinae, aquatic moths). In the past, Paracymoriza has been treated as a synonym of Parthenodes Guenée, 1854 which is an unrelated genus which has, in contrast, an exclusively new world distri- bution. Paracymoriza comprises 28 currently recog- nized species in the Oriental Region (SPEIDEL & MEY 1999; SPEIDEL 2003). The new species occurs in Lom- bok, Indonesia and seems to be closely associated with humid ecosystems where the specimens of the type se- ries have been found. 2. TAXONOMY Paracymoriza naumanniella sp. nov. (Figs. | A, 1 B, 2 a, 2 b) Material: Holotype ©: “Indonesia, North Lombok, Gunung Rinjani NP, Air terjun, near village Senaru, ca. 500 - 600 m, 08°18' S / 116°24' E, 28. Dezember 2003, TF, leg. Mei-Yu Chen & Ulf Buchsbaum”. “DNATAX 02820”; genitalia slide no.: M3450 Zoologische Staats- sammlung Miinchen (Munich, Germany). 11 paratypes: same data, one y with genitalia slide no. M3451 in Zoologische Staatssammlung Múnchen, 2 paratypes in Museum fúr Naturkunde, Berlin (one of them with additional label “DNATAX02824”) and 2 paratypes in coll. W. Speidel, Bonn (one with additional label “DNATAX02822”). 1 In commemoration of Clas Michael Naumann zu Königsbrück (26.06.1939 — 15.02.2004) DNA is stored in the frozen DNA collection of the ZSM under storage numbers DNATAX02820 (holotype), DNATAX02821 (paratype), DNATAX02822 (para- type), DNATAX02823 (paratype), DNATAX02824 (paratype), and DNATAX02825 (paratype). Description: Length of forewing: 4 9 — Ilmm, @ 10 mm; Y 9— 11 mm, 9 10,2 mm. Wingspan: 6 17—22 mm, 6 17,8 mm; Y 19 — 23 mm, @ 21 mm. Body dark grey, abdomen with white intersegmental rings. White ground-colour of the wings strongly re- duced, therefore pattern not very contrasting. Forewing: Inner, medial and submarginal areas dark grey, basal area almost black. Basal, proximal and distal fascia narrow, whitish. Distal fascia strongly incurved close to inner termen, with a dent directed towards the termen on vein CuP. Some marginal white spots along the termen, fringe blackish. Hindwing: In contrast to other small Paracymoriza spe- cies, the wing pattern is clearly visible, though the me- dial area is strongly suffused like in the forewing. Basal. proximal and distal fascia narrow, whitish. The distal fascia with outwardly directed dent. Discal spot black, very close to proximal fascia. ' genitalia (Fig. 3): Valva distally rounded, elongate, reminiscent of the other members in the genus. however only with two large, acute inwardly directed. apical, se- tae and several smaller hairs along the distal margin. Uncus elongate, rounded towards end, gnathos slender with very inconspicuous teeth. Aedeagus comparatively strong, without cornuti, bulbus ejaculatorius arising from this center. 228 Bonner zoologische Beitráge 53 (2004) 05cm B Fig. 1: Paracymoriza naumanniella sp. n. Holotype (male) (A) and one Paratype (female) (B). Indonesia, North Lombok, Gu- nung Rinjani NP, Air terjun, near village Senaru, ca. 500 - 600 m, 08°18' S / 116°24' E, 28. Dezember 2003, TF, leg. Mei-Yu Chen & Ulf Buchsbaum. Fig. 2a-b: Paratypes of the new species still alive at the type locality in Lombok (see above). Photo: Ulf Buchsbaum. Y genitalia (Fig. 4): Ovipositor short, with long, slen- der, fragile apophyses. Ductus bursae rather long, gradually widening towards corpus bursae, with an un- usual evagination about in its middle. Corpus bursae round, ball-shaped, without a clear signum. Colliculum situated terminally, with the slender ductus seminalis originating inside and in a short distance of the colli- culum. Ostial region strongly sclerotized, large, cup- shaped, narrowing strongly towards the ductus bursae. Egg (Fig. 7a, b): Oval, with micropylar zone at the nar- row end of the egg (size: 310 x 180 pm). Distribution (maps, Figs. 5a and 5b): Hitherto only known from the type locality in Lombok. Genetic data: We sequenced the mitochondrial gene cytochrome oxidase subunit I in order to provide a mo- lecular dataset for subsequent re-identification of the new species as recommended by TAUTZ et al. (2003). This gene has proven its usefulness for DNA taxonomy purposes in many studies (e.g. HEBERT et al. 2003). Genetic characterization: Method: DNA was extracted from abdominal tissue of some of the type specimens of P. naumanniella sp. nov. using Qiagen (Hilden, Ger- many) tissue kit according to the protocol of KNOLKE et al. (2005). Mitochondrial (mtDNA) cytochrome ox1- dase subunit I (COI) gene was amplified with PCR us- ing protocols and primers as in SIMON et al. (1994). Di- rect sequencing of dye labelled templates was carried Wolfgang SPEIDEL, Ulf BUCHSBAUM & Michael A. MILLER: A New Paracymoriza Species from Lombok (Indonesia) 229 0,1 mm Fig. 3: Holotype male genitalia. out using an ABI 377 automated sequencer (Applied Biosystems). Up to 10 single sequences per individual were assembled and aligned to the COI gene of Bombyx mori (L., 1758) (Lepidoptera, Bombycidae; GenBank accession number NC 002355). Result: The following sequence of P. naumanniella sp. nov. shows the partial COI gene of the holotype: >Paracymoriza naumanniella Holotype _ DNATAX02820 1230bp CGAAAATGACTTTATTCTACAAATCATAAAGAT ATTGGAACTTTATATTTTATTTTTGGAATTTGAT CGGGAATAGTTGGAACATCTTTAAGATTATTAA TTCGAGCTGAATTAGGAAATCCTGGATCATTAA TTGGAGATGATCAAATTTATAATACTATTGTTA CTGCETCATGCTTTTATTATAATTTTTTTTATAGT AATGCCAATTATAATTGGAGGATTTGGAAATTG ATTAGTTCCATTAATATTAGGTGCTCCTGATAT GGCTTTCCCACGAATAAATAATATAAGATTTTG ACTIETTCCTCCATCICTTACACTTTTAATTTCA AGAAGAATCGTAGAAAATGGAGCAGGAACAGG ATGAACAGTGTACCCCCCACTATCATCTAATAT CGCTCATGGAGGAAGATCTGTAGATTTAGCTAT TTTTTCCTTACATTTAGCTGGTATTTCCTCAATT TTAGGAGCAATTAATTTCATTACTACAATTATT AACATACGAATTAATAATTTATTTTTTGATCAA ATACCTCTTTTTATTTGAGCGGTAGGTATTACA GETTITACTICTTCITCITICITIACCAGTTITAGC TGGAGCAATTACTATGTTATTAACAGATCGAAA TTTAAATACATCTTTTTTTGATCCTGCTGGAGGA 0,5 mm Fig. 4: Paratype female genitalia. GGAGATCCAATTTTATATCAACATTTATTTTGAT TTTTTGGGCATCCCGAAGTATATATTTTAATTTI ACCAGGATTTGGAATAATCTCTCATATTATTTCT CAAGAAAGAGGAAAAAAAGAAACTTTTGGATC TTTAGGAATAATTTATGCTATAATAGCAATTGG ATTATTAGGATTTGTTGTTTGAGCACATCATAT ATTTACTGTAGGTATAGATATTGATACTCGAGC ATATTTTACTTCTGCAACTATAATTATTGCTGTA 230 Bonner zoologische Beitráge 53 (2004) CCAACAGGAATTAAAATTTTTAGATGATTAGCA ACCTTACATGGAACTCAAATTAATTATAGACCT TCTACTTTATGAAGATTAGGATTTGTATTTTTAT TTACTGTAGGGGGATTAACTGGAGTTGTTTTAG CTAATTCTTCAATTGATGTAGCTCTTCATGATAC TTATTATGTAGTAGCACATTTTCATTATGTTCTA TCTATAGGAGCAGTATTTGCTATTTTAGGAGGT - TTTATTCATTGATATCCATTATTTACAGGAT- TAACTCTTAAACCTTATTACTTAAAAATTCAA The complete COI gene of the paratypes is shown for DNATAX02821: >Paracymorıza naumanniella Paratype _ DNATAX02821_1530bp CGAAAATGACTTTATTCTACAAATCATAAAGATA TTGGAACTTTATATTTTATTTTTGGAATTTGATCG GGAATAGTTGGAACATCTTTAAGATTATTAATTC GAGCTGAATTAGGAAATCCTGGATCATTAATTGG AGATGATCAAATTTATAATACTATTGTTACTGCT CATGCTTTTATTATAATTTTTTTTATAGTAATGCC AATTATAATTGGAGGATTTGGAAATTGATTAGTT CCATTAATATTAGGTGCTCCTGATATGGCTTTCCC ACGAATAAATAATATAAGATTTTGACTTCTTCCT CCATCTCTTACACTTTTAATTTCAAGAAGAATCG TAGAAAATGGAGCAGGAACAGGATGAACAGTGT ACCCCCCACTATCATCTAATATCGCTCATGGAGG AAGATCTGTAGATTTAGCTATTTTTTCCTTACATT TAGCTGGTATTTCCTCAATTTTAGGAGCAATTAA TTTCATTACTACAATTATTAACATACGAATTAAT AATTTATTTTTTGATCAAATACCICTTTTTATTTG AGCGGTAGGTATTACAGCTTTACTTCTTCTTCTTT CTTTACCAGTTTTAGCTGGAGCAATTACTATGTT ATTAACAGATCGAAATTTAAATACATCTTTTTTT GATCCTGCTGGAGGAGGAGATCCAATTTTATATC AACATTTATTTTGATTTTTTGGGCATCCCGAAGTA TATATTTTAATTTTACCAGGATTTGGAATAATCTC TCATATTATTTCTCAAGAAAGAGGAAAAAAAGA AACTTTTGGATCTTTAGGAATAATTTATGCTATA ATAGCAATTGGATTATTAGGATTTGTTGTTTGAG CACATCATATATTTACTGTAGGTATAGATATTGA TACTCGAGCATATTTTACTTCTGCAACTATAATTA TTGCTGTACCAACAGGAATTAAAATTTTTAGATG ATTAGCAACCTTACATGGAACTCAAATTAATTAT AGACCTTCTACTTTATGAAGATTAGGATTTGTAT TTTTATTTACTGTAGGGGGATTAACTGGAGTTGT TTTAGCTAATTCTTCAATTGATGTAGCTCTTCATG ATACTTATTATGTAGTAGCACATTTTCATTATGTT CTATCTATAGGAGCAGTATTTGCTATTTTAGGAG GTTTTATTCATTGATATCCATTATTTACAGGATTA ACTCTTAAACCTTATTACTTAAAAATTCAATTTAT TACAATATTTATTGGAGTAAATTTAACTTTTTTCC CACAACATTTTTTAGGTTTAGCTGGAATACCACG ACGATATTCTGATTATCCTGACATTTATATTTCAT GAAATATTATTTCTTCATTAGGATCTTATATTTCA TTATTAGCAATTATAATAATAATAATAATTATTT GAGAATCAATAATTAACCAACGAATTATTTTATT TTCATTAAATTTATCATCTTCTATTGAATGATATC AAAATTTACCCCCTGCAGAACATTCATATAAT- GAACTTCCTATTTTAAGAAAT Species variation: In addition to the holotype we se- quenced four of the paratypes. The sequence variation within the set of types is shown in Table 1. There are six variable sites within the complete sequences of COI of the paratypes and five variable sites within all type specimens due to the reduced sequence lengths avail- able for comparison (1.230bp). Therefore the species variation is 0.39% sequence divergence within the com- plete gene and 0.41% in the short dataset. Table 1: Variable sites and positions. Abbreviation: n.a. - sequence data not available. varable sites: positions COI 69 309 351 708 1113 1283 DNATAX02820 (holotype) G T TG A na. DNATAX02821 (paratype) G DNATAX02823 (paratype) G o» © a A A G A HA 8 HHO 8 dE ab DNATAX02824 (paratype) A C DNATAX02825 (paratype) G T The new sequences are deposited under the accession numbers AJ 852519 (DNATAX02820, holotype), AJ 852520 (DNATAX02821, paratype), AJ 852521 (DNATAX02823, paratype), AJ 852522 (DNATAX 02824, paratype), and AJ 852523 (DNATAX02825, paratype), respectively, at EBI/GenBank. Remarks on the molecular studies. The aim is to provide an additional character set in the diagnosis of the new species. There is at the moment no possibility to include related species in a differential diagnosis on DNA se- quence level due to the missing COI data of other spe- cies of the family Crambidae. Biological data: The imagines of the new species are so far found only in December. The biotope is a primary rain forest, near a waterfall, with very high humidity due to the mist. The moths rest on the algae covered sides of stones and rocks situated close to or in the water. Eventually they jump onto the surface of the water and afterwards back to the rock. The moth are highly hydrophobic, can be completely submerged under waves, but are not trapped by the surface-tension of the water. They seem to be very connected to their restricted habitat and to leave it very rarely and only within little distances. Wolfgang SPEIDEL, Ulf BUCHSBAUM & Michael A. MILLER: A New Paracymoriza Species from Lombok (Indonesia) 23] A «SUNDA-INSELN . Javasee ‘ a GROSSE SUNDA-IN SEUWN 5 , ; 1 | : | 19 PAZIFISCHER A OZEAN MA 115S CH | ver AR CoH PI % + INDISCHER OZEAN Fig 5: Type locality. 5 a: Enlarged view to show the situation of the Island Lombok (Indonesia) east of Bali: 5 b: type locality of P. naumanniella on Lombok Island near the village Senaru below the mountain Gunung Rinjani. 159) Ww N Bonner zoologische Beitráge 53 (2004) Fig. 6: Biotope at type locality beside the waterfall. 6 a: Here, most moths rest under the rocks near the water surface. Photo: Buchsbaum; 6 b: One of the authors (U. B.) collecting specimens of the new species. Photo: Mei-Yu Chen. Ulf Wolfgang SPEIDEL, Ulf BUCHSBAUM & Michael A. MILLER: A New Paracymoriza Species from Lombok (Indonesia) 233 | Paracymoriza naumanniella Vergrößerung= 250X Paracymoriza naumanniella Vergrößerung= 910X Fig. 7: SEM photos of the egg of Paracymoriza naumanniella. 7 a: Egg (total view); 7 b: surface structure of the egg with large! magnification. 234 Relationships: The closest relatives of the new species can only be ascertained after a revision of the genus. A similar wing-pattern is found in the P. albifascialis Hampson, 1891 species group (SPEIDEL & MEY 1999), especially in the strongly toothed distal fascia. It is therefore tentatively placed in this species group, but differs from all other members of the group by the in- fuscated medial area of both wing-pairs and the small size. Differences from the Paracymoriza distinctalis (Leech, 1889) group include the slender wing-shape (broader in the P. distinctalis species group) and the size (larger in the P. distinctalis species group). Members of the P. nigra (Warren, 1896) species group show the same small size, however the pattern is almost absent from the hindwing. Etymology: The species is named in honour of the late Prof. Dr Clas M. NAUMANN, one of the most important contemporary lepidopterists who has successfully intro- duced and used the methods of Phylogenetic Systemat- ics in lepidopterology. 3. DISCUSSION The approach to include genetic data in new species de- scriptions is facing more and more acceptance within the scientific community. Proponents of the so-called ‘DNA taxonomy’ plea for the use of genetic markers as a unique determination tool (TAUTZ et al. 2003). The question which gene to use is tending to the well investigated COI gene of the mitochondrion (HEBERT et al. 2003). Critics of the choice refer to the risk of misleading results due to the analysis of paralogous sequences because of independent evolving nuclear insertions of mitochondrial genes (THALMANN et al. 2004). Nevertheless these rare excep- tions should not keep taxonomists from producing genetic markers of newly described species. There is a major chance in the offering of genetic markers: Unknown imma- ture stages can easily be associated with adult descriptions. Breeding of aquatic moths in the field is rather difficult and in the present case the early stages of P. naumanniella sp. nov. are unknown. The genetic data will enable to associate larvae in the future. The new species may have aquatic lar- vae, which probably live in the current water of small creeks close to the locations where the imagines were found. The assignment of unknown larvae to the species they belong, is only possible by analysis of their genome when rearing of the caterpillars is not feasible. Prof. Dr. Clas M. NAUMANN had been aware of the chances genetic markers offer for taxonomy. In his late years he hardly hesitated to become a proponent of DNA taxonomy, and his main focus still laid on the phylogenetic aspect which genetic information of taxa provides. Bonner zoologische Beitrage 53 (2004) Acknowledgements. We thank Dr Axel Hausmann (Mu- nich) for his discussion about the manuscript. We are grateful to Mei-Yu Chen (Meifeng, Taiwan) for her help in the field-work and photographic assistance and Dr. Wolf- ram Mey (Berlin) kindly allowed the use of the large refer- ence collection in the Museum fiir Naturkunde, which is especially rich in South East Asian material. Special thanks also to Enrico Schwabe (Munich) for his kind help with SEM photos. REFERENCES HEBERT, P. D. N., CYWINSKY, A., BALL, S. L. & DEWAARD, J. R. (2003): Biological identifications through DNA barcodes. Proceedings of the Royal Society London Series B 270(1511): 313-321. KNOLKE, S., ERLACHER, S. I., MILLER M. A., HAUSMANN, A. & SEGERER, A. H. (2005): Providing data for both morphology- and DNA-based taxonomy: A combined procedure for genitalia dissection and DNA extraction in Lepidoptera. Insect Systematics and Evolution 35: 401-409. SIMON, C., FRATI, F., BECKENBACH, A., CRESPI, B., LIU, H. & FLOCK, P. (1994): Evolution, weighting and phy- logenetic utility of mitochondrial gene sequences and a compilation of conserved polymerase chain reaction primers. Annals of the Entomological Society of America 87(6): 651-701. SPEIDEL, W. & MEY, W. (1999): Catalogue of the Oriental Acentropinae (Lepidoptera, Crambidae). Tijdschrift voor Entomologie 142: 125-142. SPEIDEL, W. (2003): New species of aquatic moths from the Philippines (Lepidoptera, Crambidae). Insecta Ko- reana 20: 7-49. TAUTZ, D., ARCTANDER, P., MINELLI, A., THOMAS, R. H., & VOGLER, A. P. (2003): A plea for DNA taxonomy. Trends in Ecology and Evolution 18: 70-74. THALMANN, O., HEBLER, J., POINAR, H. N., PAABO, S. & VIGILANT, L. (2004): Unreliable mtDNA data due to nuclear insertions: a cautionary tale from analysis of humans and other great apes. Molecular Ecology 13: 321-339, Authors’ addresses: Dr Wolfgang SPEIDEL (correspon- ding author), Zoologisches Forschungsmuseum Alexan- der Koenig, Adenauerallee 160, D-53113 Bonn, Ger- many; e-mail: w.speidel.zfmk@uni-bonn.de; UIf BUCHSBAUM, Zoologische Staatssammlung München, Münchhausenstraße 21, D-81247 München, Germany; e-mail: UlfBuchsbaum.Lepidoptera@zsm.mwn.de; Mi- chael A. MILLER, Zoologische Staatssammlung Mün- chen, Münchhausenstraße 21, D-81247 München, Ger- many; e-mail: miller@zsm.mwn.de Bonner zoologische Beitráge | Band 53 (2004) Heft 1/2 Seiten 235-254 Bonn, Juni 2005 Adult Morphology and the Higher Classification of Bia Hübner (Lepidoptera: Nymphalidae) ' / Richard I. VANE-WRIGHT ” & Michael BOPPRÉ ” ; ” Department of Entomology, The Natural History Museum, London, UK ? Forstzoologisches Institut, Albert-Ludwigs-Universität, Freiburg i. Br., Germany Abstract. The South American nymphalid Bia Húbner, 1819, treated for over 150 years by most lepidopterists as a member of the Satyrinae, has been shown by recent work on early stages and DNA to share characters with the Morph- inae: Brassolini. Examination of the wing patterns and androconial organs of Bia, described in detail for the first time, reveals unusual features otherwise only known from brassolines. In particular, the tufted posterior androconial organ of the hindwing forming palisades is a synapomorphy for Bia and several genera of Brassolini, including Caligo. The ge- nus Bia is formally transferred from the Satyrinae to the Morphinae: Brassolini as the sole member of the subtribe Biina Herrich-Scháffer, 1864, stat. nov., co-ordinate with Brassolina Boisduval, 1836, and Naropina Stichel, 1925. Key words. Systematics, Satyrinae, Morphinae, Brassolini, Ca/igo, androconia, Neotropics, butterflies The brothers Kratos and Zelos, and their sisters Nike and Bia, were the personifications of strength, rivalry, victory and force. These four winged gods stood beside the throne of Zeus 1. INTRODUCTION The genus Bia Hübner, 1819, has long been a puzzle to systematists. At present only one species is recognised, Bia actorion (Linnaeus, 1763). However, our investiga- tions and those of Gerardo LAMAS (pers. comm., Lima 2004) indicate that there may be two or more sibling species, and this will be addressed in a future paper (LAMAS, BOPPRE, HOARE & VANE-WRIGHT in prep.). In general facies Bia is markedly divergent from other butterflies, and both sexes are instantly recognisable as members of the genus (Figs 1-4). The butterflies are re- stricted to lowland and lower-montane forests of South America, occurring in dense undergrowth where the canopy is not fully closed. Individuals fly along trails and in clearings in damp, marshy areas, and are active from dawn to dusk. They often settle on rotting fruits, on which they feed, or on vegetation about a metre above the ground, when they may reveal their yellowish and blue upper side pattern. Mark and recapture studies indicate that males can live for at least 20 days, and are generally loyal to a particular patch of forest. In flight, the iridescent blue patches sported by most individuals flash conspicuously. If threatened, the butterflies dash into the base of a bush, where they are very difficult to reach. As they settle after an escape flight, the wings are closed to reveal only the cryptic, ripple-pattern under- side (Fig. 5). This, together with the sudden disappear- 1 In commemoration of Clas Michael Naumann zu Kénigsbriick (26.06.1939 — 15.02.2004) http://www.theoi.com/Ouranos/Kratos.htm! ance of the distinctive blue colour, makes them difficult to locate. Interactions between males are frequent, and courtship is lengthy and complex, including tandem flight patterns, contact during flight, and male flight over perched females that apparently may respond by flashing their wings. Until very recently their early stages and host plants were unknown (BARTLETT 1876: HALL 1939; MASTERS 1970; FREITAS et al. 2002; Keith WILLMOTT, pers. comm., London 2004). Bia adults have tri-carinate antennae and small forelegs in both sexes, and the genus undoubtedly belongs to the Nymphalidae sensu ACKERY et al. (1999). Its systematic position within the family has, however, been very un- certain. With no convincing evidence to support MILLER's (1968) suggestion of a relationship to the melanitine Satyrinae, D'ABRERA (1988: 846: 2001: 340), for example, has continued to locate Bia amongst the Pronophilina, the dominant group of typical Satyr- inae found in mountainous regions of South America. This reflects a convention first adopted by KIRBY (1871) in his 'Catalogue', and subsequently followed by WEY- MER (1911: 276) in 'Seitz', and by GAEDE (1931: 524) in 'Lep. Cat.'. MIELKE & CASAGRANDE (1998) list the Biini immediately after the Pronophilini. FREITAS & BROWN (2004). in contrast. conclude that Bia should be placed as a monobasic subfamily (Biinae) within their “satyroid clade”, reflecting a view going back to HERRICH-SCHAFFER (1864) that gives Bia very high taxonomic rank. In most of their analyses, Bia ap- peared in various relationships with the Satyrinae, Mor- Bonner zoologische Beitráge 53 (2004) 236 Richard I. VANE-WRIGHT & Michael BopPrÉ: Adult Morphology and the Higher Classification of Bia Hübneı 23 phini, Brassolini and Calinaginae, although in a succes- sive weightings analysis it appeared as the stem group of the Brassolini (FREITAS & BROWN 2004: fig. 2). Re- cent publications by BROWER (2000) and FREITAS et al. (2002) have provided, respectively, valuable new data on the molecular systematics and early stages that are consistent with the idea that Bia is a member of the Brassolini, one of the two South American tribes that belong to the Morphinae. This view, that Bia is a bras- soline, was first put forward by CLARK (1947), tenta- tively supported by DEVRIES et al. (1985), and recently accepted by YOSHIMOTO (2003). Here we re-investigate the adult morphology of Bia and question why its membership of the Brassolini was not recognised previously. CLARK (1947, 1948) failed to provide any evidence, and inaccurate or incomplete sub- sequent work has obscured its natural relationships. The peculiarities of the androconial systems reported here demonstrate that, even without the evidence now avail- able from knowledge of DNA sequences and early stage morphology, the clear relationship of Bia to the owl but- terflies (Caligo Húbner, 1819) and other Brassolini has literally been “staring us in the face” for over 200 years. 2. SYSTEMATIC HISTORY LINNAEUS (1763a,b) described Papilio actorion from “Indiis”, for which HUBNER (1819: 51) introduced the genus Bia, with Papilio actoriaena Hübner, 1819 (an objective synonym of Papilio actorion: HEMMING 1964), as the only included species. GODART (1824: 446), however, consigned P. actorion to Morpho Fabri- cius, 1807, in which he also included many species now placed in the Amathusiini and Brassolini. In his outstanding contribution to The Genera of Diur- nal Lepidoptera, WESTWOOD (1850: 321) accepted Hübner's genus for actorion, noting Bia as a “very inter- esting ... butterfly” belonging to the “Nymphalidae”. It must be appreciated, however, that Westwood's classifi- cation of the Nymphalidae differed significantly from current practice. He likened Bia not only to various but- terflies in the “Satyridae”, but in particular among his “Nymphalidae” to such genera as Siderone Hübner, 1823 (now in Charaxinae), Heferopsis Westwood, 1850 (now Satyrinae), Kallima Doubleday, 1849 (Nymphal- inae) and Amathusia Fabricius, 1807, Zeuxidia Hübner, 1826, and Discophora Boisduval, Amathusiini). 1836 (Morphinae In contrast, HERRICH-SCHAFFER (1864) suggested that Bia should have very high taxonomic-rank, placing it as the sole member of a new family, the Biidac [as "Bi- ina”], one of just 16 family groups into which he di- vided the entire Rhopalocera. In so doing, he compared Bia with butterflies now placed in the Brassolini and Danaini, but not the Satyrinae. As noted by WESTWOOD (1850), Bia has the bases of forewing veins Sc, Cu and 2A conspicuously inflated. This apparently persuaded WALLACE (1854) to place Bia in the Satyridae: “the beautiful Bia Actorion, which, though classified with the Nymphalidae, exactly agrees with this family [Satyridae] in its haunts and mode of flight ... [and] in many structural points.” Wallace con- cluded that it formed “a very satisfactory link connect- ing the two families.” In assigning Bia to the satyrines, he has been followed by a majority of lepidopterists ever since — e.g. FELDER (1861), DIETRICH (1862), KIRBY (1871), DRUCE (1876), MULLER (1877), STAUD- INGER (1888), SCHATZ € ROBER (1889), WEYMER & MAASSEN (1890), WEYMER (1911), GAEDE (1931), HALL (1939), EHRLICH (1958), HAYWARD (1958, 1964), FORSTER (1964), MILLER (1968), D'ABRERA (1988, 2001), HARVEY (1991), MIELKE & CASAGRANDI (1998), RACHELI & RACHELI (2001). However, at least two authors before the recent period linked this curious little butterfly firmly with the Morphinae — but in differ- ent ways. REUTER (1896), in his remarkable but often neglected thesis, placed Bia in the Morphinae, as one of three tribes: Morphini, Amathusiini and Biini (GODART 1824, by including actorion as a discrete subgroup “IIA” within Morpho, set a precedent for this). REUTER sepa- rated the Morphinae from the Brassolini, including the latter within the much larger Satyrinae. On the other hand, and possibly taking a lead from HERRICH- SCHAFFER, CLARK (1947, 1948) unhesitatingly placed Bia as a brassolid, but without giving any reason. Frus- tratingly, in the first of these two papers, CLARK stated confidently but without explanation, “Brassolidae [are] easily recognisable by adult characters”; no justification at all was given in his second paper. For various rea- sons, both REUTER (1896, 1898) and CLARK (1947, 1948) have largely been ignored. Figs 1-9: Bia actorion (L.) sensu lato. Adult butterflies and wing venation (both specimens from Suapure, Venezuela). 1 male upperside (upper arrow: anterior alar organ; lower arrow: posterior alar organ; BMNH(E) #693091): 2 female upperside (BMNH(E) #693105); 3 underside of | (upper arrow: border ocellus in forewing cell M,: mid arrow: border ocellus in hindwing cell Rs; lower arrow: border ocellus/diagonal white stripe in hindwing cell Cu;,); 4 underside of 2: 5 live individual at rest (Vene- zuela, Bolivar State, Jasper Falls 27.x.2000); 6 forewing radial venation from anterior apex of discal cell to separation of Ry and Rs (subcostal and radial veins labelled; BMNH electron micrographs #E3/273—5, composite SEM): 7 detail of 6 to show origin of radial veins from discal cell; 8 hindwing precostal area (upper arrow: precostal vein; lower arrow: precostal cell: BMNH electron micrograph #E3/270, SEM); 9 detail of 8 (BMNH electron micrograph #E3/271). Scale bars: 1-4: 10 mm: 6: | mm: 8-9: | mm. 7: 0.5 mm: i) Ww MILLER (1968) followed conventional wisdom in ac- cepting Bia as a member of the Satyridae (he regarded them as a family) in which, like Reuter, he also included the Brassolini but not Morphini or Amathusiini. MILLER used the name Biinae to designate one of seven subfam- ily divisions for the group, and further subdivided the Biinae into three named tribes: the Melanitini, Antir- - rheini, and the monobasic Biini, commenting that “Bia is far too aberrant to be referred to either of the other two biine tribes.” He also suggested that “within the Sa- tyridae the brassolines are allied to the New World Bi- inae, particularly through such genera as Narope [Brass- oliniJ” (MILLER 1968: 23). Miller thus united Bia with the new world Antirrheini (Antirrhea Hübner, 1822, and Caerois Hübner, 1819), and the old-world Melanitini: Melanitis Fabricius, 1807, Cvllogenes Butler, 1868, Gnophodes Westwood, 1851, Parantirrhoea Wood-Mason, 1880, and Bletogona C. & R. Felder, 1867 (for placement of this last genus, see UÉMURA 1987). In addition, but in a very ambiguous manner, Miller also listed Manataria Kirby, 1908, at the end of his account of the Biinae. This peculiar genus represents a small group of South American brown but- terflies of very uncertain affinity that he likened to the old world Elymniini: Lethina, as well as some members of his Biinae. In contrast, FORSTER (1964; see also RACHEL] & RACHELI 2001) had earlier placed Mana- taria within the Satyrini: Euptychima, the dominant group of lowland Satyrinae found in Latin America. MIELKE & CASAGRANDE (1998) understandably listed Manataria at the end of the Satyrinae as “tribe uncer- tain”. VANE-WRIGHT (1972a) recognised that the three higher taxa linked within the Biinae by MILLER (1968) repre- sent an unnatural assemblage. Based on evidence from eggs, larvae and adults, DEVRIES et al. (1985) formally transferred the Antirrheini to the Morphini, as a sub- tribe. They also suggested that Bia, mainly on the evi- dence of its external abdominal androconia similar to those found in Caligo and related genera, might belong to the Brassolini, and these views were echoed by ACK- ERY (1984: 16, 1988: 104). In BROWER's (2000) mo- lecular investigation, Bia grouped with Caligo, and these two genera then grouped with Opsiphanes Doub- leday, 1849, a result consistent with CLARK's assertion and the suggestion of DEVRIES et al. This contention is further supported by the work of FREITAS et al. (2002) on the early stages. Currently, of the subgroups included by MILLER in the Biinae, only the evening browns and their relatives of the Old World tropics (Melanitini), together with the peculiar New World Manataria, appear to belong se- curely to the Satyrinae as currently conceived (ACKERY 1988; BROWER 2000; WAHLBERG et al. 2003). YOSHI- 8 Bonner zoologische Beiträge 53 (2004) MOTO (2003) has formally raised the Melanitini (to in- clude Manataria) to tribal rank within the Satyrinae. Following DEVRIES et al. (1985), the Antirrheina are now widely accepted to as a subtribe of the Morphini (ACKERY 1988; HARVEY 1991; BROWER 2000). But what conclusions should be drawn with respect to Bia? Appreciating the peculiarity of Bia is confounded by MILLER's description of the adult insect, which is inac- curate with respect to the labial palpi and forewing ra- dial venation (the latter error has recently been repeated by YOSHIMOTO 2003), and incomplete most notably with respect to the androconial organs. In the following sections we first correct MILLER’s (1968) account of the palpi and forewing venation. This is followed by obser- vations on its wing patterns and an extensive account of the androconial organs. We then review recently pub- lished work on the early stages, hostplant relationships and molecular systematics, before offering a general discussion. Finally, we summarise a revised provisional classification for Satyrinae and Morphinae (Appendix I). 3. THE LABIAL PALP AND FOREWING RADIAL VENATION Labial palp. MILLER’s (1968: 33) account of the adult morphology of Bia is fundamentally incorrect on two points. First, regarding the labial palp, he states that “the third segment ... is very long, over half the length of the second segment”. Such an arrangement would be highly autapomorphic, but is simply not the case. As first shown by SCHATZ & ROBER (1889: pl. 39), the third segment is much shorter, about one quarter the length of the second, as found in very many Nymphalidae. Forewing radial venation. In contrast, as MILLER cor- rectly appreciated, the forewing radial venation of Bia is highly autapomorphic, but his description (“forewing radial veins arise from a single branch”) and illustration (MILLER 1968: 33, fig. 29) are inaccurate. The forewing radial system comprises two main branches, Rj. and R3+4+5, Which arise in very close proximity at the ante- rior apex of the discal cell (Figs. 6, 7). After about 0.5 mm, Rj divides. R; then fuses with the subcostal vein for about | mm before separating again and finally run- ning free to the costa (such an anastomosis occurs in many butterflies). R runs free to the costa, but for the first 2 mm or so of its length it remains extremely close to R345. After this parallel section, R3.4:5 gently di- verges before separating, at about 5 mm from the apex of the discal cell, into R3 and Ry,s5; about 2 mm or so further on the latter separates into Ry and Rs, with all the separate branches of R eventually running free to the costa. The real peculiarity of this system is the ‘joint’ origin (Fig. 7) of two branches of the radius as R¡+> and R3.4:5, and their extremely close, parallel course that continues as R, and R3+4+5 (Fig. 6). Richard I. VANE-WRIGHT & Michael Boppre: Adult Morphology and the Higher Classification of Bia | mth ae Figs 10-14: Bia actorion (L.) sensu lato. Hindwing androconial organs. 10 anterior (A) and posterior (B) alar organ: 11 posterior hair tuft fully erect; 12 pocket between 2A and 3A exposed with tuft closed (BMNH(E) #693030); 13 tuft partly erect revealir part of scale patch (BMNH(E) #693232); 14 tuft fully erect, patch of modified scales (ms) visible (BMNH(E) 693196). Sc bars: 10-14: 5 mm. 4. WING PATTERN The underside pattern of Bia is very reduced compared with the nymphalid groundplan (NUHOUT 1991: 24). Almost the entire area of both wings is covered by a ripple pattern (Figs. 3-5) (NIJHOUT 1991: 37), relieved only by marginal and submarginal bands on both wings, three or four specialised border ocelli and the parafocal elements on the forewings, and some very reduced ocelli and a few other markings on the hindwings, in- cluding the small but characteristic diagonal white stripe in cell Cuy, (Figs. 3-5). dic The forewing border ocelli, although small, are distinc- tive, occurring very close to the wing margin, with the two or three anterior ocelli (Ry and Rs, and in some in- dividuals, R;) being reduced to white 'pupils' only (Figs 3, 4). The posterior ocellus (in cell M;), although b developed, is somewhat oblate, with the proximal sid little drawn out to form a blunt point. The parafo elements of the forewing are not overwhelmed by ripple pattern (cf. NIJHOUT 1991: 37), bu tinctive line that deviates more or less markedly intervenous stripe in cell M;. On the hindwin o der ocelli are reduced to vague spots, present only 240 Bonner zoologische Beitráge 53 (2004) gan with hairs from ten anterior tuft rows broken off to reveal a patch made of modified scales (ms) (22, 23), surrounding naked zone (nz), tuft hairs (th), and covering scales (cs) of wing area adjacent to vein 3A; 16 bases of a five rows of palisade-forming tuft hairs; 17, 18 bases of a row showing conjoined sockets; 19 fine structure of hair; 20 part of patch with hairs lifted off to re- veal sockets; 21 detail from sockets with hairs removed; 22, 23 details of scales and sockets comprising patch (ms in 15). Scale bars: 15: 1 mm; 16: 200 um; 17: 50 um; 18: 10 um; 19: 2 um; 20: 100 um; 21: 10 um; 22: 100 um; 23: 10 um. three adjacent anterior cells, R;, Rs and M,, plus a di- agonal whitish stripe in Cu, (which is also a specialised border ocellus — see Discussion). Although the three an- terior hindwing ocelli occur in cells that, in terms of se- rial homology, correspond to the forewing cells that al- ways have border ocelli, unlike the forewing, on the hindwing the ocelli are located far from the margin (Figs. 3, 4). 5. ANDROCONIAL SYSTEM Alar androconial organs. According to MILLER (1968: 34), “there is a patch of mealy scales on the upper end of the cell along crossvein rs-ml, and a long hair tuft lies along 2A.” As long ago and more accurately pointed out by MULLER (1877), the males of Bia possess a “tuft of long pale leather-brown hairs near the inner margin of the hind-wings, which can be erected or de- pressed at will, and when at rest, are enclosed in a long pocket, and also by a patch with long black silky hair 3 Richard I. VANE-WRIGHT & Michael BOPPRÉ: Adult Morphology and the Higher Classification of Bia Hübneı 241 Figs 24, 25: Caligo arisbe Húbner, 1822. Hindwing posterior androconial organs (from BARTH 1953: figs 6, 7). Barth has called the structure 'apparatus assisting evaporation of the secretion’. An: analis of hindwing; dotted: naked area: BOR: ring of bristles: BO: bristles. [Note: obviously, the scale bar in 25 is incorrect. | Figs 26-32: Caligo eurilochus (Cramer). Macrophotographs (26-28) and scanning electron micrographs (29-32) of hindwing posterior androconial organs. 26 hair tuft surrounded by a large shiny zone; 27, 28 partly (27) and fully (28) erected hair tuft; 29-31 tuft rows showing conjoined sockets of palisade-forming tuft hairs; scanning electron micrographs: 32 fine structure of hair. Scale bars: 26-28: 5 mm; 29: 100 um; 30, 31: 20 um; 32: 2 um. near the anterior margin of the hind-wings. This lat- ter patch is covered by a bare spot on the under side of the fore-wings, close to the inner margin.” (cf. Figs. 1, 10). Close examination of the hair tuft of the posterior alar organ (Figs. 11-21) reveals a peculiar arrangement of the hairs from which it is formed. The tuft comprises several rows of transversely inserted hairs, the length of the hairs, the distance between the rows, and the number per row all diminishing posteriorly (Figs. 11, 15; th), so that the entire tuft fits into a pocket formed between veins 2A and 3A. The whole organ, in its retracted state, is about 6-7 mm in length. When the hairs are erect, single lines become apparent, forming palisades (Figs. 11, 12, 14). The fine structure of these hairs (Fig. 19) is typical for many Lepidoptera androconia but their bases are peculiar in being conjoined (Figs. 18, 21). Under- neath the hair tuft there is a large patch of modified scales (Figs. 14, 15; ms) that was overlooked by MULLER (1877). The scales are densely packed and partly upstanding, but do not show any peculiar features under SEM (Figs. 22, 23) or have sockets (Fig. 23) sug- gestive of glandular nature. Under a strong electron beam, these scales twist, something that happens to some scales but is relatively unusual. Adjacent to the posterior organ on the side abutting vein 3A is an exten- sive area of the wing with covering scales that are less dense and with scattered hairs (Fig. 15: cs), unlike the main areas of the wing. 242 In several specimens, the upperside forewing cell Cuy; has a mane composed of hairs that are significantly longer and more densely packed than those found on the rest of the wing. This may represent another androconial organ, perhaps characteristic of one or more of the sib- ling species of which Bia may be composed, and is sub- ject to further study. MÜLLER (1877) found all kinds of “hair-tufts and felted patches” on the wings of male butterflies, including various Satyrinae and Morphinae. For Caligo he noted “Hind-wing of the male with a small tuft of hair near the inner margin, opposite to the middle of the abdomen.” However, he apparently did not realize that the ar- rangement of the posterior hair tuft in Bia shares some peculiar features with Caligo (BARTH 1953; cf. figs. 24, 25), Penetes Doubleday, 1849, Catoblepia Stichel, 1902, Opsiphanes (ELTRINGHAM 1926; BARTH 1952), Blepolenis Rober, 1906, and Caligopsis Seydel, 1924, and some other brassolines. The most striking similarity is that the rows of tuft hairs arise from conjoined sock- ets (Figs. 24, 25, 27-32), a configuration currently un- known elsewhere in the butterflies. However, each row in these other genera comprises only a single line of scales (Fig. 29), not a double or triple line as in Bia (Figs. 17, 20, 21). Also, the number of rows of hairs (Figs. 27, 28) forming the posterior alar organ is always less, sometimes as few as 3 rather than 11-15 found in Bia. The major difference is the lack of a scale patch, but the surrounding zone (Figs. 24, 26) 1s comparable, normally much larger and conspicuous as a shiny, na- creous area (= “Reibefláche” of STICHEL 1909). The anterior alar androconial organ of Bia consists in part of a pencil of hairs about 6-7 mm long (Fig. 33) in- serted on the upperside close to the base of the hindwing discal cell (Fig. 10), and aligned approximately with the radial sector. It is evident that this pencil can be erected, as the hair sockets are modified to form an obvious ‘click’ mechanism (Fig. 36) comparable to that ob- served in the forewing alar organ of the morphine Antir- rhea (Vane-Wright 1972b). These hairs do not other- wise exhibit special morphological peculiarities (Fig. 37), but when decumbent (Fig. 33) they virtually cover an extensive patch of modified scales (Fig. 34). This scale patch was not mentioned by MULLER (1877), but MILLER (1968: 34, fig. 29) referred to it (or the organ as a whole) as “a patch of mealy scales on the upper end of the cell.” Probably a dual organ in the terminology of BOPPRE & VANE-WRIGHT (1989: 123), the hairpencil and its patch lie directly opposite a completely naked area on the underside of the forewing (MULLER 1877). A dual anterior alar organ of this type located in the hindwing discal cell is not typical for the Brassolini, but many members of the tribe have androconial organs of various sorts, including hairpencils located at various positions on the wings (cf. STICHEL 1909). Caligo, for Bonner zoologische Beitráge 53 (2004) example, has a conspicuous area of scales on the upper surface of the hindwing discal cell (Figs. 70-74) but, unlike Bia, this patch in Caligo is not associated with a hairpencil. Eryphanis Boisduval, 1870, has a patch and a hairpencil (ELTRINGHAM 1926), but the latter does not rest on the former. Abdominal androconial organs. STICHEL (1909) men- tions for many brassolines, including Caligo, Penetes, Opsiphanes, Catoblepia, and Eryphanis, “Reibewiilste” or “drüsenartige Wülste” (rubbing or glandular bulges) that occur laterally on the male abdomina; no further characterisation is given. Brassolis Fabricius, 1807, Dy- nastor Doubleday, 1849, Dasyophthalma Westwood, 1851, Narope Doubleday, 1849, Opoptera Aurivillus, 1882, and Selenophanes Staudinger, 1887, lack them. These structures were not mentioned by MULLER (1877), but they have been described in considerable de- tail by BARTH (1952, 1953), and also by WASSERTHAL & WASSERTHAL (1977; as “scent pads”). Some of these structures are figured here for Caligo eurilochus (Cramer, 1775) (Figs. 63-69). For the first time we describe lateral abdominal pads in Bia (Figs. 42, 44-46). Unfortunately, the condition of the specimens available to us is not suited for detailed study. However, in contrast to Caligo, the pads of Bia are located on the tergites (Figs. 42-46), not within the pleurae (Fig. 63). Moreover, in Bia the pads are com- prised of three relatively simple scale types (Figs. 47— 60), none of which matches the single highly specialised type (Figs. 64-69) of Caligo. The abdominal pads of Caligo can be protruded (WASSERTHAL & WASSER- THAL 1977). One set specimen of Bia in the collection of the BMNH shows the pads protruded, appearing as warty, shiny structures (Figs. 61-62). Another differ- ence between Caligo and Bia concerns the resting posi- tion: in Caligo, when the butterfly is at rest, the pads are enclosed by the anal area of the hindwings, and thus must come automatically in contact with the posterior alar organs. In Bia, however, the posterior alar organ at rest is enfolded, and contact with abdominal pads would require a special behaviour. While Caligo exhibits dual androconial organs, those of Bia appear to be binate (BOPPRE & VANE-WRIGHT 1989). Although there are many differences in detail, the abdominal pads of Bia are grossly similar those found in Brassolini, and andro- conial organs of this general type are unknown from other taxa. 6. EARLY STAGES AND HOSTPLANT RELATIONSHIPS Until the publication by FREITAS et al. (2002), the life cy- cle of Bia was undescribed. Here we summarise their re- sults with reference to features of the early stages consid- ered likely to be of significance for higher classification. | | Richard I. VANE-WRIGHT & Michael BOPPRÉ: Adult Morphology and the Higher Classification of Bia Hübner 243 PD puras | = Th Wt HARV LL TG) LE SPL) LULL ELIS VA ANETTE hate == a Zi r A . Y FE , 4 a . =! 5 s i ,. : 4 es. , CA Ce, = ns zes 35 Es AA Rad 5 Fee a> r, “2 = - ear 33 a 22; - A es = + 227 23 + as: S Fr “en! = %: eee 2 ey y AA A SS — nr 2 m... m a See wee o gS See SN m . ee —— a pt ee ee gen WEB ne nn a ie Figs 33-41: Bia actorion (L.) sensu lato. Macrophotographs (33, 34) and scanning electron micrographs (35-41) of hindwing an- terior androconial organs. 33 hair tuft in resting position, obscuring 34 patch with modified scales; scanning electron micro- graphs: 35, 36 hair bases exhibiting 'click' mechanism; 37 fine structure of hair; 38-41 patch scale bases and scale fine structure are unexceptional but the scale undersides are not perforated (40, U). Scale bars: 33, 34: 2 mm; 35: 100 um; 36, 37: 5 um: 38: 20 um; 39: 50 um; 40: 20 um; 41: 10 um. Egg. Spherical, with 25-30 longitudinal ribs and as many as 50 transverse ridges (FREITAS et al. 2002: 120, fig. la). The eggs are thus comparable to those of Brassolini, which have 30-60 transverse ridges (e.g. Narope: CASAGRANDE 2002: figs. 1,2), and are unlike those of Satyrinae, which never have as many (FREITAS 1999), First instar larva. The head capsule lacks scoli but has numerous long, branched or plumose setae (FREITAS et al. 2002: figs. Ib,c). A very similar condition can be seen in some Brassolini (e.g. Narope: CASAGRANDE 2002: fig. 4). FREITAS et al. (2002: 119) note that the newly hatched larvae are active, moving around the hostplant unlike “the sluggish behaviour of typical sa- tyrines”. Later instar larvae. Later instars have three pairs of scoli on the head capsule (FREITAS ef al. 2002: 121, figs lij.k), typical of most Brassolini other than Brassolis (cf. DEVRIES 1987: fig. 32 E, 1-8; CASAGRANDE 2002: fig. 3e). According to FREITAS et al. (2002: 121), Sa- tyrinae only have one pair of such scoli; however, while this is generally the case, arguably Elymnias Hübner, 1818, also has three pairs (IGARASHI & FUKUDA 1997: 85-89). The form of the head scoli in Bia is, however, highly autapomorphic, especially the dorsal pair (FREITAS et al. 2002: fig. Ik). The bifid caudal projec- = S S N em un vo oD Bes} = = oO faa} © < o Bl &b = 9 e) N a) = S 2 m Richard I. VANE-WRIGHT & Michael BOPPRÉ: Adult Morphology and the Higher Classification of Bia Hübneı 245 tions (FREITAS et al. 2002: figs. Ib,f,g,h,i) are like those seen in many Brassolini (e.g. Caligo: CASAGRANDE 1979), and are thus grossly similar to all members of the Satyrine clade as conceived by FREITAS & BROWN (2004), including Amathusiini, Calinaginae and Apaturinae. The numerous secondary body setae give a “hairy” appearance, as in many Brassolini (e.g. Caligo: CASAGRANDE & MIELKE 2000a: fig. 3) and Amathusiini (IGARASHI & FUKUDA 1997). Pupa. Squat and sculptured (FREITAS et al. 2002: fig. I l,m), and thus quite similar to e.g. Opsipha- nes (DEVRIES 1987: fig. 32B) and Dasyopthalma (CASAGRANDE & MIELKE 2000b: figs 6-8; 2003: figs. 4-6). Hostplants. Astrocaryum G.Mey, 1818, and Geonoma Willd., 1805 (Arecaceae) (FREITAS et al. 2002). Arec- aceae are recorded as foodplants of species of the bras- soline genera Brassolis, Opsiphanes, Catoblepia and Dasyophthalma, and are also utilised by some species of Morphini, Amathusiini and Satyrinae (ACKERY 1988). Among the Brassolini, Geonoma is recorded as the host of Dasyophthalma species (CASSANGRANDE & MIELKE 2000b, 2003), both Geonoma and Astrocaryum are re- corded as hosts for Opsiphanes (PENZ et al. 2000), and Astrocaryum as a host for Brassolis (ACKERY 1988). 7. MOLECULAR EVIDENCE BROWER (2000) carried out a cladistic analysis of 103 species of Nymphalidae based on sequence data ob- tained from a 378 base-pair region of the wingless gene. In addition to Bia, his sample included species repre- senting 6 genera conventionally included in the Mor- phinae sensu lato (Morpho, Caerois, Antirrhea, Amathusia, Caligo and Opsiphanes), and 13 genera in- cluded in the Satyrinae (Haetera Fabricius, 1807, Melanitis, Lethe Hübner, 1819, Mycalesis Hübner, 1818, Tisiphone Hübner, 1819, Megisto Hübner, 1819, Oressinoma Westwood, 1852, Taygetis Hübner, 1819, Cercyonis Scudder, 1875 , Corades Doubleday, 1848, Lymanopoda Westwood, 1851, Pedaliodes Butler, 1867, and Steroma Westwood, 1851). In his preferred solution (a most parsimonious cladogram produced us- ing the successive approximations weighting option in PAUP 3.1: SWOFFORD 1991), all 20 of these genera, in- cluding Bia, formed a monophyletic group. This was di- vided into two subclades, one including the 13 genera conventionally included in the Satyrinae plus Amathu- sia. The remaining five conventional morphines, plus Bia, formed the other group. Within this latter clade, Bia grouped as sister to Caligo, with Opsiphanes as sister to these two, with these three forming the sister group to (Morpho (Antirrhea + Caerois)). WAHLBERG et al. (2003) presented results from a cladis- tic analysis of 54 Nymphalidae, based on sequence data for one mitochondrial gene (COI, 1450 bp) and two nu- clear gene sequences (EF-la, 1064 bp; and wingless, 412-415 bp). His species sample represented three con- ventional morphine and nine conventional satyrine gen- era, including Manataria but not Bia. Although all of these genera grouped within a single major subclade, this grouping also encompassed both of their exemplar Charaxinae, and the enigmatic Calinaga Moore, 1857 (Calinaginae). This last genus (not available to Brower) grouped as sister to the two Charaxinae, and in some analyses these three together appeared as the sister group to (Caligo + Morpho). The remainder of the clade formed a paraphyletic assemblage of the nine conven- tional Satyrinae including Melanitis and Manataria, to- gether with Stichophthalma C. & R. Felder, 1862 (Amathusiini). Manataria appeared as sister to Melani- tis, and did not group with any of the Satyrini or Elym- niini included in the analysis. This offers support for the inclusion of Manataria within the Melanitini as dealt with by YOSHIMOTO (2003), and not in the lethines (Elymniini) as vaguely speculated by MILLER (1968), or in the Euptychiina (Satyrini) as suggested by FORSTER (1964). 8. DISCUSSION MILLER (1968: 33) made two errors in his account of Bia. First, he stated that the third segment of the labial palpus was abnormally long, exceeding half the length of the second. As revealed even by his own diagram (MILLER 1968: fig. 30), this is simply inaccurate. The gross morphology of the Bia palp is commonplace and unremarkable, being directly comparable to brassolines such as Aponarope Casagrande, 1982 (CASAGRANDE 2002: fig. 103), and many other nymphalids. MILLER’s description of the forewing venation was also wrong: two branches of the radius (R;¡.> and R3.4-s) arise from the cell, not one. MILLER (1968: fig. 29) was mis- led because the anterior of the two branches arising from the cell, R).. and its continuation as the basal part of free R,, lies parallel to R3.¿+s, the two sections ini- tially running very close together (Fig. 7). He was none- theless correct to regard the venation as very peculiar. Figs 42-52: Bia actorion (L.) sensu lato. Scanning electron micrographs of abdominal androconial organs. 42 lateral view of ab- domen from segment 3 to apex, showing position of lateral pads on tergites of segments; 43 channel-like strukture formed by pleurae; 44-46 pads on segment 4-6; 47 short (type 1) and long (type 2) androconial scales clothe all three pads: 48 detail of 47: 49 small marginal scales (type 3); 50, 51 scale bases of type 1; 52 scale base of type 2. Scale bars: 42: | mm: 43: 100 um: 44, 45: 200 um; 46: 100 um; 47: 50 um; 48: 10 um; 49: 100 um: 50: 20 um: 51, 52: 5 um. 246 Bonner zoologische Beitráge 53 (2004) Figs 53-62: Bia actorion (L.) sensu lato. Scanning electron micrographs of abdominal androconial organs. 53-56 details of scale type 1; 57, 58 details of scale type 3; 59, 60 details of scale type 2; macrophotographs: 61, 62 dorsal views of abdomen of unique museum specimen in which the pads are exerted. Scale bars: 53: 10 um; 54: 5 um; 55, 56: 2 um; 57: 20 um; 58: 2 um; 59: 50 um; 60: 2 um; 61-62: | mm. Although the forewing radial vein configuration of Bia is unique among the butterflies, it can be compared in some ways with the Brassolini: Naropina (genera Narope and Aponarope), in which three branches of the radius arise in close proximity from the discal cell, with either a single anastomosis of Sc+R, (CASAGRANDE 1989: fig. 5), or a double anastomosis giving a short section Sc+Rj)42, with Sc+R, and R> eventually running separately to the costa (STICHEL 1904: pl. 1, fig. 5; CASAGRANDE 1989; 1996: fig. 29). The venation of Bia is also comparable to that of certain amathusiines, such as Discophora, in which the forewing radial system similarly arises from the discal cell as two branches that run closely parallel. However, the two branches in Discophora are R, and Rs (not Ry42 and R3+4+5). In this genus R, forms a long anastomosis with Sc before sepa- rating, then forms an anastomosis with R> before they separate and run free to the costa (BASCOMBE et al. 1999: fis. 9.38). But even if MILLER had been right regarding the palp, two such autapomorphies would have told us little about relationships. The odd venation even when correctly de- scribed, simply underscores the generic distinctness of this peculiar nymphalid. This would also be true with respect to the absence of tibial spurs, another unusual feature of Bia observed by MILLER (1968: 33, fig. 31). In this context it may be significant that Narope also Richard I. VANE-WRIGHT & Michael BOPPRÉ: Adult Morphology and the Higher Classification of Bia Húbnes 24 lacks tibial spurs, whereas its close relative Aponarope does not (CASAGRANDE 2002: figs. 30a,b, 103a,b). However, before drawing any detailed conclusions about the relationships of Bia, we first discuss a series of wider questions regarding its higher classification. Does Bia belong to the satyrine clade? In nymphalid butterflies other than Bia, ripple patterns (NIJHOUT 1991) are found in the Nymphalina (e.g. Nymphalis Kluk, 1802, Aglais Dalman, 1816, Polygonia Hübner, 1819), Satyrinae (many species, including Melanitis, Elymnias and Ypthima Húbner, 1818), Morphinae (all Brassolini; Morphini: Antirrheina; and a few Amathusi- ini, including Thauria Moore, 1894, and Discophora), and certain Charaxinae (including Palla Húbner, 1819, and Anaea Hiibner, 1819). Thus the capacity to produce underside ripple patterning appears to be a characteristic (with the exception of the Nymphalina) of the satyrine clade sensu WAHLBERG et al. (2003) as based on mo- lecular evidence, or the satyroid clade sensu FREITAS (1999) other than the supposedly basal Apaturinae (FREITAS & BROWN 2004) as based on early stage char- acters. Other than in the Nymphalina, ripple patterns are unknown elsewhere in the Nymphalidae, including the very small subfamily Calinaginae (considered internal to the satyrine clade by BROWER 2000, WAHLBERG et al. 2003, and FREITAS & BROWN 2004), and the Apaturinae (not included in the satyrine clade by BROWER 2000 or WAHLBERG et al. 2003). Based on many characters including features of the thorax, EHR- LICH (1958) placed Bia in the Nymphalidae: Satyrinae. With the recent addition of molecular and early stage data, there is now little doubt that Bia belongs to the sa- tyrine clade sensu WAHLBERG et al. (2003), and its rip- ple pattern (Figs. 3-5) is further evidence from adult morphology consistent with this conclusion. Why isn't Bia a satyrine? As reviewed above, most au- thors have included Bia within the Satyrinae, including EHRLICH (1958), MILLER (1968), and HARVEY (1991). The emergent view, however, is that Bia belongs to the Morphinae: Brassolini (CLARK 1947, 1948; DEVRIES et al. 1985; BROWER 2000; FREITAS et al. 2002; VANE- WRIGHT 2003; YOSHIMOTO 2003). The question then naturally arises, what are the distinguishing features of the Satyrinae, and does Bia exhibit them or not? Unfortunately, from a morphological perspective, no uniquely diagnostic features for the Satyrinae have been recognised (DEVRIES et al. 1985; HARVEY 1991: ACK- ERY et al. 1999). Traditional but non-unique characters include the closed hindwing discal cell, feeding on monocots, and the fleshy, bifid larval tail (MILLER 1968; ACKERY et al. 1999). Although Bia has all of these features, none is diagnostic for Satyrinae with re- spect to Morphinae. EHRLICH (1958) listed a number of characters for all subfamilies of the Nymphalidae that he recognised. For the Morphinae and Satyrinae the only clear separation he gave was another traditional character, the inflated forewing veins Sc, Cu and 2A never clearly seen in the Morphinae, but present in many Satyrinae. In this respect Bia is a typical satyrine and unlike the morphines. However, the expression, of this character varies widely. For example, it is virtually unexpressed in Satyrinae: Melanitini, while only vein Sc is inflated in Satyrinae: Ragadiini. Moreover, inflated forewing veins occur elsewhere in the Nymphalidae, well outside the satyrine clade (EHRLICH 1958: ACKERY et al. 1999). Why don't the Brassolini belong to the Satyrinae? Even if Bia were most closely related to the Brassolini, we must also consider the possibility that the Brassolini are simply nested, to the exclusion of the Morphini and Amathusiini, within the Satyrinae, as proposed by MILLER (1968). DEVRIES et al. (1985) re-affirmed EHR- LICH's (1958) position by including the Brassolini and Amathusiini within the Morphinae, doing so primarily on the basis of three putative larval characters. How- ever, DeVries later retained the Morphinae and Brasso- linae as separate subfamilies, and commented that the latter were “closely related to the Satyrinae” (DEVRIES 1987: 245). HARVEY (1991), partly due to an error in in- terpreting ACKERY (1988) (see ACKERY et al. 1999), also kept the Brassolinae as a separate subfamily. Given all this uncertainty, and the fact that the larval characters introduced by DEVRIES et al. (1985) remain unverified for many relevant taxa, we must question whether or not it is correct to link the brassolines with the morphines. and whether or not they can legitimately be excluded from membership of the Satyrinae, either alone, or to- gether with the Morphini. Do the Brassolini belong to the Morphinae? In the molecular investigations of both BROWER (2000) and WAHLBERG et al. (2003), their exemplar Brassolini grouped exclusively with their exemplar Morphini. while the two Amathusiini (Amathusia in BROWER, Sti- chophthalma in WAHLBERG et al.) appeared elsewhere. either within a monophyletic Satyrinae (BROWER 2000: fig. 4), or as part of a paraphyletic assemblage made up of Satyrinae, Morphinae, Charaxinae and Calinaginae (WAHLBERG et al. 2003: fig. 4). This is consistent with the conclusion of EHRLICH (1958) that the Satyrinae and Morphinae sensu lato are closely related, as also sug- gested by KUZNETZOV & STEKOLNIKOV (2001). who linked the Morphinae sensw EHRLICH (1958) with the Satyrinae as a monophyletic family. The conclusions of FREITAS & BROWN (2004: fig. 5) also suggest that the Brassolini are more closely related to the Morphini (and the Amathusiini) than they are to the Satyrinae. SCOTT (1985) considered that the larval “fuzzy head” characterised the Morphinae sensu EHRLICH (1958) as a 248 Bonner zoologische Beitráge 53 (2004) 4 far Sr Figs 63-69: Caligo eurilochus (Cramer). Macrophotograph (63) and scanning electron micrographs (64-69) of abdominal andro- conial organs. 63 lateral view of abdominal segments 4-6 showing pad, located in the pleural area between the tergites and ster- nites; scanning electron micrographs: 64-69 scales making up pad; they exhibit unusual scale structure (65), and, in particular, highly specialised sockets (66-68); note tube leading from interior and funnel-like base that fits over the specialised sockets (69). Scale bars: 63: 2 mm; 64: 20 um; 65: 5 um; 66, 67: 20 um; 68: 10 um; 69: 20 um. S384 A u N fs aot Ee FF “8 Ge 5 = y Ua Figs 70-74: Caligo eurilochus brasiliensis (Cramer). Macrophotographs (70, 71) and scanning electron micrographs (72-74) of hindwing anterior androconial organs. 70 patch immediately anterior to vein Rs (note also very small precostal cell): 71 detail of patch; scanning electron micrographs: 72 scales comprising patch (U: underside); 73 detail of scale surface; 74 scale sockets. Scale bars: 70-71:5 mm; 72: 50 um; 73: 2 um; 74: 20 um. Richard I. VANE-WRIGHT & Michael BOPPRÉ: Adult Morphology and the Higher Classification of Bia Hübner 249 monophyletic group. If so, this raises doubts about in- clusion of the Amathusiini within the Satyrinae. Cer- tainly the larvae of various Amathusiini do have “fuzzy” heads (e.g. IGARASHI & FUKUDA 1997: pls. 126-133), comparable to Brassolini and Morphini. However, it is evident that the head capsule of Melanitis, for example, is also quite “fuzzy”, and more work is needed on this character. Another character linking the Amathusiini to the Brassolini is the presumed repugnatorial neck gland of the larvae (ELIOT, in CORBET & PENDLEBURY 1992: 137). Our conclusion is that, despite the weakness of the present evidence, we should maintain the Morphinae sensu EHRLICH (1958), to include Morphini, Amathusi- ini and Brassolini. Is Bia a brassoline? In BROWER's (2000) analysis, and in some of the analy- ses of FREITAS & BROWN (2004), Bia groups with the Brassolini to form a monophyletic group, either with just the Morphini (BROWER 2000; FREITAS & BROWN 2004: fig.1) or with the Morphini + Amathusiinae in ad- dition (FREITAS & BROWN 2004: fig. 3). So the Brassol- ini do not appear to belong to the Satyrinae sensu stricto. Given the evidence from early stages reported by FREITAS et al. (2002) that seem so suggestive that Bia is a brassoline (summarised in section 7 above), it 1s perhaps surprising that FREITAS & BROWN (2004: fig. 5) placed Bia as a monobasic subfamily separate from both Morphinae and Brassolinae. Apparently they did so be- cause Bia behaved ambiguously in their analyses: “it appeared in three different positions in the trees” (FREITAS & BROWN 2004: 372). Can any strong support or challenge to the hypothesis that Bia is a brassoline be drawn from our re-examination of adult morphology? ACKERY et al. (1999) stated that the “Brassolini ... cur- rently lack convincing autapomorphies”. Although MILLER‘s (1968) arguments for including the Brassolini within the Satyrinae were unconvincing, he did identify one relatively distinctive character for the group appar- ently overlooked by ACKERY et al. — the basal separa- tion of the hindwing veins Sc and R, to produce a dis- tinct precostal cell (STICHEL 1909). MILLER correctly pointed out that this feature recurs in a few groups in- cluded in the Satyrinae (e.g. Elymnias: SCHATZ & ROBER 1889: pl. 39), and essentially the same character is found in some other butterflies, including many Papil- ionidae (SMITH & VANE-WRIGHT 2001), various Char- axinae (SCHATZ & ROBER 1888: pls. 28, 29), Parthenos Hiibner, 1819 (Limenitidinae: SCHATZ & ROBER 1887: pl. 25), various Danaini (ACKERY € VANE-WRIGHT 1984), and even Morpho itself (SCHATZ & ROBER 1885: pl. 1, fig. 1). Thus, although characteristic of all Brasso- lini, the precostal cell is not uniquely diagnostic for the group. Within the Brassolini, as demonstrated by STICHEI (1904, 1909), the precostal cell varies significantly in size and form. In some genera (e.g. Opoptera) it is very large (STICHEL 1904: pl. 2, fig. 1), and most brassolines approach this condition. In Eryphanis, Caligopsis and Caligo, however, the precostal cell is much smaller, with a narrow, ovoid 'lumen' (STICHEL 1904: pl. 2, figs. 4, 5), unlike the widely open 'parallelogram' seen in other genera. Bia has a slight basal separation of these veins, as correctly observed over 150 years ago by WESTWOOD (1850), but this can only be appreciated readily by examination of a cleared wing preparation or use of SEM. The form this takes in Bia (Figs. 8, 9) is like a miniaturised Eryphanis or Caligo (Fig. 70). How- ever, given the homoplasious distribution of this charac- ter as noted above, to include Bia within the Brassolini on this basis would be unconvincing. The configuration of the forewing ocelli of Bia closely approximates that seen in several brassoline genera, no- tably Opoptera, Catoblepia, and many species of Opsi- phanes. This is also true for the curvilinear path of the parafocal elements that occurs in some of the species belonging to these genera, including the deviation in cell M3. Overall, this gives a reminiscent 'Gestalt' to both the upperside and underside pattern of the forewing apex of Bia and these three genera. On the hindwing underside the position of the ocellar marking in cell R, also corresponds closely to that occupied by the large and fully-developed border ocellus in underside cell R; of the same genera. The suggestion of a border ocellus in hindwing cell Rs is unusual in most brassolines, while an ocellus in M, is only seen in a few genera, notably Brassolis and Dasyophthalma. However, in many spe- cies of Narope small border ocelli similar to those of Bia occur in all hindwing underside cells R¡-Cu;, (e.g. CASAGRANDE 2002: figs. 29, 95). Most Brassolini have an extremely well developed border ocellus in hindwing underside cell Cu,,, reaching its maximum development both in size and basal displacement to give the huge eyespot characteristic of the owl butterflies (Caligo). Of this there is no obvious trace in Bia, unless we interpret the curious diagonal white stripe that occurs in cell Cu), adjacent to the tail-like extension formed around Cu), (also unique to this genus: the tail of Opoptera is formed around M;) as a modified remnant of the ocellar pupil. This appears to be confirmed by the very similar underside white stripe that occurs in cell Cu, of some species of Narope, such as N. cyllastros Doubleday, 1849, and N. cyllene Felder, 1859 (CASAGRANDE 2002: figs. 28, 29, 34, 35; cf. fig. 50). Male genitalia are widely used in insect systematics, but these highly plastic structures are often difficult to in- terpret for higher taxonomy (SMITH € VANE-WRIGH1 2001). Most members of the satyrine clade have rela- tively simple male genitalia, and this is true for Bia 250 Bonner zoologische Beitráge 53 (2004) (HAYWARD 1958, 1964). Indeed, the genitalia of Bia are quite similar to Narope, except that the ganthos is di- rected ventrally (as in Brassolina), unlike the upswept structure found in Naropina (CASAGRANDE 1996: figs. 10-12, 31, 32). As with the precostal cell, the wing patterns of Bia and perhaps even the male genitalia are suggestive of a rela- tionship with the Brassolini, but are not wholly convinc- ing. In contrast, several features of the androconial or- gans provide what we consider to be strong evidence for such a relationship. Notably, hair-tufts arranged in palisades with conjoined sockets and abdominal pads occur in many Brassolini, and in Bía, but not in the Sa- tyrinae. 9. CONCLUSIONS Evidence from all life stages, including several adult characters described here, and DNA sequence data, supports the view that Bia is a member of the morphine tribe Brassolini. Even though Bia is very small for a brassoline (forewing length 25-32 mm) and highly aut- apomorphic, it may ultimately prove to be internal to the tribe as a whole, and not sister to the rest of the group as suggested by one of the analyses made by FREITAS & BROWN (2004: fig. 2). Until recently there has been no accepted subtribal clas- sification for the Brassolini. However, CASAGRANDE (1996, 2002) has separated the Naropina Stichel (to in- clude only Narope and Aponarope) from all of the re- maining genera, which she included in the Brassolina. In addition to its marked autapomorphic features (e.g. forewing radial venation, inflated forewing veins, min- ute hindwing precostal cell, basally fused dorsal horns of larval head), Bia shares putative synapomorphies with both the Naropina (e.g. loss of tibial spurs; unique form of hindwing underside border ocellus in cell Cu;a; possibly the plumose hairs on larval head) and the Bras- solina (e.g. tufted alar organs composed of palisade rows and abdominal pads, as found in Caligo and sev- eral other genera). In the circumstances, we propose, pending more extensive analysis, to place Bia in the Bi- ina Herrich-Schaffer, 1864, as a third subtribe of the Brassolini Boisduval, 1836 (see Appendix I). However, it seems quite possible that the Biina will ultimately be subsumed within the Brassolina, or subsume the Naropina. BOPPRE (1984) commented that “androconial organs are . analogous structures, convergently evolved many times ... [and] of limited taxonomic value, although they certainly provide good characters in some groups”. Despite this rather cautious view, our experience over the intervening 20 years suggests that detailed investiga- tions of androconial organ morphology (e.g. BOPPRE & VANE-WRIGHT 1989), even though loss and independent gain of these organs are indeed frequent evolutionary phenomena, can provide extremely valuable insights into systematic relationships (e.g. VANE-WRIGHT et al. 2002; cf. HALL € HARVEY 2002). This can also be true of androconial chemistry, as in the Danaini (VANE- WRIGHT & BOPPRE 1993; SCHULZ et al. 1993), although SCHULZ et al. (2004) found relatively little evidence for phylogenetic relationships from their analyses of Ithomiini pheromones. In the case of the two Neotropi- cal tribes of Morphinae, abdominal coremata are diag- nostic for the Morphini, while palisade alar organs and abdominal pads appear to be autapomorphic for Brassol- ini (including Bia), even though they are not expressed by all members of the group. We conclude that a combined morphological and chemical investigation into the scent organs of Bia and other Brassolini would be a most interesting and poten- tially instructive challenge. Such a study would be in the best tradition of Clas NAUMANN, our dear departed friend, inspiration and mentor, to whose memory this paper is most respectfully dedicated. To him, gaining knowledge and understanding was always more impor- tant than merely accumulating information. Acknowledgements. We are most grateful for the skilful and painstaking assistance of Ottmar Fischer (FZI) with SEM and macro-photography of androconia and for his comments on the manuscript. We thank Julie Harvey, Kim Goodger, Phil Ackery, Tony Hoare (BMNH) and Gerardo Lamas (Lima) for help with various matters. Special thanks are also due to Shayleen James (BMNH), who prepared Figs. 12-14 with great care. Other original figures were provided by: 1-4 BMNH photo studio; 5 Tony Hoare; 6-9 RIVW. Zusammenfassung. Die südamerikanische Nymphaliden- Gattung Bia Hübner, 1919, wurde für mehr als 150 Jahre von den meisten Lepidopterologen als Mitglied der Satyr- inae betrachtet. Neuere Berichte zu Präimaginalstadien sowie DNA-Analysen haben jedoch gemeinsame Merk- male mit den Morphinae: Brassolini aufgedeckt. Unter- suchungen der Flügelmuster und der androconialen Organe von Bia, hier erstmals im Detail vorgestellt, zeigen ungewöhnliche Merkmale, die sonst nur von Brassolinen bekannt sind. Insbesondere das büschelförmige posteriore androconiale Organ der Hinterflügel, das Palisaden bildet, stellt eine Synapomorphie für Bia und verschiedene andere Gattungen der Brassolini, inclusive Caligo, dar. Die Gat- tung Bia wird daher formal von den Satyrinae zu den Mor- phinae: Brassolini übertragen, als einzigem Taxon des Sub- tribus Biina Herrich-Scháffer, 1864, stat. nov., zusammen mit Brassolina Boisduval, 1836, und Naropina Stichel, 1925. REFERENCES D'ABRERA, B. (1988): Butterflies of the Neotropical Re- gion Part V Nymphalidae (Conc.) & Satyridae: 679— 877. Hill House, Melbourne. Richard I. VANE-WRIGHT & Michael BoPPRÉ: Adult Morphology and the Higher Classification of Bia Húbnes 25] D'ABRERA, B. (2001): The Concise Atlas of Butterflies of the World. Hill House, Melbourne. ACKERY, P. R. (1984): Systematic and faunistic studies on butterflies. Symposia of the Royal Entomological So- ciety of London 11: 9-21. ACKERY, P. R. (1988): Hostplants and classification: a re- view of nymphalid butterflies. Biological Journal of the Linnean Society 33: 95-203, ACKERY, P. R. & VANE-WRIGHT, R. I. (1984): Milkweed Butterflies. Cornell University Press, New York. ACKERY, P. R., DE JONG, R. & VANE-WRIGHT, R. I. (1999): The butterflies: Hedyloidea, Hesperioidea and Papil- ionoidea. Pp. 263-300 in: KRISTENSEN, N. P. (ed.) Handbook of Zoology 4 (35): 263-300. de Gruyter, Berlin. BARTLETT, E. (1876) in: DRUCE, H. (1876): List of the but- terflies of Peru, with descriptions of new species. Pro- ceedings of the Zoological Society of London 1876: 205-250, 2 pls. BARTH, R. (1952): Die Hautdrúsen des Mánnchens von Opsiphanes isagoras Fruhst. (Lepidoptera, Brassol- idae). Zoologische Jahrbiicher Anatomie 72: 216-230. BARTH, R. (1953): Das abdominale Duftorgan des Mánn- chens von Caligo arisbe Hbn. (Lepidoptera, Brassol- idae). Memorias do Instituto Oswaldo Cruz 51: 220- 226. BASCOMBE, M. J., JOHNSTON, G. & BASCOMBE, F. S. (1999): The Butterflies of Hong Kong. Academic Press, London. Boppre, M. (1984): Chemically mediated interactions be- tween butterflies. Symposia of the Royal Entomologi- cal Society of London 11: 259-275. BOPPRE, M. € VANE-WRIGHT, R. I. (1989): Androconial systems in Danainae (Lepidoptera): functional mor- phology of Amauris, Tirumala, Danaus and Euploea. Zoological Journal of the Linnean Society 97: 101- 133. Brower, A.V.Z. (2000): Phylogenetic relationships among the Nymphalidae (Lepidoptera), inferred from partial sequences of the wingless gene. Proceedings of the Royal Society of London (B) 267: 1201-1211. CASAGRANDE, M. M. (1979): Sobre Caligo beltrao (Il- liger). I: Taxonomia, biologia, morfologia das fases imaturas e distribuigdes espacial e temporal (Lepido- ptera, Satyridae, Brassolinae). Revista Brasileira de Biologia 39: 173-193. CASAGRANDE, M. M. (1989): Espécie nova de Narope do sul do Brasil (Lepidoptera, Nymphalidae, Brassolinae). Revista Brasileira de Zoologia 6(1): 125-129. CASAGRANDE, M. M. (1996): Notas sistemáticas sobre Brassolinae. 1. Tribos (Lepidoptera, Nymphalidae). Revista Brasileira de Zoologia 12(3): 671-699. CASAGRANDE, M. M. (2002): Naropini Stichel, taxonomia e imaturos (Lepidoptera, Nymphalidae, Brassolinae). Revista Brasileira de Zoologia 19(2): 467-569. CASAGRANDE, M. M. & MIELKE, O. H. H. (2000a): Larva de quintero estádio e pupa de Caligo martia (Godart) (Lepidoptera, Nymphalidae, Brassolinae). Revista Brasileira de Zoologia 17(1): 75-79. CASAGRANDE, M. M. & MIELKE, O. H. H. (2000b): Larva de quintero estádio e pupa de Dasyophthalma rusina rusina (Godart) (Lepidoptera, Nymphalidae, Brassol- inae). Revista Brasileira de Zoologia 17(2): 401-404. CASAGRANDE, M. M. & MIELKE, O. H. H. (2003): Larva de quarto e quinto estadios e pupa de Dasvophthalma creusa creusa (Hibner) (Lepidoptera, Nymphalidae, Brassolinae). Revista Brasileira de Zoologia 20(1) 157-160. CLARK, A. H. (1947): The interrelationships of the several groups within the butterfly superfamily Nymphal- oidea. Proceedings of the Entomolgical Society of Washington 49: 148-149. CLARK, A. H. (1948): Classification of the binamtiies with the allocation of the genera occurring in North Amer- ica north of Mexico. Proceedings of the Biological So- ciety of Washington 61: 77-81. [Note: this paper is of- ten incorrectly cited as being published in the non- existent journal “Proceedings of the Biological Society of London”, or in the Proceedings of the Entomologi- cal Society of Washington; the pagination may also be given incorrectly as “77-84” due to an error on the first printed page. | CORBET, A. S. & PENDLEBURY, H. M. (1992): The Butter- flies of the Malay Peninsula. Fourth Edition, revised by ELIOT, J.N. Malayan Nature Society, Kuala Lum- pur. COWAN, C. F. (1970): Annotationes Rhopalocerologicae 1970. Privately published by the late C.F. Cowan, Clunbury Press, Berkhamsted, UK. DEVRIES, P. J. (1987): The Butterflies of Costa Rica and Their Natural History. Papilionidae, Pieridae, Nym- phalidae. Princeton University Press, New Jersey. DEVRIES, P. J., KITCHING, I. J. & VANE-WRIGHT, R. I. (1985): The systematic position of Antirrhea and Caerois, with comments on the classification of the Nymphalidae (Lepidoptera). Systematic Entomology 10: 11-32. DIETRICH, K. (1862): Zur Systematik der Schmetterlinge. Stettiner Entomologische Zeitung, Stettin 23: 466- 479. DRUCE, H. (1876): List of the butterflies of Peru, with de- scriptions of new species. Proceedings of the Zoologi- cal Society of London 1876: 205-250, 2 pls. EHRLICH, P. R. (1958): The comparative morphology, phy- logeny and higher classification of the butterflies (Lepidoptera: Papilionoidea). University of Kansas Science Bulletin 39: 305-370. ELTRINGHAM, H. (1926): On a new organ in the abdomen of Eryphanis polyxena, Meerb. (Lepidoptera). Trans- actions of the Entomological Society of London 74: 367-369, 1 pl. FELDER, C. (1861): Ein neues Lepidopteron aus der Familie der Nymphaliden und seine Stellung im natúrlichen System, begründet aus der Synopse der übrigen Gat- tungen. Novorum Actorum Academiae Caesareae Leopoldino-Carolinae germanicae Naturae Curio- sorum 28(3): 1-50, 1 pl. FORSTER, W. (1964): Beiträge zur Kenntnis der Insekten- fauna Boliviens XIX: Lepidoptera III. Satyridae. Ver- öffentlichungen der Zoologischen Staatssammlung Miinchen 8: 51-188. FREITAS, A. V. L. (1999): Nymphalidae (Lepidoptera). filogenia com base em caracteres de imaturos, com experimentos de troca de plantas hospedeiras. PhD Thesis, Universidade Estadual de Campinas, Campi- nas, Sáo Paulo. [Not seen.] FREITAS, A. V. L. & BROWN, K. S. (2004): Phylogeny of the Nymphalidae (Lepidoptera). Systematic Biology 53: 363-383. bo Nn 159) FREITAS, A. V. L., MURRAY, D. & BROWN, K. S. (2002): Immatures, natural history and the systematic position of Bia actorion (Nymphalidae). Journal of the Lepi- dopterists' Society 56: 117-122. GAEDE, M. (1931): Satyridae. Lepidopterorum Catalogus 20(43,46,48): 1-759. W. Junk, Berlin. GODART, J. B. (1824): In LATREILLE, P. A. & GODART, J. B., Histoire Naturelle. Entomologie, ou Histoire Naturelle des Crustacés, des Arachnides et des Insec- tes. Encyclopédie Methodique 9(2): 329-828. HALL, A. (1939): Catalogue of the Lepidoptera Rhopalo- cera (butterflies) of British Guiana. The Agricultural Journal of British Guiana 10: 25—252. HALL, J. P. W & HARVEY, D. J. (2002): A survey of an- droconial organs in the Riodinidae (Lepidoptera). Zoological Journal of the Linnean Society 136: 171— 197. HARVEY, D. J. (1991): Higher classification of the Nym- phalidae. Pp. 255-273 in: NIJHOUT, H.F. The Devel- opment and Evolution of Butterfly Wing Patterns. Smithsonian Institution Press, Washington D.C. HAYWARD, K. J. (1958): Dibujos de los genitales mascu- linos de algunos satiridos neotropicales (Lep. Rhop. Satyridae). Acta Zoologica Lilloana 16: 511—517. HAYWARD, K. J. (1964 (1963”)): Dibujos de los genitales masculinos de algunos satiridos neotropicales. III (Lep. Rhop. Satyridae). Acta Zoologica Lilloana 19: 61-81. HEMMING, F. (1964): Annotationes Lepidopterologicae (3): 75-112. Published privately by the author. HERRICH-SCHAFFER, [G. A. W.] (1864-1871): Prodromus Systematis Lepidopterorum. Versuch einer systema- tischen Anordnung der Schmetterlinge. Correspon- denz-Blatt des zoologisch-mineralogischen Vereines zu Regensburg 18: 89-112, 123-136, 148-152, 173- 174; 21: 100-106, 124-128, 138-144, 161-172; 22: 119-138, 172-176; 23: 56-64, 67— 77, 130-141, 163— 172, 184-204; 24: 154-160; 25: 103-104. HUBNER, J. (1816-1826): Verzeichnis bekannter Schmet- terlinge [sic]. Jacob Hübner, Augsburg. IGARASHI, S. & FUKUDA, H. (1997): The Life Histories of Asian Butterflies, volume 1. Tokai University Press, Tokyo. KIRBY, W. F. (1871): A Synonymic Catalogue of Diurnal Lepidoptera. John Van Voors, London. KIRCHBERG, E. (1942): Genitalmorphologie und natürliche Verwandtschaft der Amathusiinae (Lep. Nymphal.) und ihre Beziehungen zur geographischen Verbreitung der Subfamilie. Mitteilungen der Münchner Ento- mologischen Gesellschaft 32: 44-87. KUZNETZOV, V .I. 8 STEKOLNIKOV, A. A. (2001): New ap- proaches to the system of Lepidoptera of world fauna (on the base of the functional morphology of abdo- men). Russian Academy of Sciences (Proceedings of the Zoological Institute 282: 1-462), St Petersburg. LAMAS, G., NIELSEN, E. S., ROBBINS, R. K., HAUSER, C. L., DE JONG R. & [VANE-WRIGHT, R. 1.] (2000): Develop- ing and sharing data globally: the 'Global Butterfly In- formation System'—GIoBIS. P. 196 in: GAZZONI, D. L. (ed.) Abstracts 21" International Congress of Ento- mology 1. EMBRAPA, Londrina, Brazil. LINNAEUS, C. (1763a): Centuria Insectorum Rariorum. Up- sala. LINNAEUS, C. (1763b): CXXI. Centuria Insectorum. Amoenitates Academicae 6: 384-415. Bonner zoologische Beitráge 53 (2004) MASTERS, J. H. (1970): Bionomic notes on Haeterini and Biini in Venezuela (Satyridae). Journal of the Lepi- dopterists' Society 24: 15-18. MIELKE, O. H. H. & CASAGRANDE, M. M. (1998): Butter- flies of the Ilha de Maracá. Pp. 355-359, 467-478 in: MILLIKEN, W. & RATTER, J. A. (eds) Maracá. The Bio- diversity and Environment of an Amazonian Rainfor- est. Wiley & Sons, Chichester, UK. MILLER, L. D. (1968): The higher classification, phylogeny and zoogeography of the Satyridae (Lepidoptera). Memoirs of the American Entomological Society (24): 174 pp. MULLER, F. (1877): Über Haarpinsel, Filzflecke und ähn- liche Gebilde auf den Flúgeln mánnlicher Schmetterlinge. Jenaische Zeitschrift für Naturwissenschaft 5: 99-114. (English translation. Pp. 604-615 in: LONGSTAFF, G. B. (1912): Butterfly Hunting in Many Lands. Long- mans, Green & Co., London.) NIJHOUT, H. F. (1991): The Development and Evolution of Butterfly Wing Patterns. Smithsonian Institution Press, Washington D.C. PARSONS, M. (1998): Butterflies of Papua New Guinea. Their Systematics and Biology. Academic Press, San Diego and London. [Note: although this book indicates “1999” as its publication date, there is no doubt that it was published and available in 1998; J. Tennent, pers. comm. | PENZ, C. M., AIELLO, A. & SRYGLEY, R. B. (2000): Early stages of Caligo illioneus and C. idomeneus (Nym- phalidae, Brassolinae) from Panama, with remarks on larval food plants for the subfamily. Journal of the Lepidopterists' Society 53(4): 142-152. RACHELI, T. & RACHELI, L. (2001): An annotated list of Ecuadorian butterflies (Lepidoptera: Papilionidae, Pieridae, Nymphalidae). Fragmenta Entomologica, Roma 33(2): 213-380. REUTER, E. (1896): Uber die Palpen der Rhopaloceren. Acta Societatis Scientiarum Fennicae 22(1): xvi + 578 pp. + [vi], 6 pls. REUTER, E. (1898): On a new classification of the Rhopa- locera. Entomologists' Record and Journal of Variation 10(2): 25-26, 75-77, 95-98, 1 pl. SCHATZ, E. & ROBER, J. (1885-1892): Die Familien und Gattungen der Tagfalter. In: STAUDINGER, O. & SCHATZ, E. (eds) Exotische Schmetterlinge 2: 284 pp + 50 pls. G. Löwensohn, Bayern. SCHULZ, S., BOPPRÉ, M. & VANE-WRIGHT, R. I. (1993): Specific mixtures of secretions from male scent organs of Kenyan milkweed butterflies (Danainae). Philoso- phical Transactions of the Royal Society (B) 342: 161-181. SCHULZ, S., BECCALONI, G., BROWN, K. S., BOPPRÉ, M., FREITAS, A. V. L., OCKENFELS, P. & TRIGO, J. R. (2004): Semiochemicals derived from pyrrolizidine al- kaloids in male ithomiine butterflies (Lepidoptera: Nymphalidae: Ithomiinae). Biochemical Systematics and Ecology 32: 699-713. SCOTT, J. A. (1985): The phylogeny of the butterflies (Papilionoidea and Hesperioidea). Journal of Research on the Lepidoptera 23(4): 241-281. SMITH, C. R. & VANE-WRIGHT, R. I. (2001): A review of the Afrotropical species of the genus Graphium (Lepi- doptera: Rhopalocera: Papilionidae). Bulletin of the Natural History Museum London (Entomology) 70: 503-719. Richard I. VANE-WRIGHT & Michael BOPPRÉ: Adult Morphology and the Higher Classification of Bia Hübner 253 STAUDINGER, O. (1888): Exotische Tagfalter [in two vol- umes]. G. Lówensohn, Bayern. STICHEL, H. (1904): Lepidoptera, Rhopalocera, fam. Nym- phalidae, subfam. Brassolinae. In: WYTSMAN, P. (ed.) Genera Insectorum 20: 48 pp. Brussels. STICHEL, H. (1909): Lepidoptera. Brassolidae. In: SEITZ, A. (ed.) Das Tierreich 25: xiv + 244 pp. R. Friedlander und Sohn, Berlin. SWOFFORD, D. L. (1991): PAUP version 3.1, program and documentation. Illinois Natural History Survey, Champaign, IL. UEMURA, Y. (1987): Description of a new species of the genus Bletogona Felder (C.) & Felder (R.) (Lepido- ptera: Satyridae). Memoirs of the Tsukada Collection, Japan (5): 18-22, 1 pl. VANE-WRIGHT, R. I. (1972a): Pre-courtship activity and a new scent organ in butterflies. Nature 239: 338-340. VANE-WRIGHT, R. I. [anonymously] (1972b). Scent organs of male butterflies. Report on the British Museum (Natural History) 1969-1971: 31-35, 4 pls. VANE-WRIGHT, R. I. (2003): Evidence and identity in but- terfly systematics. Pp. 477-513 in: Bocas, C. L., WATT, W. B. & EHRLICH, P .R. (eds) Butterflies: Ecol- ogy and Evolution Taking Flight. University of Chi- cago Press, Chicago. VANE-WRIGHT, R. I. & BOPPRE, M. (1993): Visual and chemical signalling in butterflies: functional and phy- logenetic perspectives. Philosophical Transactions of the Royal Society (B) 340: 197-205, 2 pls. VANE-WRIGHT, R. I., BOPPRE, M. ACKERY, P. R. (2002): Miriamica, a new genus of milkweed butterflies with unique androconial organs (Lepidoptera: Nymphal- idae). Zoologischer Anzeiger 241: 255-267, WAHLBERG, N., WEINGARTNER, E. & NYLIN, S. (2003): Towards a better understanding of the higher systemat- ics of Nymphalidae (Lepdidoptera: Papilionoidea) Molecular Phylogenetics and Evolution 28: 473-484 WALLACE, A. R. (1854): On the habits of the butterflies of the Amazon Valley. Transactions of the Entomological Society of London, New Series 2: 253-264, WASSERTHAL, T. H. & WASSERTHAL, W. (1977): Ultra- structure of a scent scale organ with pressure disgharge in male Caligo eurilochus brasiliensis (Fldr.) (Lepi- doptera: Brassolidae). Cell and Tissue Research 177: 87-103. WESTWOOD, J. O. (1850-1852): Pp. 251-534 in: DOUBLI DAY, E. & WESTWOOD, J. O. The genera of diurnal Lepidoptera 2. Longman, Brown, Green and Long- mans, London. WEYMER, G. (1910-1911): 4. Family Satyridae. Pp. 173 283, 19 pls. in: SEITZ, A. (ed.) Die Grosschmetterlinge der Erde 5. A. Kernen, Stuttgart. WEYMER, G. & MAASSEN, J. P. (1890): Lepidopteren ge- sammelt auf einer Reise durch Colombia, Ecuador, Perú, Brasilien, Argentinien und Bolivien in den Jahren 1868-1877 von Alphons Stiibel. Asher & Co., Berlin. YOSHIMOTO, H. (2003): Notes on the Biini and Melanitini (Lepidoptera, Nymphalidae). Butterflies, Japan (37): 49-55. Authors' addresses: Dr. R. I. VANE-WRIGHT (corre- sponding author), The Natural History Museum, De- partment of Entomology, Cromwell Road, London SW7 SBD, UK; e-mail: dickvanewright@btinternet.com; Prof. Dr. Michael BOPPRÉ, Forstzoologisches Institut, Albert-Ludwigs-Universitát, D-79085 Freiburg, Ger- many; e-mail: boppre@fzi.uni-freiburg.de 254 APPENDIX I Classification the satyrine clade sensu EHRLICH (1958) and KUZNETZOV & STEKOLNIKOV (2001), within the Nymphalidae Rafinesque, 1815, following break-up of the Biinae sensu MILLER (1968). Despite various changes, the current system still owes much to Miller (see HARVEY 1991). However, Miller overlooked some potentially important genera that clearly belong here, such as Penthema Doubleday, 1848, and Xanthotaenia Westwood, 1858, and these need to be located (KIRCH- BERG, 1942, firmly included Xanthotaenia im the Amathusiini). The Melanitini currently include Melani- tis, Cyllogenes, Gnophodes, Parantirrhoea, Bletogona and Manataria (UEMURA 1987; WAHLBERG et al. 2003; YOSHIMOTO 2003). In future it seems possible that the Amathusiini could be relocated within the Satyrinae, while the grouping as a whole will probably be ex- panded to subsume the Charaxinae Guenée, 1865, and the Calinaginae Moore, 1895 (WAHLBERG et al. 2003). The suggestion of FREITAS & BROWN (2004) that the Apaturinae Boisduval, 1840, also belong here is contra- dicted by current molecular evidence (BROWER 2000; WAHLBERG et al. 2003). Type genera are given in square brackets (for further de- tails regarding these generic names see http://www.nhm. ac.uk/entomology/butmoth/index.html). The family group names, authorities, dates and type genera are in accord- ance with the as yet unpublished 'GloBIS' system for the nomenclature and classification of the butterflies (LA- MAS et al. 2000, in prep.). Note that there is currently no widely accepted sub-tribal system for the Amathusiini. The work of KIRCHBERG (1942) will be invaluable in trying to formulate any effective subdivision. PARSONS (1998) has suggested that Morphopsis Oberthiir, 1880, Taenaris Hübner, 1819 (including Morphotenaris Fruhstorfer, 1893), Hyantis Hewitson, 1862, and Faunis Hübner, 1819, may form a subgroup (for which the old- est available family-group name would be Hyantına Róber, 1905 [Hvantis Hewitson, 1862]). The Disco- phorina Stichel, 1902, are recognised by BASCOMBE et al. (1999). Note that YOSHIMOTO (2003), following MILLER (1968), incorrectly attributed the following family-group names to Miller: Melanitini, Mycalesina, Ypthimina and Coe- nonymphina. By following Miller, Yoshimoto also mis- attributed Lethina to Clark, 1948; Melanargiina to Verity, 1920; gave the original date for Satyrinae Boisduval incor- rectly as 1836; and misspelled Antirrheini as “Antirrhint”. (Antirrhini Miller, 1968, is an objective synonym and homonym of Antirrhaeidi Reuter, 1896, the latter based on Westwood's invalid emendation “Antirrhaea”—see COWAN 1970—and which is properly corrected to Antir- Bonner zoologische Beitráge 53 (2004) rheini Reuter, 1896, or Antirrheina Reuter, 1896, depend- ing on adopted rank.) MORPHINAE Newman, 1834 [Morpho Fabricius, 1807] MORPHINI Newman, 1834 [ Morpho Fabricius, 1807] ANTIRRHEINA Reuter, 1896 [Antirrhea Hiib- ner, 1822] MORPHINA Newman, 1834 [Morpho Fabricius, 1807] BRASSOLINI Boisduval, 1836 [Brassolis Fabricius, 1807] BIINA Herrich-Scháffer, 1819] stat. nov. NAROPINA Stichel, 1925 [Narope Doubleday, 1849] BRASSOLINA Boisduval, 1836 [Brassolis Fab- ricius, 1807] AMATHUSIINI Moore, 1894 [Amathusia Fabricius, 1807] SATYRINAE Boisduval, 1833 [Satyrus Latreille, 1810] HAETERINI Herrich-Scháffer, 1864 [Haetera Fab- ricius, 1807] MELANITINI Reuter, 1896 [ Melanitis Fabricius, 1807] ELYMNIINI Herrich-Schäffer, 1864 [Elymnias Húbner, 1818] LETHINA Reuter, 1896 [Lethe Hiibner, 1819] ZETHERINA Reuter, 1896 [Zethera C. Felder, 1861] ELYMNIINA Herrich-Scháffer, 1864 [Elymnias Hübner, 1818] MYCALESINA Reuter, 1896 [Mycalesis Húbner, 1818] ERITINI Miller, 1968 [Erites Westwood, 1851] RAGADINI Herrich-Schäffer, 1864 [Ragadia Westwood, 1851] SATYRINI Boisduval, 1833 [Satyrus Latreille, 1810] HYPOCYSTINA Miller, 1968 [Mypocysta Westwood, 1851] YPTHIMINA Reuter, 1896 [Ypthima Hübner, 1818] EUPTYCHIINA Reuter, 1896 [Euptychia Hüb- ner, 1818] COENONYMPHINA Tutt, 1896 [Coenonympha Hiibner, 1819] MANIOLINA Grote, 1897 [Maniola Schrank, 1801] EREBIINA Tutt, 1896 [Erebia Dalman, 1816] DIRINA Verity, 1953 [Dira Hübner, 1819] PRONOPHILINA Reuter, 1896 [Pronophila Doubleday, 1849] SATYRINA Boisduval, 1833 [Satyrus Latreille, 1810] MELANARGIINA Wheeler, 1903 [Melanargia Meigen, 1828] 1864 [Bia Hübner, Bonner zoologische Beiträge Band 53 (2004) Heft 1/2 Seiten 255-282 Bonn, Juni 2005 Revision of the vincta Species-group of Monolepta Chevrolat, 1837 from _ Africa, Arabia and the Near East (Coleoptera: Chrysomelidae, Galerucinag) | Thomas WAGNER Universitat Koblenz-Landau, Institut fiir Integrierte Naturwissenschaften — Biologie, Koblenz Abstract. Specimens of Monolepta Chevrolat, 1837 from Africa, the Arabian Peninsula and the Near East, which are characterized by yellow elytra with transverse black bands at base and in the apical third, are revised. This coloration is typical for most specimens of M. vincta Gerstaecker, 1871, a widely distributed and the most abundant species of Monolepta in Africa. Nine junior synonyms of this species could be found: M. alternata Chapuis, 1879, syn. n.; M in- signis Weise, 1903, syn. n.; M. sjöstedti Weise, 1909, syn. n.; M. ugandaensis Laboissiere, 1920, syn. n.; M. lusingensis Laboissiere, 1920, syn. n.; M. bouvieri Laboissiere, 1920, syn. n.; M. striola Laboissiere, 1920, syn. n.; M. consociata Laboissiere, 1920, syn. n.; M. rugifrons Laboissiere, 1920, syn. n.; M. femoralis Laboissiere, 1940, syn. n. Further spe- cies herein revised are M. melanogaster (Wiedemann, 1823) (= M. bizonata Chevrolat, 1837, syn. n.); M. buguetii Chevrolat, 1837; M. lepida Reiche, 1858. Newly described species are M. sharonae sp. n., M. ronbeeneni sp. n., and M. naumanni sp. n. Lectotypes are herein designated for: M insignis Weise, 1903; M. sjóstedti Weise, 1909; M. lusingensis Laboissiere, 1920; M. femoralis Laboissiere, 1940. Distribution maps and an identification key for these species are given. Key words. Afrotropical region, Arabian Peninsula, Israel, taxonomy, biogeography, synonyms, lectotype, new species 1. INTRODUCTION In the last catalogue of the Galerucinae (WILCOX 1973), 180 species of Monolepta Chevrolat, 1837 from tropical Africa were listed. Most of these species were described between 1890 and 1950 (WAGNER 2003a). With very few exceptions, the descriptions by preceeding authors were based on external characters only. The allocation to Monolepta and other genera of the “Monoleptites” (WILCOX 1973) was mostly typological. In an ongoing revision of this group, the afrotropical species of Monolepta turned out as polyphyletic, and many species need to be transferred to other groups (WAGNER 2004). Many synonyms particularly in widely distributed spe- cies, but also many undescribed species could be found (WAGNER 2000a, b; 2001, 2002, 2003b). In this paper, afrotropical Monolepta, which have yel- low to yellowish-red elytra, usually two transverse black elytral bands at base and in the apical third, are revised. Head and prothrorax are mostly yellow to red, rarely black; abdomen is yellow, in a few species sometimes entirely black or rarely only last abdominal segments black. The revision includes the most abundant afro- tropical Monolepta species, M. vincta, which is de- scribed from Mombasa, Kenya. Wide distribution, high abundance and the high polymorphic coloration have * Revision of afrotropical Monolepta Chevrolat, 1837 — Part V. 19" contribution to the taxonomy, phylogeny and biogeography of afrotropical Galerucinae | In commemoration of Clas Michael Naumann zu Königsbrück (26.06.1939 — 15.02.2004) led to a high number of synonyms. A further two spe- cies of this coloration type have been listed in the De- jean catalogue by CHEVROLAT (1837), M. buquetii from Senegal and M. bizonata from the Cape of Good Hope. The latter could be identified as a junior synonym of M. melanogaster described by WIEDEMANN (1823) also from the Cape of Good Hope using specimens with en- tirely black abdomen. The fourth valid species is Monolepta lepida described by REICHE (1858) from Palestine, a species which occurs also throughout the Arabian Peninsula and in northeastern Africa. Since the original description of most species are very short and lack the description of genitalic patterns, redescriptions are given for all species. Monolepta buquetii was never described, but there exist an indication in the Dejean catalogue only. Finally, three species with this peculiar coloration are described as new. 2. METHODS A standard set of figures is given for each species. These include illustrations of the coloration (dorsal view), including the right antenna, where black colora- tion is indicated by black, yellow coloration by white. red coloration by dot-shading, and brown by black shad- ing. The basal four antennal articles of two different males and females, dorsal and lateral view of the me- dian lobe including the endophallic structures, and ven- tral view of the median lobe without the endophallic structures (for classification see WAGNER 2000), the spermathecae of three (if available) different females. and bursa-sclerites usually of one female are figured. 256 Morphometric measurements were made for external characters. Absolute measurements are: Total length from the clypeus to apex of the elytron, length of elytron, maximal width of both elytra (usually in the middle or posterior third of the elytra), and width of pronotum. Relative measurements are: Length to width of pronotum, maximal width of both elytra to length of elytron, length of second to third antennomere, and length of third to fourth antennomere. The number of specimens measured is given in the description under “total length”. 3. MATERIAL The subsequent redescriptions and descriptions are based on labelled specimens from the following collec- tions. Acronyms used and responsible curators in brack- ets: Bishop Museum, Honolulu (BPBM; A. Samuelson); The Natural History Museum, London (BMNH; S. Shute, M. Brendell, M. Cox); private collection Ron Beenen, Nieuwegein, The Netherlands (CBe); private collection Jan Bezdek, Brno, Cech Republic (CBz); pri- vate collection Manfred Döberl, Abensberg, Germany (CDö); private collection Uwe Heinig, Berlin, Germany (CHe); private collection Horst Kippenberg, Herzogen- aurach, Germany (CKi); private collection Joachim Mauser, Germany (CMa); private collection Vladimir Medvedev, Moscow, Russia (CMe); Deutsches Ento- mologisches Institut, Múncheberg (DEI; L. Behne, L. Zerche); Hungarian Museum of Natural History, Buda- pest (HNHM; O. Merkl); Instituto de Investigacáo Científica Tropical, Lisboa (IICT; L. F. Mendes); Insti- tute Royal des Sciences Naturelle de Belgique, Brussels (IRSNB; M. Cludts, D. Drugmand); Museo Civico di Storia Naturale, Genova (MCSN; R. Poggi); Museu de Catalunya, Barcelona (MCTB; G. Masó); Museo ed In- stituto di Zoologia Sistematica, Universitá di Torino (MIZT; M. Daccordi); Musée National d' Histoire Naturelle, Paris (MNHN; N. Berti); Museum für Naturkunde der Humboldt Universität zu Berlin (ZMHB; J. Frisch, H. Wendt, M. Uhlig); Musée Royal d’Afrique Centrale, Tervuren (MRAC; M. de Meyer); Museum of Zoology, Helsinki (MZHF; H. Silfverberg); Museo Zoologico “La Specola”, Firenze (MZUF; L. Bartolozzi); Naturhistorisches Museum Basel (NHMB; E. Sprecher-Ubersax, M. Brancucci); Naturhistorisches Museum Wien (NHMW; H. Schönmann); Naturhis- toriska Riksmuseet, Stockholm (NHRS; B. Viklund); National Museums of Kenya, Nairobi (NMK; W. Ki- nuthia, K. Maes); National Museum of National His- tory, Washington (USNM; D. Furth); National Museum of Namibia, Windhoek (NMNW; E. Marais); Oxford University Museum of Natural History (UMO; G. McGavin); South African National Collection, Plant Protection Research Institute, Pretoria (SANC; E. Grob- belaar); South African Museum, Cape Town (SAMC; Bonner zoologische Beitráge 53 (2004) M. Cochrane); Transvaal Museum, Pretoria (TMSA; S. Gussmann); Zoologisches Forschungsinstitut und Mu- seum Alexander Koenig, Bonn (ZFMK; M. Schmitt, K. Ulmen); Zoological Institute St. Petersburg (ZISP; A. Kirejtshuk); Zoological Institute University of Kopen- hagen (ZMUC; M. Hansen); Zoologisches Institut und Zoologisches Museum der Universitat, Hamburg (ZMUH; R. Abraham). For location data, geographical coordinates were given in degree and minute. These coordinates were mostly taken from the Alexandria Digital Library Gazetteer Server. Locations of the former Belgian Congo were in particular taken from a gazetteer compiled by Ugo Dall’ Asta (MRAC). 4. REDESCRIPTIONS AND DESCRIPTION OF SPECIES Monolepta melanogaster (Wiedemann, 1823) = Galleruca melanogaster Wiedemann, 1823: 77. = Monolepta bizonata Chevrolat, 1837: 407, syn. n. Redescription. Total length. 3.80-5.60 mm (mean: 4.81 mm; n= 12). Head. Red, frons often yellowish-red (Fig. 1), about 15 % of specimens examined with dark brown or black vertex (Fig. la). Antennae pale yellow to reddish- yellow, last antennomere or its tip only brown, rarely black, antennomeres 4 to 10 about 4 times long than broad at apex. Length of antennomeres two to three 0.86-1.00 (mean: 0.96), length of antennomeres three to four 0.30-0.41 (mean: 0.35). Third antennomere par- ticularly in large males very broad (Fig. 2a). Thorax. Prothorax reddish-yellow (Figs la, b), in about 20 % of material examined yellow (Fig. 1c). Pronotal width 1.15-1.70 mm (mean: 1.42 mm), pronotal length to width 0.60—0.65 (mean: 0.63), very finely punctured. Elytra pale yellow to yellow with broad black base which is often smaller than in (Figs la, b) and can be reduced to broad black margins like in Fig. Ic in about 10 % of material examined, subapical elytral transverse black band usually broad, rarely smaller, but not re- duced to a circular spot. Elytral length 2.90-4.20 mm (mean: 3.67 mm), maximal width of both elytra 1.90- 2.85 mm (mean: 2.58 mm), maximal width of both elytra to length of elytron 0.68-0.72 (mean: 0.70). Scutellum red to yellowish-red. Meso- and metathorax, coxa, trochanter an basal two thirds of femur red to red- dish-yellow, outer parts of legs yellow. Abdomen. Yellow, in 12 % of material examined black. Female genitalia. Spermatheca with spherical nodulus, comparatively broad middle part and cornu (Fig. 3). Thomas WAGNER: Revision of the vincta Species-group of Monolepta Chevrolat 25 Dorsal part of bursa sclerites triangular (Fig. 4a), ventral part slender, outer margin serrate to undulate (Fig. 4b). Male genitalia. Median lobe conical, straight, apical part lanceolate, slightly widened beyond apex, apical half with significantly visible short hairs (Fig. 5a). Tec- tum pointed, ventral groove parallel-sided, slender (Figs 5b, c). Dorsal pair median endophallic spiculae strong, slightly bent inwards, other median spiculae slender, straight, ventral spiculae separated in two portions, hooked at apex, two pairs of slightly curved lateral spi- culae, both of same size and orientation (Fig. Ib). Distribution. Widely distributed and abundant in southern Africa. In Central Africa northwards towards northeastern Congo and western Uganda (Fig. 6). Diagnosis and discussion. On average the largest spe- cies of the coloration type is characterised by two trans- verse black bands. Specimens with a total length of more than 5 mm from Central or southern Africa belong to this species. Only some specimens of M. lepida, which is restricted to Somalia, Eritrea and Egypt, and occurs also in Arabia and the Near East, can be up to 5.5 mm. Monolepta melanogaster has shorter basal anten- nomeres than M. lepida (length of second to third an- tennomeres: 0.86-1.00, M. lepida: 0.75-0.88; length of third to fourth antennomeres: 0.30-0.41, M. lepida: 0.46-0.54), has a broader pronotum (pronotal length to width: 0.60-0.65; M. lepida: 0.63-0.67) and both spe- cies are allopatrically distributed. Smaller specimens can be very similar to M. vincta, which is presumably most closely related to M. melanogaster. It can be distinguished on average by broader elytra (maximal width of both elytra to length of elytron: 0.68—0.72; M. vincta: 0.69-0.75), and shorter antennae. The median lobe is more homogeneously conical with slightly widened apex in M. melanogaster (Fig. 5b), while in M. vincta the apical part is more slender (Figs 23b, 24b). The endophallic armature of both species is very similar and only size and shape of the lateral spiculae show some constant differences (Figs 5b, 23b, 24b). Some specimens can be hardly distinguished by external and genital morphology. Some doubt remains if these are two distinct species, a question which might be only solved by using molecular data. At this stage I prefer a specific differentiation supported mainly by geographic distribution, and variation of body size and colour pat- tern. From two locations, Garamba National Parc in north-eastern Congo, and Kibale Forest in western Uganda large series were collected. Both species occur syntopicly at these sites and seem to be genetically separated. Monolepta vincta shows high polymorphic coloration (Figs 18c, e) at this site, while M. melano- gaster occurs only in one coloration type (Garamba Parc: Fig. Ic; Kibale Forest: Fig. Ib) and can be casily distinguished by significantly larger body size and by constant and comparatively high differences in male genital characters. This might be interpreted as character displacement of closely related distinct species when they occur syntopely. In South Africa and Namibia, and most parts of Zimbabwe, Zambia and southern Mogam- bique no specimens of “typical” M. vincta could be found and all specimens of the peculiar coloration type are assigned to M. melanogaster. In these countries many M. melanogaster are much smaller and on aver- age of the same size as M. vincta. Only specimens from some populations, particularly from coastal areas of the Cape Region are larger. Type material. Type specimens of Galleruca melano- gaster Wiedemann, 1823 were not available, but species identity can be derived from the original description and several specimens from old collections bearing identifi- cation labels. — Monolepta bizonata. Holotype: 7 “e. coll. Chev.' / Monolepta bizonata Chev. C. b. Sp. Holotype” (BMNH); type locality: South Africa, Cape of Good Hope, 33.56S/18.28E. CHEVROLAT (1837: 407) listed this species in the third edition of the Dejean cata- logue. The species was never described, but the name is available by indication (article 12.2, ICZN rules). Further material examined. Angola: | ex., Tchivinguire, Huila, 15.00S/13.20E, X1.1974 (NMNW); | ex., Cahama, 15 km E. 16.18S/14.28E, XII.1974 (NMNW). — Botswana: | ex., Betchua- naland, Gaberones, 24.40S/25.55E, IX.1915, Ellenberger (MNHN); | ex., Serowe, X1.1985, P. Forchhammer (NMHB); 10, Serowe, VII.-1X.1988, P. Forchhammer (SANC). — Burundi: | ex., Bururi, 3.58S/29.35E, P. Giraudin, ex coll. Breuning (MRAC). — Congo: | ex., Kasai, Kondué, 4.578/23.31E, E. Luja (MRAC); 1 ex., Kisantu, 5.07S/15.05E, P. Goosens (MRAC): | ex., Amadi, 3.35N/26.47E, 111.1913, van den Plas (MRAC); | ex., Malela, 2.28S/26.09E, X11.1913, L. Burgeon (MRAC); 1 ex., Al- bertville, 5.56S/29.12E, XII.1918, R. Mayné (MRAC): 3 ex. Haut-Uele, Moto, 2.27N/26.25E, 1920, IV.-V.1923, L. Burgeon (MRAC); 2 ex., Haut-Uele, Abimva, 3.09N/29.50E, V.—VII.1925, L. Burgeon (MRAC); 2 ex, Lomami, Kaniama, 7.34S/24.11E, 1931, R. Massart (MRAC); I ex., Equateur, 00.05S/18.14E, IV.1932, P. Hulstaert (MRAC); 1 ex., Eala, 0.04N/18.17E, IV.1936, Henrard (MRAC); 5 ex., Lukolela, 1.03S/17.12E, 1937, R. Massart (IRSNB); 5 ex., Bambesa, 3.28N/25.43E, 1V.1937, 1.1940, J. Vrijdagh (4 IRSNB, | MRAC): | ex., Bas Congo, May- idi, 5.11S/15.09E, 1942, van Eyen (MRAC); I ex., Lusinga, 8.56S/27.12E, 1760 m, IV.1947, G. F. de Witte (IRSNB): | ex.. Kalanga, 9.34S/27.08E, Grotte de Kakontwe, VIII.1948, N. Leleup (MRAC); 1 ex., Basoko, 1.14N/23.36E, 1V.1948, L. G. Benoit (MRAC); 1 ex., Stanleyville, Yaolema, 0.43N/24.28E, IV.1949, R. Laurent (MRAC); 102 ex., P. N. Garamba, 3.40N/29.00E, several locations, V.1950, IX.-X11.1951, V.1952 (54 ex. IRSNB, 48 ex. MRAC) 1 ex., Tshuapa. Bokuma, 0.065/18.42E, V.1952, P. Lootens (MRAC); 2 ex., Elisabethville, 11.40S/27.28E, 1.1956-1.1957, Ch. Seydel (MRAC). — Congo, Republic: | ex., reg. de Brazzaville, 4.00S/15.00E, 1906, Dupey (MNHN). — Malawi: 6 ex., Blantyre, 15.47S/35.00E, 1910, Dr. J E. S. Old (BMNH); 3 ex., Mlanje, 16.058/36.29E, X11.1912, S. A. Neave (BMNH) 5 ex. 50 km N Zomba. 15,225/35,22E, VI11.2000, L. Schmidt (ZFMK). — Mogambique: Liwonde, 258 Bonner zoologische Beitráge 53 (2004) 5a C Figs 1-5: Monolepta melanogaster (Wiedemann, 1823) 1: colour pattern; 2: basal antennal articles (a: Í, b: Y); 3: three different spermathecae; 4: bursa-sclerites (a: dorsal, b: ventral); 5: median lobe (a: lateral, b: dorsal, c: dorsal. Scale for colour pattern dif- ferent from same scales for basal antennal articles and genital structures: each 1 mm. Same for all following figures. | ex., Beira, 19.49S/34.52E, (SAMC); | ex., Chibababa, Lower Buzi R., 19.52S/34.45E, XII.1906, C. F. M. Swynnerton (BMNH); 3 ex., Pomene, 22.59S/35.35E, V.1975, beaten, costal bush, A. Strydom (TMSA). — Namibia: | ex., Kapako, Okavango, 18.015/21.22E, X11.1954 (NMNW); | ex., Andara, SE 1821 Ab, Kavango, 18.045/21.27E, VII1.1971 (NMNW); 11 ex., Nyangana, Okavango, 18.005/20.41E, 1.1985, H. Roer (ZFMK); 5 ex., E- Caprivi, Katima Mulilo, 17.29S/24.17E, 111.1992, U. Góllner, M. Uhlig, lux (ZMHB); 1 ex., Kavango, Mahango GR, 18.14S/- 21.43E, X.1993, F. Koch (ZMHB); 2 ex., Popa falls, 18.07S/- 21.34E, X11.1993, M. Uhlig (ZMHB); | ex., Okavango, Bagani, Popa falls, 18.07S/21.34E, 1.1995, F. Kantner (CKa); 1 ex., Popa Camp, 11.1998, U. Góllner (ZMHB). — South Africa: 3 ex., Cap b. Sp., 33.568/18.28E, 1817, Wied.”, Mus. “melanogaster Thomas WAGNER: Revision of the vincfa Species-group of Monolepta Chevrolat 259 Westerman (ZMUC); | ex., ex coll. Laferté, „Monolepta bizonata Chevr.; C. B. Sp. Verraux“ (BMNH); I ex., Cap Diege, coll. Haag, “Monolepta Er. / bizonata Chey.” (DEI); | ex., coll. Clavareau, coll. Chapuis, “Monolepta pulchella Klug / Monolepta duplicata Sahlb. = pulchella 1, Klug (MRAC); | ex., Cape Col- ony, coll. Clavareau, Monolepta pulchella Klug, Determinat Ja- coby (MRAC); 2 ex., “Galleruca atrofasciata Diege Cap” (Halle); 1 ex., “alternans, N, Crio. bifasciata, Fab.?, Krebs, Pr. B. Sp., Li- chtenst. / 30377 (ZMHB); 8 ex., Cap, ex coll. J. Weise (ZMHB); | ex., Cap, coll. de Bonvouloir (MNHN); 4 ex., P. B. Sp. (UMO); | ex., Cap B. Spei, Victoria (NHRS); I ex., Cape Town, coll. Jaco- by (BMNH); 2 ex., Alexandria, Dorn Nek, J. L. Drege (SAMC); 5 ex., Caffraria, J. Wahlberg (NHRS); I ex., Caffra, coll. Fry (BMNH); 2 ex., Capland, Willamore, Dr. Brauns (BMNH); 3 ex., Kapland (ZMHB); | ex., Dunbrody, Cape Colony, 33.28S/25.33E, coll. Jacoby (BMNH); 4 ex., Transvaal, Fruhsdorfer (ZMHB); 4 ex., Transvaal (BMNH, MNHN, ZMHB, ZMUH); 6 ex., Trans- vaal, Lydenburg, 25.06S/30.27E, F. Wilms (ZMHB); 4 ex., Transvaal, Lydenburg (IRSNB); | ex., Transvaal, Pietersburg (BMNH); 1 ex., Grahamstown, 33.175/26.32E, C. le Doux (ZMHB); 5 ex., Barberton, 25.48S/31.03E, P. Rendaff (BMNH); 4 ex., Natal (1 BMNH, 3 MNHN); | ex., Natal, Marshall, Jacoby coll. (BMNH); 4 ex., Natal, Estcourt, 29.008/29.53E (SAMC); | ex., Natal, Frere, 28.53S/29.46E (SAMC); I ex., Wydah, R. P. Ménager (MNHN); I ex., Port Natal, 28.30S/30.30E, Boheman (ZMHB); 3 ex., Zoutpansberg, 28.10S/32.15E, Shilouvane, Junod, coll. Clavareau (MRAC); | ex., Port Elisabeth, 33.50S/25.45E, 1882, S. D. Bainslow, “Monolepta bifasciata, named 1899 by M. Jacoby” (UMO); 2 ex., Natal, Frere, 28.53S/29.46E, VI., X.1892, G. A. K. Marshall, on roses (BMNH); 11 ex., Natal, Estcourt, VIT.-X.1892, IX., IX., X1.1896, G. A. K. Marshall (BMNH); | ex., Transvaal, Hammanskraal, 25.24S/28.16E, 1893, E. Simon (MNHN); | ex., Oranje FS, Mateles Pic, 2400 m, 1.1896, J. Thode (ZMHB); | ex., Maritzburg, 29.38S/30.24, 11.1896, P. Cregoe (MNHN); 6 ex., Natal, Malvern, 29.53S/30.55E, 1.1896, IH., VILL, X.1897, X11.1899, G. A. K. Marshall (BMNH); | ex., Natal, Um- gent, 29.48S/31.03E, VIL1897, G. A. K. Marshall (BMNH); | ex., C. Bon. Spei, 1900, coll. Fry (BMNH); | ex., Transvaal, Sterkfon- tein, 24.42S/30.15E, 1900, H. P. Thomasset (BMNH); | ex., Na- tal, Upper Tongaat, 29.34S/30.45E, Barwon, 1600 ft, X.1901, G. A. K. Marshall (BMNH); | ex., Lower Tugela, 29.14S/31.30E, X.1901, Monolepta melanogaster Wied. (SAMC); | ex., Natal, Lower Tugela, 1902, E. Reynolds (BMNH); 2 ex., Durban, 29.518/31.01E, 1902, F. Muir (BMNH); | ex, Transvaal, Zout- pansberg, 28.10S/32.15E, X11,1907, Knothe (ZMHB); 1 ex., Escourt, Mrs. E. J. Turner, 1909 (BMNH); | ex., Natal, Nw. Hanover, 28.035/29.39E, X.1914 (SANC); 4 ex., East London, 33.02S/27.54E, IX., XII.1915, 1923, R. Ellenberger (MNHN); 2 ex., Pietermaritzburg, Fort Napier, 29.38S/30.25E, 1919 (ZMHB) ; 7 ex., Transvaal, Piet Retief, 27.00S/30.49E, 11.1920, feeding on petals of bean flowers (3 BMNH, 4 SANC); | ex., Na- tal, Durban, VII1.1920, A. F. J. Gedye (NMK); | ex., Umatata, Transkei, 31.35S/28.47E, 11.-111.1923, R. E. Turner (BMNH); 4 ex., Port St. John, 31.255/29.30E, Pondoland, IV., VII.1923, R. E. Turner (BMNH); | ex., Natal, Weenen, 2840 ft, 28.05S/30.06E, VI1.-IX.1923, H. P. Thomasset (BMNH); | ex., Aliwal North, Cape Prov., 30.415/26.42E, X11.1923, R. E. Turner (BMNH); 1 ex., Transvaal, Machadodorp, 25.40S/30.15E, IV.1926, M. Mos- sop (SANC); 4 ex., S. Zululand, Gingindhlovu, V.1926, R. E. Turner (BMNH); | ex., Zululand, Eshowe, 2853S/31.28E, Turner (BMNH); 2 ex., Natal, Drakensberg, 28.435/28.53E, van Reenen (BMNH); | ex., Oranje FS, Harrismith, 28.17S/29.08E, 111.1927, R. E. Turner (BMNH); 2 ex., Somerset East., Cape Prov., 32.438/25.35E, IX.1930, R. E. Turner (BMNH); I ex., Natal NaFauresmith, 29,458/25.19E, V.1932, Dr. Henrici (1 BMNH, 4 SANC); 7 ex., Natal, Dundee, 28,105/30,13E, VI11.1932 (2 BMNH, 5 SANC); 4 ex., Katberg, E. Cape Prov., 32,325/26.411 1.1933, R. E. Turner (BMNH); | ex., Bainskloof, 33,355/19.081 IX.1938, L. D. Brongerama (NNML); I ex., Natal NP, 60m y Ladysmith, X.1938, L. D. Brongerama (NNML); | ex., Zululand Mkuzi, X11.1945, DDT killed (SANC); 2 ex., Pretorius Kop. X.1950, H. K. Munro (SANC); | ex., Schoemanskloof, E. T., VI.1951, H. K. Munro (SANC); | ex., Natal, Richmond, X11.1954, ex coll. Breuning (MRAC); | ex., Natal, Durban, 1.1955, K. Mannonen (Helsinki); 3 ex., Natal, Indaleni, 29.54S/30.17E, X.1956, Rev. W. Hunt, ex coll. Breuning (MRAC); | ex., Natal, Southbroom, 30.55S/30.19E, VI1.1956, ex. coll. Breuning (MRAC); | ex., Politzi, VI1.1966, P. Pali. (SANC); 1 ex., 32 m SW Loius Trichardt, X.1966 (SANC); I ex., Cape Prov., Kei Bridge, 32.31S/27.58E, 1.1970, Cl. Besnard (MRAC); 2 ex., Zululand, Ndugu GR, 26.55S/32.19E, IV.1970, C. Besnard (MRAC); | ex., Zululand, Hluhluwe GR, 28.05S/32.04E, IX.1970, O. Bourquin (TMSA ); 2 ex., Transvaal, Debegeni Falls, 5 km SI Magoebaskloof, 11.1973, A. Prinsloo & T. Bouweri (SANC); | ex., Transvaal, Tswane, 111.1973, A. Prinsloo (SANC); | ex., Zu- luland, Dukuduku Forest, 28.22S/32.19E, IV.1974, S. Endrödy- Younga, indig. forest litter (TMSA); 1 ex., Zululand, Empangeni, 28.45S/31.54E, IV.1975, P. E. Reavel (TMSA); 5 ex., Zululand, St. Lucia, Mission Rock, 28.22S/32.35E, X11.1975, S. Endródy- Younga, at black light (TMSA); | ex., Natal, St. Lucia Mission Rocks, 28.22S/32.25E, XII.1975, S. Endrödy-Younga, at uv-light (TMSA); | ex., Transvaal, Naboomspruit, Libertas, 24.27S 28.33E, X1.1976, C. G. E. Moolman (SANC); 1 ex., Natal, Nyala GR nr Empangeni, 28.42S/31.46E, VIII.1977, P. Reavell (SANC): l ex., Austenburg, Kloof, IV.1978, coll. M. Keeping (TMSA): | ex., Transvaal, Pretoria, 25.45S/28.12E, X11.1978, C. G. Moolman (SANC); 2 ex., Transvaal, Barberton, 25.48S/31.03E, 111.1979, C. Moolman (SANC); 1 ex., Transvaal, Grahamstown, 111.1979, G. L. Prinsloo (SANC); 2 ex., Transvaal, Mogol NR, Ellisras Distr. 23.58S/27.45E, X1.1979, 11.1984, C. D. Eardley (SANC); I ex., Transkei, Dwesa Forest, 32.17S/28.50E, XI1.1979, S. Endrödy- Younga, flowering Acacia (TMSA); 4 ex., Transvaal, Mogoto NR, Zebediela, 24.15S/29.13E, X.1979, 111.1980, G. L. Prinsloo et al. (SANC); 1 ex., Percy Fyfe NR, 24.03S/29.09S, I11.1980, C. Kok (SANC); 8 ex., Kwa-Zulu, lake Sibaya, E shore. 27.22S/32.43E, 1.1981, C. D. Eardley (SANC); 1 ex., Natal, Not- tingham Road, 29.22S/29.59E, 1.1981, S. J. v. Tonder & C. Kok (SANC); | ex., Natal, Cape Vidal, 28.10S/32.32E, 1.1981, R. G. Oberprieler (SANC); 1 ex., Transvaal, Pretoria, Roodeplaat, 25.41S/28.18S, 1.1983, C. Moolman (SANC); 1 ex., Cape Prov- ince, Knysna, 34.02S/23.03E, X1.1983, R. Oberprieler (SANC): 1 ex., Natal, Ngoye Forest, 1984, inside cumbers (BMNH): 2 ex., Natal, Pietermaritzburg, Town Bush Forest, 29.36S/30.23E. 11.1984, C. G. E. Moolman (SANC); | ex., Natal, Mt. Nhlosane nr Dargle, 1600 m, 29.33S/29.56E, 11.1984, C. G. E. Moolman (SANC) 1 ex., Natal, Wembesi nr Estcourt, 29.04S/29.46E, 11.1984, C. G. E. Moolman (SANC); 1 ex., Natal, Balgowan, 29.238/30.20", 11.1984, R. Cberprieler & C. G. E. Moolman (SANC); 3 ex., Franskraal distr., Suikerbosnandfarm. W. Cape. IX.1984, W. Wittmer (NHMB); 2 ex., Kogmanskloof. 200 m, Cape Prov., X.1984, W. Wittmer (NHMB); I ex., S. Cape Prov.. Herbertsdale, 34.01S/21.46E, X.1984, R. Müller (TMSA): 5 ex., E Cape Prov., 35 km N Grahamstown, 32.575/26.09E, X.1984, S. Endrödy-Younga, ground & vegetation (TMSA): | ex.. Cape Province, Mitchell’s Pass nr Ceres, 33.23S/19.27E, X1.1984, C. Eardley (SANC); | ex., Cape Province, Saasveld nr George, 33.578/22.35E, 11.1985, W. Breytenbach (SANC); 1 ex., S. Natal, Weza-Lovedale, 30.40S/29.41E, 111.1985, S. Endródy-Younga, sift. indig. for. litt. (TMSA); I ex., Transvaal, Messina, V.1985, 260 Naudé (SANC); 1 ex., SW Cape, Clanwilliam, 32.09S/18.53E, IX.1985, on vegetation on flood plain (TMSA ); I, Transvaal, Let- sitele, 23.525/30.24E, X.1985, S. Kamburow (SANC); 1 ex., Transvaal, Bourkes Luck, 24.40S/30.48E, 11.1986, C. D. Eardley (SANC); 1 ex., Transvaal, Lotus Trichardt, 800 m,23.03S/29.49E, 11,1986, D. d’Hotman (TMSA); | ex., E. Cape, Grahamstown, 33.19S/26.31E, VI.1886, S. van Noort (SAMC); | ex., Meirings- poort, 33.248/22.33E, IX.1986, N. MacPherson (SAMC); 5 ex., Transvaal, Entabeni FR, 33.00S/30.16E, 1.1987, C. D. Eardley (SANC); I ex., Transvaal, Hans Merensky NR, 23.42S/30.44E, 1.1987, B. Grobbelaar (SANC); 5 ex., Transvaal, Swadini, Blyde- poort NR, 24.32S/30.54E, 1.1987, B. Grobbelaar (SANC); l ex., Transvaal, Modjadji NR, 23.38S/30.20E, 1.1987, C. D. Eardley (SANC); 5 ex., Transvaal, Nelshoogte Knuckles Rocks Forest, 25.47S/30.50E, 11.1987, S. Endrödy-Younga, beating in forest (TMSA); 1 ex., Transvaal, Uitsoek waterfall, 25.16S/30.33E, 11.1987, S. Endrödy-Younga, beating (TMSA); 7 ex., OFS, Tus- sen Die Riviere Res. nr Bethulie, 30.30S/26.12E, III.-IV.1987, B. Grobbelaar (SANC); 2 ex., Transvaal, Wyllieespoort, Ingwe Mo- tel, 22.58S/29.57E, 11.1988, G. D. Butler (SANC); 1 ex., Trans- vaal, Kruger NP, Satara, 24.23S/31.46E, 350 m, II.1988, B. Grob- belaar (SANC); 2 ex., Transvaal, Meiringspoort, Groot Swartberge, 33.335/22.19E, X1.1988, R. Oberprieler (SANC); 2 ex., Cape Province, Suurberg NR, 33.16S/25.45E, X1.1988, R. Oberprieler (SANC); 1 ex., Cape Province, Phanton Pass nr Knysna, 34.00S/22.59E, X11.1988, R. Oberprieler (SANC); 2 ex., Cape Province, Brenton-on-Sea, Knysna, 34.048/23.01E, X11.1988, E. Grobbelaar (SANC); 3 ex., Transvaal, Barberton, 25.50S/31.02E, 1.1988, E. Holm & E. Marais (NMNW); 1 ex., Natal, 27 km S Empangeni, 28.57S/31.43E, 11.1989, B. Grobbe- laar & E. v. d. Linde (SANC); 1, Gankapoordam, Cape Prov., V111.1989, G. Minet (NHMB); | ex., 23 km N East London, IX.1989, W. Wittmer & S. Gussman (NHMB); 4 ex., Transkei, Port St. John, Silaka, 31.335/29.30E, XI.-X11.1987, S. Endródy- Y ounga, at uv-light (TMSA ); | ex., Transvaal, Montrose State Fo- rest, Barberton, 25.50S/31.02E, 1.1988, E. Holm & E. Marais (NMNW); 1 ex., Transvaal, Uakhutswa River, Ofcolaco, 24.05S/30.22E, 1.1989, Werner (cDö); 4 ex., Transvaal, Witpoort- jie, Botanical Garden, 26.07S/27.50E, 11.1989, B. Grobbelaar (SANC); 3 ex., Transvaal, Lekgalameetse NR, 24.10S/30.14E, 11.1989, V. M. Uys / N. Verheijen (SANC); 6 ex., Natal, Cathkin Peak Hotel, 29.00S/29.27E, IV.1989, E. v. d. Linde (SANC); 1 ex., Natal, Overstone nr Fawn Leas Farm, 29.22S/30.36E, X.1989, E. v. d. Linde (SANC); 3 ex., Transvaal, Weltevreden Farm nr. Nelspruit, 25.34S/31.10E, 11.1989, 1.1990, R. Ober- prieler, V. M. Uys (SANC); 2 ex., Natal, St Lucia Estuary, 28.17S/32.25E, 11.1989, B. Grobbelaar & E. v. d. Linde (SANC); l ex., 20 km S Mkuze, 27.56S/32.13E, Natal, IX.1989, W. Witt- mer & S. Gussmann (MNHB); | ex., Transvaal, Roodeplast dam nr Pretoria, 24.41S/28.18E, X.1989, J. S. Donaldson (SANC); 1 ex., Cape Province, Keurboomrivier Mond nr. Plettenberg Bay, 34.02S/23.24E, 11.1990, M. Jonsson (SANC); 2 ex., Cape Prov- ince, Swartberg pass, 33.19S/22.03E, 11.1990, M. Jonsson (SANC); 2 ex., Cape Province, Kango Mts Ressort nr Oudtshoorn, 33.31S/22.21E, 11.1990, M. Jonsson (SANC); 2 ex., Beacon Bay, Cape Prov., X.1990, W. Wittmer (NHMB); 2 ex., 70 km W Grahamstown, X.1990, W. Wittmer (NHMB); | ex., Tran- skei, Lusikisiki, 31.21S/29.35E, X1.1991, Richter (MIZT); 2 ex., Pretoria, 25.45S/28.12E, X1.1992, W. Wittmer (NHMB); | ex., Wonderdal Farm, Graaff-Reinet, 32.15S/24.32E, 800 m, X1.1992, W. Wittmer (NHMB); | ex., Port Elisabeth, Addo Elephant NP, 30.50S/25.45E, X11.1992, F. Koch (ZMHB); 1 ex., Groenfontain, Bonner zoologische Beiträge 53 (2004) SW of Naboomsprui, 24.38S/28.31E, 11.1993, E. Grobbelaar (SANC); 1 ex., Dragon Peaks Park, 1150-1450 m, 29.02S/ 29.26E, X1.1993, J. Deckert, lux (ZMHB); 4 ex., Natal, Drakens- berg, 29.03S/29.24E, IX.1993, 11.1994, U. Göllner, F. Koch (ZMHB); 6 ex., Natal, Itala GR, 27.30S/31.20E, 1.1994, 11.1995, U. Göllner, F. Koch (ZMHB); 1 ex., Natal, Spa ca. 20 km SES Paulpietersburg, 27.32S/30.57E, 11.1994, U. Góllner (ZMHB); 1 ex., Natal, Sodwana Bay NP, 27.37S/32.41E, 1.-11.1994, U. Góll- ner (ZMHB); 1 ex., Natal, Mkunza NP, 27.36S/32.13E, 11.1994, F. Koch (ZMHB); 3 ex., KwaZulu-Natal, St. Lucia Park, 28.12S/32.25E, 11.1995, F. Koch (ZMHB); | ex., KwaZulu-Natal, N'dumu GR, 26.55S/32.19E, 11.1995, F. Koch (ZMHB); 1 ex., KwaZulu-Natal, Pongola River, Makanes Drift, 27.01S/32.18E, 11.1995, F. Koch (ZMHB); 5 ex., Cape Prov., Citrusdal, Olifants- river, 32.36S/19.08E, IV.1995, U. Göllner (ZMHB); 2 ex., Bont- bok NP, 34.04S/20.27E, IV.1995, J. Deckert, U Göllner (ZMHB); l ex., KwaZulu-Natal, Sodwana Bay NP, 27.37S/32.41E, X1.1995, F. Koch (ZMHB); 1 ex., Bophutatswana, Pilansberg NP, 25.158/27.13E, 1200-1500 m, X1.1996, M. Hartmann (NME); 3 ex., Legalameetse NR, Makhutsi Camp, 24.12S/30.18E, 1200 m, 1.1997, E. Grobbelaar (SANC); 1 ex., Tshipise, Nonet NR, 22.37S/30.10E, 320 m, X1.1998, M. Hartmann (NME); 1 ex., Ka- roo, Graaf-reinet Camp, 32.14S/24.32E, 500 m, XI.-XII.1996, M. Hartmann (NME); 1 ex., W-Cape, Greyton, 34.03S/19.37E, X.1999, M. Snizek (MIZT). — Uganda: 1 ex., Tero Forest, SE Buddu, 0.50S/31.40E, IX.1911, S. A. Neave (BMNH); 1 ex., Bugiri, 0.34N/33.45E, 1400 m,VIII.1957, P. Basilewsky & N. Leleup (MRAC); 176 ex., Kibale Forest, 0.50N/31.06E, VIL, IX.- X.1983, VI, IX.1984, IV.-V.1985, IX.1986, M. Nummelin (Hel- sink1); 2 ex., Ft. Portal, Kibale Forest, 0.45N/31.00E, V.-IX.1992, coll. Maus (ZFMK); 5 ex., Kibale Forest, Kanyaware, 1600 m, VI1.-V111.1998, L. Schmidt (ZFMK). — Zambia: | ex., Victoria Falls, 17.55S/25.51E, 3000 ft, IX.1905, B. E. Poulton (UMO); 11 ex., Victoria Falls, Palm Kloof, IX.1905, B. E. Longstaff (UMO); l ex., Lofu River, 8.345/30.44E, 3500 ft, VIIL 1909, S. A. Neave (UMO); 2 ex., Upper Luangwa, 14.49S/19.06E, 1880-2000 ft, 111.1908, S. A.Neave (BMNH, UMO); 1 ex., Mid-Lungwa, 14.00S/19.00E, VIII.1910, S.A. Neave (BMNH); 1 ex., Niamadzi R., nr. Nawalia, VIII.1910, S. A. Neave (BMNH); 1 ex., Mwengwa, 13.005/27.40E, VUI.1913, H. C. Dollman (BMNH). — Zimbabwe: | ex., Mashonaland, Jacoby coll. (BMNH); 2 ex., Matabele, Hard af Seg. (NHRS); 2 ex., Salisbury, 17.43S/31.05E, Jacoby coll. (BMNH, SAMC); 1 ex., Umchaki River, 11.1897 (BMNH); 5 ex., Salisbury, H.1895,VHI.1898, 11.1899, VI.— VII.1900, G. A. K. Marshall (BMNH); 6 ex., Salisbury, I., II., V., VIII.1898 (BMNH); 2 ex., Salisbury, VIII.1898, on Mosasa, Jacoby coll. (BMNH, SAMC); | ex., Old Umtali, 19.00S/32.40E, X.1897, G. A. K. Marshall (BMNH); 3 ex., Mashonaland, Chir- inda, 20.05S/31.28E, X.1905, G. A. K. Marshall (BMNH); | ex., Selukwe, 19.40S/30.02E, 1915, A. Ellenberger (MNHN); 1 ex., near Fort Victoria, 20.10S/30.49E, IX.1931, J. Ogilvie (BMNH); 2 ex., Umtali, 19.00S/32.40E, Xmas Pass, V.1932, J. Ogilvie (BMNH); 1 ex., Imbezi Valley, HI.1957, N. L. H. Krauss (BMNH); | ex., Melsetter, 19.48S/32.52E, 1700 m,VII.1960, N. Leleup (MRAC); 1 ex., Mpika, Muchinga Mts., 11.15S/31.27E, 1700 m, VII.1960, N. Leleup (MRAC); 1 ex., Khami ruins, Bula- wayo, 20.15S/28.30E, 1350 m, X1.1987, W. Wittmer (NHMB); 1 ex., Kyle RP, Lake Mutirikwi, 20.13S/31.00E, XIL1993, F. Koch (ZMHB); 1 ex., Chimanimani, 19.47S/32.50E, X11.1998, S. Becvár (CBz); l ex., 15 km SE of Muzarabani, 16.20S/31.10E, XII.1998, S. Beévar (CBz). Thomas WAGNER: Revision of the vincta Species-group of Monolepta Chevrolat 261 etme M. buquetii + 1 $ 2-4 we 5-15 Fig. 6: Distribution of Monolepta melanogaster (Wiedemann, 1823), and M. buquetii Chevrolat, 1837 Monolepta buquetii Chevrolat, 1837 Description. Total length. 3.00-4.20 mm (mean: 3.58 mm; n = 8). Head. Including mouth parts yellowish-red, in some specimens labrum and frons yellow, vertex yellowish- red, vertex very broad (Fig.7). Antennae pale yellow throughout, rarely tip of last antennomere dark brown, antennomeres four to eleven short, less than 4 times long than broad at apex, third antennomere very short in males (Fig. 8a), more elongated in females (Fig. 8b). Length of antennomeres two to three 1.00-1.40 (mean: 1.17), length of antennomeres three to four 0.20-0.36 (mean: 0.30). Thorax. Prothorax entirely pale yellow. Pronotal width 0.95-1.20 mm (mean: 1.11 mm), pronotal length to width 0.54-0.58 (mean: 0.57), pronotum very finely punctured. Elytra pale yellow with narrow transverse black band at base, and each elytron with transverse black spot in the apical third, often much smaller than in Fig. 7a. Rarely elytra with broad black basal and outer margins reaching towards the apical third and some- times joining with the subapical spot (Fig. 7b). Elytral length 2.55-3.15 mm (mean: 2.68 mm), maximal width of both elytra 1.60-2.10 mm (mean: 1.92 mm), maximal width of both elytra to length of elytron 0.69-0.73 (mean: 0.72). Scutellum yellow. Meso- and metathorax reddish-yellow, legs completely pale yellow. Abdomen. Yellow to reddish-yellow. Female genitalia. Spermatheca with spherical nodulus, comparatively broad middle part and short. slender cornu (Fig. 9). Dorsal part of bursa sclerites with nar- rowed, hook-like base (Fig. 10a), ventral part slender, outer margin very finely serrate (Fig. 10b). Male genitalia. Median lobe very slender in the apical half (Fig. 11), apex usually rounded (Figs 11b, d), rarely pointed (Fig. 11c). Tectum and ventral groove of me- 262 dian lobe narrow (Figs b, c). One pair of median endo- phallic spiculae strong, significantly bent at apex, ven- tral spiculae slender, apically hooked, lateral spiculae very small, triangular (Fig. 1 1b). Distribution. Known from savannas of West and north- ern Central Africa (Fig. 6). Diagnosis. Monolepta buquetii is one of the smallest Monolepta species known from Africa. Size and colora- tion is most similar to M. vincta and M. sharonae sp. n. Males can be easily distinguished by the very short third antennomere (Fig. 8a). Generally, short antenna, broad head and the transverse pronotum are good external, and shape of median lobe and spermatheca are excellent genital diagnostic characters of M. buquetii. Type material. Holotype: © “E. Coll. Laferte / Monolepta Buquetii Dj. Senegal / Holotype” (BMNH); type locality: Senegal. CHEVROLAT (1837: 407) listed this species in the third edition of the Dejean catalogue. It was not described, but the name is available by indi- cation (article 12.2, ICZN rules). Further material examined. Congo: P. N. Garamba, 3.40N/ 29.00: 4 ex., l/a/l, 1/b/3, 1/c/1, 1.-11.1950, H. de Saeger (2 ex. IRSNB, 2 ex. MRAC); 5 ex., V/a/l, I/b/1, U., IX.1950, G. De- moulin (IRSNB); | ex., H/fe/18, X1.1951, H. de Saeger (IRSNB); I ex., I/gd/4, 111.1952, H. de Saeger (IRSNB); 2 ex., PFSK.25/3, V1.1952, H. de Saeger (IRSNB); 12 ex., P. N. Garamba, Ndelele, 4.22N/29.47E, IL.—IIL., V11.1952, H. de Saeger (8 ex. IRSNB, 4 ex. MRAC); 2 ex., 2/source, VI.1952, H. de Saeger (IRSNB). — Ivory Coast: 4 ex., Comoé NP, 8.75N/3.80W, VI—VII.1997, C. Mody (ZFMK). — Senegal: | ex., Guede, 11.1946 (BMNH). Monolepta lepida Reiche, 1858 Redescription. Total length. 3.80-5.30 mm (mean: 4.68 mm; n = 10). Head. Pale yellowish-red (Fig. 12a), frons sometimes yellow (Fig. 12b). Antennae pale yellow, tip of terminal antennomere black, antennomeres four to eleven about 4 times longer than broad at apex. Basal antennomeres elongated, third antennomere significantly longer than second particularly in males (Fig. 13a), length of anten- nomeres two to three 0.75—0.88 (mean: 0.84), length of antennomeres three to four 0.46—0.54 (mean: 0.49). Thorax. Prothorax pale yellow to yellowish-red (Fig. 12). Pronotum narrow, width 1.10-1.55 mm (mean: 1.33 mm), pronotal length to width 0.63-0.67 (mean: 0.66), very finely punctured. Elytra pale yellow, base with narrow transverse black band (Fig. 12a), rarely black coloration reduced to spots at humeri (Fig. 12b), subapical transverse elytral band narrow. Some speci- mens from Oman have a yellowish-red head, pronotum and elytra with larger, more circular subapical elytral spots. Elytral length 2.80-3.80 mm (mean: 3.41 mm), maximal width of both elytra 2.00-2.70 mm (mean: 2.34 mm), maximal width of both elytra to length of Bonner zoologische Beitráge 53 (2004) elytron 0.65-0.70 (mean: 0.68). Scutellum yellow to yellowish-red. Meso-, metathorax, and legs usually yel- low, rarely yellowish-red, but tibia and tarsus always pale yellow. Abdomen. Pale yellow to yellow. Female genitalia. Spermatheca with spherical nodulus, comparatively broad middle part and cornu (Fig. 14). Dorsal part of bursa sclerites slender, with row of hooks at base (Fig. 15a), ventral part very slender, outer mar- gin very finely serrate (Fig. 15b). Male genitalia. Median lobe conical, straight, apical part spoon-like widened (Fig. 16a), ventrally bent (Fig. 16b). Tectum slender, pointed, ventral groove parallel- sided, very slender (Fig. 16c). Dorsal pair of median endophallic spiculae stronger, very long, other median spiculae slender, long, ventral spiculae separated in two portions, hooked at apex, lateral spiculae slender, long, bifurcate (Figs 16a, b). Distribution. Most specimens are known from the Ara- bian Peninsula and this species reaches the Palaearctic Region in Israel and Jordan. A few specimens are known from Eritrea and Somalia (Fig. 17), and occurs most likely also in Egypt and eastern Sudan. Diagnosis. Most similar to M. vincta and both species occur sympatrically in north-east Africa. Including M. melanogaster, these three species are most likely a mo- nophyletic group, which can be derived from external characters, coloration, and male genital patterns, par- ticularly the type of the endophallic armature. In com- parison to M. vincta, M. lepida is on average larger, and has reduced black elytral coloration, while syntopic M. vincta often has broad transverse black elytral bands. Good diagnostic external characters are the elongated second and third antennomeres in M. lepida (length of second to third antennomeres: 0.75—0.88, M. vincta: 0.86-1.00; length of third to fourth antennomeres: 0.46— 0.54, M. vincta: 0.27-0.35) and the narrow pronotum (pronotal length to width: 0.63-0.67, M. vincta: 0.57— 0.64). O « Type material. Holotype: Y “Vog. de Saulcy, Jerusa- lem / Monolepta lepida Reiche 1858 / Muséum Paris 1952 coll. R. Oberthur / Holotype” (MNHN); type lo- cality: Israel, Jerusalem, “des bords du Jourdain” which is about 30 km east of the centre of Jerusalem. REICHE (1858) gave a detailed description of external characters and coloration including an excellent figure (pl. 1, fig. 10). His description based most likely on a single specimen “M. Azambre, a qui dois un bel individu de cette espéce, ...” which is considered as holotype. The original description was repeated in JOANNIS (1866). Further material examined. Eritrea: 2 ex., Agorgat, 15.33N/ 37.53E, 1.1906, D. Figini (MCSN); 23 ex., Assab, 13.00N/42.44E, 1907, Katona (HNHM). — Israel: 2 ex., Wadi Arugod, Ein Gedi, Thomas WAGNER: Revision of the vincta Species-group of Monolepta Chevrolat 6 /a 10a b Figs 7-11: Monolepta buquetii Chevrolat, 1837 7: colour pattern; 8: basal antennal articles (a: ©, b: 11a b E d ); 9: two different sper- mathecae; 10: bursa-sclerites (a: dorsal, b: ventral); 11: median lobe (a: lateral, b: dorsal, c: dorsal, variation with pointed apex, d: ventral, without endophallic structures). 31.27N/35.23E, 111.1963, W. Wittmer (CMe). — Jordan: | ex., Wadi Schaib, 30.30N/36.00E, 100 m, X1.1959, J. Klapperich (NHMB). — Oman: 7 ex., Dhofar, 17.00N/54.10E, X.1979, T. B. Larsen (NHMB); 1 ex., Dhofar, Ain Rzat, Salalah, 17.01N/54.06E, 11.1989, W. Wittmer (NHMB); 2 ex., 30 km W Salalah, Wadis nr. Al Mughsayi, 17.00N/53.50E, 11.1998, G. Wewalka (NHMB). — Saudi Arabia: | ex., Jebel Ibrahim, 20.25N/41.11E, 1540 m, VIII.year?, W. Büttiker (NHMB); 3 ex., Shileam, X1.1935, R. C. M. Darling (BMNH); 2 ex., Wadi Johan, Abha, 18.12N/42.30E, 2150 m, IV.1976, Wittmer & Büttiker (NHMB); I ex., Wadi ad Dilla, 1120 m, X.1979, W. Büttiker (NHMB); | ex., Al Alayyah, 19.36N/41.58E, 1950 m, X.1979, W. Büttiker (NHMB); 6 ex., Thanomah, 1950 m, IV.1980, W. Büt- tiker (1 CMe, 5 NHMB); | ex., Riyadh, 24.40N/46.43E, V.1980, W. Büttiker (NHMB); 2 ex., An Nimas, 19.07N/42.08E, VIL.1981 (CMe); | ex., Jizan, 16.57N/42.33E, 111.1984, A. S. Talhouk (NHMB); | ex., Bani Rizam, 18.20N/42.28E, 2230 m, 1X.1984, W. Büttiker (CMe); | ex., Harithi, 21.18N/40.18E, IV.1985, W. Büttiker (NHMB). — Somalia: | ex. Br. Somaliland, 10.00N/48.00E, B.M. 1929-398, W. A. Mactadyen (BMNH); 1 ex., Burao, Br. Somaliland, 9.00N/46.00E, V11.1942, J. R. Audy (BMNH); 3 ex., Run, Garee, VIII.1964, Miss. Biol. (NHMB): 5 ex., Run, 8.47N/48.56E, VII.1969 (NHMB): 1 ex., Taleh, 9.09N/48.26E, IV.1980 (NHMB). — Yemen: 2 ex., Jebel Jihaf, Wadi Leje, 13.45N/44.42E, 6300-6700 ft, X.1937, Scott & Brit- ton (BMNH); | ex., Wadi Natid, Kirsh, 13.22N/44.32E, 2300 ft. X11.1937, Scott & Britton (BMNH). Monolepta vincta Gerstaecker, 1871: 83 Monolepta alternata Chapuis, 1879: 23: syn. n. = Monolepta insignis Weise 1903: 212: syn. n. = Monolepta sjóstedti Weise, 1909: 212: syn. n. = Monolepta ugandaensis Laboissiere, 1920a: 52: syn. n. Monolepta lusingensis Laboissiere, 1920b: 98: syn. n. Monolepta bouvieri Laboissiere, 1920b: 98: syn. n. Monolepta striola Laboissiere, 1920b: 98-99: syn .n. 264 Bonner zoologische Beitráge 53 (2004) 16a b C Figs 12-16: Monolepta lepida Reiche, 1858 12: colour pattern; 13: basal antennal articles (a: Ö, b: Y); 14: two different sper- mathecae; 15: bursa-sclerites (a: dorsal, b: ventral); 16: median lobe (a: lateral, b: dorsal, c: ventral, without endophallic struc- tures). = Monolepta consociata Laboissiere, 1920b: 99; syn. n. = Monolepta rugifrons Laboissiere, 1920b: 99; syn. n. = Monolepta femoralis Laboissiere 1940b: 66; syn. n. Redescription. Total length. 3.25—4.75 mm (mean: 4.06 mm; n = 25). Head. An entirely yellowish red to red head is typical for specimens from lowland areas of eastern and west- ern Africa (Figs 18b, g), in montane regions of East Af- rica and along the Albertine Rift head often with yellow frons and black vertex (Figs 18a, d, e), frons can be also brownish (Fig. 18f) or head black throughout as in some type specimens of M. bouvieri and M. sjóstedti from Mt. Kilimandjaro, of M. femoralis from the Ruwenzori Mountains, and in most specimens from the Ethiopian Highlands including the type of M. alternata (Fig. 18c). Antennae pale yellow, only last antennomere or at most apical parts of the tenth antennomere brownish or black, antennomeres four to eleven about 4 times longer than broad at apex. Basal antennomeres comparatively short (Fig. 19), length of antennomeres two to three 0.86—1.00 (mean: 0.95), length of antennomeres three to four 0.28-0.37 (mean: 0.32). Thorax. Prothorax pale yellow in about 60 % of spe- cimens examined (Figs 18 a-d), others with yellowish- red to red prothorax (Figs 18e, g), very rarely proster- num black as in some types of M. alternata or prothorax Thomas WAGNER: Revision of the vincta Species-group of Monolepta Chevrolat 265 , ” q , e. at Fig. 17: Distribution of Monolepta lepida Reiche, 1858, and M. naumanni sp. n. black throughout (Fig. 18f) as in some specimens from Uganda and Rwanda. Pronotal width 0.95-1.35 mm (mean: 1.17 mm), pronotum broad, pronotal length to width 0.58—0.64 (mean: 0.61), very finely punctured. Elytra pale yellow, to yellow, extension of black parts highly variable. Mostly reduced in specimens from montane regions of East and Central Africa, with nar- row black elytral base (Fig. 18a), which is sometimes significantly extended along the outer elytral margins and the suture (Fig. 18d), subapical elytral band can be reduced to circular spots (types of M. sjóstedti: Fig. 18a), or subapical black spots can be extended towards the outer elytral margins (types of M. insignis and M. lusingerensis; Fig. 18d). Most specimens from mid alti- tude areas and lowland sites in West, Central and East M. naumanni sp. n. 7 E ee A 4-8 A 9-14 M. lepida o 1-2 O 3-8 (e) 9-23 Africa with more extended basal band (type of M. vincta; Fig. 18b, or types of M. consociata, M. rugifrons and M. striola; Fig. 18e). The black coloration is most extended in some specimens from the Ethiopian High- lands, and in few specimens from the Congo Basin and western Africa (Fig. 182). Elytral length 2.50-3.60 mm (mean: 3.09 mm), maximal width of both elytra 1.70 2.35 mm (mean: 2.12 mm). maximal width of both elytra to length of elytron 0.68-0.75 (mean: 0.72). Scutellum yellow, yellowish-red or black. Meso- and metathorax, and legs usually yellowish-red, with excep- tion of the pale yellow tibiae and tarsi. Specimens with black head from montane regions sometimes with brownish or rarely black meso- and metathorax and basal two thirds of femora can be also black, rarely un- onner zoologische Beitráge 53 (2004) Thomas WAGNER: Revision of the vincta Species-group of Monolepta Chevrola ( 23a b C 24a b C 25a b Figs 18-25: Monolepta vincta Gerstaecker, 1871 18: colour pattern; examples based mainly on coloration of type specimens a: M. sjóstedti, b: M. vincta, c: M. alternata, M. bouvieri, M. femoralis: d: M. insignis, M. lusingerensis; e: M. rugifrons, M striola, M. consociata; f: rare type with black pronotum from Rwanda, Uganda: g: one coloration type from Ethiopian Highlands: 19: basal antennal articles (a: $, b: Y); 20: three different spermathecae; 21: bursa-sclerites (a: dorsal, b: ventral); 22: bursa- sclerites, variation (a: dorsal, b: ventral); 23: median lobe, typical shape from Central and southern-east Africa (a: lateral. b: dor- sal, c: dorsal without endophallic structures); 24: median lobe, variation, mainly Ethiopian Highlands (a: lateral, b: dorsal, c: \ tral, without endophallic structures); 25: median lobe, very rare variation (a: lateral, b: dorsal, c: ventral, without endopha structures). 268 Bonner zoologische Beitráge 53 (2004) derside reddish and only femora with exception of the apex black like in some types of M. insignis. Abdomen. Pale yellow to yellow, in some specimens from montane regions with dark underside abdomen en- tirely or last sternites black. Female genitalia. Spermatheca with spherical nodulus, slender middle part and long, slender cornu (Fig.20). Dorsal part of bursa sclerites triangular, with strongly hooked base (Figs 21a, 22a), ventral part slender, outer margin finely serrate (Figs 21b, 22b). Male genitalia. Median lobe conical, straight, apical part nearly parallel-sided, slightly ventrally bent (Figs 23a, 24a), few specimens from the Congo Basin with very slender apical part of median lobe (Fig.25), some specimens from the Ethiopian Highlands have broader median lobes (Figs 24b, c). Tectum, pointed, ventral groove slender, slightly conical towards orifice (Figs 23b, c, 24b, c). Dorsal pair of median endophallic spicu- lae strong, curved or hooked at apex (Figs 23a, 24a, 25a), other median spiculae slender, long; ventral spicu- lae separated into two portions; two pairs of short lateral spiculae, the more basal pair slender, the more apical one triangular (Figs 23b, 24b). Distribution. This species is widely distributed and abundant in most parts of tropical Africa, but with in- creasing rarity in southern Africa towards the South, and not known from the Republic of South Africa (Fig. 26). Diagnosis. Monolepta vincta is most similar and very closely related to M. melanogaster and M. lepida. The latter two species are on average larger, and thus speci- mens smaller than 3.8 mm total length belong to M. vincta. However, there is a high overlap in body size. Monolepta lepida, which is sympatric with M. vincta in Somalia and Eritrea, can be distinguished by the elon- gated second and third antennomeres (length of second to third antennomeres: 0.75-0.88, M. vincta: 0.86-1.00; length of third to fourth antennomeres: 0.46-0.54, M. vincta: 0.27-0.35) and the narrow pronotum (pronotal length to width: 0.63-0.67, M. vincta: 0.57-0.64). Monolepta melanogaster 1s allopatric with M. vincta in South Africa and Namibia, while but both species are sympatric in Malawi, Congo, and most regions of Zim- babwe, southern Zambia, southern Mocambique and Uganda. Monolepta vincta can be distinguished on av- erage by more slender elytra, shorter antennae, and by slight differences in shape of the median lobe and endo- phallic armature (for details see diagnosis for M. melanogaster). Monolepta vincta, M. melanogaster, and M. lepida are most likely a monophyletic group, which can be derived from external characters, coloration and male genitalic patterns. Identification of M. lepida and especially of M. melanogaster from areas where these species occur sympatrically with M. vincta should be confirmed by examination of the male genitalia, while females can sometimes not be allocated to species with- out doubt. Other species similar to M. vincta are the herein de- scribed M. sharonae sp. n., which has the same colour pattern as the type of M. vincta (Fig. 18e). Both species can be clearly distinguished by examination of the male genitalia only, which are in particular different in endo- phallic armature (Figs 23, 24, 37). Monolepta sharonae sp. n. is on average smaller (total length up to 4.2 mm, M. vincta up to 4.7 mm), the basal antennomeres are longer (antennomere 3 to 4: 0.43-0.50, M. vincta: 0.28— 0.37), and the transverse post antennal suture is deeply incised in all known specimens of M. sharonae Sp. n., while this character is very rarely found in M. vincta. Less similar are M. buquetii which is also coloured like the type of M. vincta (Fig. 18e), but is on average smaller, has shorter antennae, and very different female and male genital characters (Figs 8, 9, 11). Monolepta ronbeeneni sp. n. is also much smaller than M. vincta, has longer antennae and more slender basal antenno- meres, a more yellowish-red elytral coloration and dif- fers also strongly in genital patterns (Fig. 31). Discussion. Wide distribution, high abundance and high polymorphism in coloration have resulted in a high number of synonyms. After this species was first de- scribed from the coastal region in Kenya, specimens with abberative coloration, in particular from Mt. Kili- mandjaro, Mt. Kenya and the Albertine Rift, were used for the description of additional species, which have turned out as synonyms. Apart from coloration, also male genital characters show some variation. Shape of the median lobe, endophallic armature, and coloration pattern varies independently and cannot be correlated to geographic distribution of single populations. It is very likely that some populations, for example some of the Ethiopian Highlands, are genetically more isolated and undergo a current speciation process. However, using morphological data only, these populations cannot be clearly distinguished from others. Molecular data might be helpful for species identity and evolutionary proc- esses of these beetles in the future. On the base of the available morphological data, all type specimens listed are considered as conspecific with M. vincta. Type material. Monolepta vincta Gerstaecker, 1871. Holotype: Y “56683 / Type / vincta, Gerst.*, Mombas, v. d. Deck.” (ZMHB); examined (coloration like Fig. 18b); type locality: Kenya, Mombasa, 4.04S/39.40E. GERSTAECKER gave no information on the number of specimens studied, and mentioned only one locality. There is only one specimen available from this site, which is considered as holotype. Thomas WAGNER: Revision of the vincta Species-group of Monolepta Chevrolat 269 M. vincta 1-7 8 - 30 31-70 (lia 158 Fig. 26: Distribution of Monolepta vincta Gerstaecker, 1871 Monolepta alternata Chapuis, 1879. Holotype: y “Abyss. Raffray / 519 / Monolepta alternata Chp. / Re- gione boschiva da Goundet ad Adoua, 1000-2000 m 1893 / Typus Monolepta alternata Chapuis, 1879 / Museo Civico di Genova / Holotypus Monolepta alter- nata Chapuis, 1879 / Monolepta vincta Gerstaecker, 1871 Th. Wagner det. 2000” (MCSN); examined (col- oration like Fig. 18c); type locality: Ethiopia, province Goundet a Adoua. CHAPUIS gave no information on the number of specimens studied, and mentioned only one locality. There is only one specimen available from this site among the material in Chapuis’ collection. This specimen is considered as holotype. Monolepta insignis Weise 1903. Lectotype: 4 “Mombo 7.99 / ex. coll. J. Weise / Type / Monolepta insignis m. Lectotypus Monolepta insignis Weise, 1903 Th. Wagner desig. 2003 / Monolepta vincta Gerstaecker, 1871 Th. Wagner det. 2003” (ZMHB); this designation; examined genwald (coloration like Figs 18a, d); type locality: Tanzania. Mombo, 4.54S/38.18E. — Paralectotypes: 12 ex. “Mom- bo” or “Mombo 7.99 / ex. coll. J. Weise”, 6 ex. addi- tionally “Type”, 1 ex. additionally “Monolepta insignis m.” (ZMHB); 2 ex. same data (NHRS). " *Kili- Typus Monolepta sjöstedti Weise, 1909. Lectotype: mandj. Sjöstedt / Kibonoto, Kulturz[one] / Type / 18. Dec./ sjöstedti m / Lectotypus Monolepta sjöstedti Weise, 1909 Th. Wagner desig. 2003 / Monolepta Gerstaecker, 1871 Th. Wagner det. 2003” (NHRS): this designation; examined (coloration like Figs 18a, b, d); type locality: Tanzania, Mt. Kilimand- Paralectotypes: | ex. “Meru Re- Meru Sjóstedt” (NHRS); 6 ex. “Kilimandjaro Sjöstedt” including 4 ex. additionally “Kibonoto, Kul- turz[one]” including 1 ex. “Type”, 2 ex. additionally “Kibonoto Nieder[ung]” (NHRS). the paralectotypes bear additional labels indicating day and month of col- vincta 270 Bonner zoologische Beitráge 53 (2004) lection. WEISE mentioned eight specimens from three locations in his original publication, which could be all located in NHRS. Monolepta ugandaensis Laboissiere, 1920a. Holotype: “ “Museum Paris Ouganda Ounyoro Méridional Ch. Alluaud 1909 / Janvier / Monolepta ougandensis {sic!| m. V. Laboissiere — Dét. 1919 / Holotypus / Monolepta vincta Gerstaecker, 1871 Th. Wagner det. 2003“. (MNHN), examined (coloration type as Fig. 18g). LABOISSIERE cited only data of one specimen “Muséum de Paris”, holotype by monotypy after article 73.1.2, ICZN rules. Monolepta lusingensis Laboissiére, 1920b. Lectotype: 2 “Oct. / Museum Paris, Afrique Orient. Angl., Ile de Lus- inga, Victoria-Nyanza, N.-E. Ch. Alluaud 1904 / Type / Monolepta lusingensis m. V. Laboissiere — Det. / Lecto- typus Monolepta lusingensis Laboissiere, 1920 Th. Wagner desig. 2003 / Monolepta vincta Gerstaecker, 1871 Th. Wagner det. 2003” (MNHN ); this designation; type locality: Lusinga Island, location? — Paralecto- types: | Y same data as lectotype (MNHN); | Y “Mu- seum Paris, Afrique Orient. Angl., Ile de Lusinga, Vic- toria-Nyanza, N. E., Ch. Alluaud 1904 / Oct. / Type / Monolepta lusinga{sic! Jensis” (ZMUH); examined (col- oration type as Fig. 18d). LABOISSIERE mentioned at least two specimens “Muséum Paris et coll. Labois- sıere” from one locality in his original publication. Monolepta bouvieri Laboissiere, 1920b. Holotype: Y “Museum Paris, Afrique Orient. All., Kilimandjaro, Zone des Foréts, Kiboscho, 1700 m, Ch. Alluaud 1904 / mars / Type / Monolepta bouvieri Laboi. V. Laboissiere — Det. / Holotypus / Monolepta vincta Gerstaecker, 1871 Th. Wagner det. 2000” (MNHN); examined (col- oration similar to Fig. 18d); type locality: Tanzania, Mt. Kilimandjaro, 3.048/37.22E. LABOISSIERE mentioned “un individu, Muséum de Paris”, holotype by monotypy after article 73.1.2, ICZN rules. Monolepta striola Laboissiere, 1920b. Lectotype: Y “Museum Paris, Ouganda Central, Ch. Alluaud 1909 / décembre / Type / Monolepta striola Labo1. V. Labois- siere — Det. / Lectotypus Monolepta striola Laboissiere, 1920 Th. Wagner desig. 2003 / Monolepta vincta Ger- staecker, 1871 Th. Wagner det. 2003” (MNHN); this designation; examined (coloration similar to Fig. 18e); type locality: Central Uganda. — Paralectotype: 1 3, same data as lectotype. LABOISSIERE mentioned at least two specimens “Muséum Paris et coll. Laboissiere” from one locality in his original publication. Monolepta consociata Laboissiére, 1920b. Holotype: $, “Afrique or. Anglaise, Mt. Kénya vers ouest, Zone in- férieure, Alluaud & Jeannel / Riviére Amboni, Vallée boisée, 1800m, Janv.-Fév. 1912- St. Et 51 / Type / Mu- seum Paris, coll. Générale / Monolepta consociata Lab. V. Laboissiere — Det. / Holotypus / Monolepta vincta Gerstaecker, 1871 Th. Wagner det. 2003” (MNHN ); ex- amined (coloration like Fig. 18e); type locality: Kenya, Mt. Kenya, 0.09S/37.19E. LABOISSIERE mentioned “un exemplaire, Muséum de Paris”, holotype by monotypy after article 73.1.2, ICZN rules. A Monolepta rugifrons Laboissiere, 1920b. Holotype: 4 “Museum Paris, Afrique Orient. Angl., Kénia N.-O., Prairies 2000 m, Ch. Alluaud 1909 / novembre / Type / Monolepta rugifrons Laboi. V. Laboissiere — Det. / Holotypus / Monolepta vincta Gerstaecker, 1871 Th. Wagner det. 2003” (MNHN); examined (coloration like Fig.18e); type locality: Kenya, Mt. Kenya, 0.098/37.19E. Labolssiére mentioned “un Í, Muséum de Paris”, holotype by monotypy after article 73.1.2, ICZN rules. Monolepta femoralis Laboissiere 1940. Lectotype: Y “Holotype femoralis [invalid indication] / Congo Belge: P.N.A., Rwindi, 1000m, 22. au 24.-xi-1934, G. F. de Witte: 792 / Lectotypus Monolepta femoralis Labois- siere, 1940 Th. Wagner desig. 2003 / Monolepta vincta Gerstaecker, 1871 Th. Wagner det. 2003” (MRAC); this designation; examined (coloration like Fig. 18c); type locality: Congo, Kivu, Rwindi, 0.47S/29.17E. — Paralec- totypes: 7 ex. “Paratype femoralis” / Congo Belge: P.N.A., Rwindi, 1000m, 22. au 24.-xi-1934, G. F. de Witte: [several numbers)” (1 ex. IRSNB, 6 ex. MRAC); 3 ex. “Paratype femoralis” / Congo Belge: P.N.A., Nde- ko (Pres Rwindi), 1082m, 27-x1-1934, G. F. de Witte: 839” (MRAC); 1 ex. “Paratype femoralis” / Congo Belge: P.N.A., Entre Kalinga et Bitshumbi, 1082m- 925m, 12-xi-1934, G. F. de Witte: 741” (MRAC). LABOISSIERE mentioned 17 specimens from the four lo- cations in his original publication of which were 12 specimens available. Further material examined. Angola: 1 ex., Dundo, 12.35S/15.12E, VI11.1953, E. Luna de Carvalho (MRAC); l ex., 5 mls SW Cacula, 12.22S/15.34E, 111.1972, Southern African Exp. B.M. 1972-1, at light (BMNH); 8 ex., Salazar, 9.18S/14.55E, 11.1972, Southern African Exp. B.M. 1972-1, at light (BMNH); 3 ex., Bruco, 15.07S/13.11E, 111.1972, Southern African Exp. B.M. 1972-1, damp leaf litter by stream (BMNH); 1 ex., 12 mls SW Luimbale, 12.15S/15.19E, 111.1972, Southern African Exp. B.M. 1972-1, at light (BMNH); 1 ex., Dolondola, 13.498/13.07E, X1.1974 (NMNW). — Benin: 1 ex., Niaouli, mais station, 6.45N/2.08E, V1.1952 (MNHN); 3 ex., Zanbbara, 10.28N/3.32E, IX.1969, G. Pierrard (MRAC). — Burkina Faso: 1 ex., Bobo- Dioulasso, 11.11N/4.08W, X.1964, R. Siffointe (MRAC); 4 ex., Quagadougou, 12.22N/1.31W, X1.1973, R. Linnavuori (MZHF). — Burundi: 2 ex., Bururi, 3.48S/29.35E, ex coll. Breuning (MRAC); 1 ex., Urundi, 11.1897, Ramsay & Hösemann (ZMHB); l ex., Mahembe, terr. Nyanza, 1400 m, 1.1953, P. Basilewsky (MRAC); 1 ex., Bururi, 11.1959, P. Giraudin (MNHN); 2 ex., Mugera, 3.048/30.40E, 1965, J. J. Rwabuneza (MRAC); 6 ex., Bururi, 1800-2000 , I1IL1953, P. Basilewsky (3 BMNH, 3 MRAC); | ex., Bujumbura, 3.228/29.31E, IX.1969, P. Giraudin, ex coll. Breuning (MRAC); 2 ex., Ngozi, 2.52S/29.56E, 11.1979, Thomas WAGNER: Revision of the vincta Species-group of Monolepta Chevrolat 27 J. Decelle (MRAC); 3 ex., Gitega, 3.29S/29.57E, 11.1979, J. De- celle (MRAC). Cameroon: 2 ex., Kamerun, Tessmann (ZMHB); 2 ex., Victoria, 4.02S/9.21E (MNHN, HNHM); I ex., Joko, 5.29N/12.19E (ZMHB); | ex., Ndzida Forest, V.1952 (MNHN); | ex., Joko, 1.1957, J. Cantaloube (MRAC); 3 ex., N’Kongsamba, 4.59N/9.53E, V.1957, J. Cantaloube (MNHN); 2 ex., Bas Logone, Birni, 11.47N/15.05E, VII1.1963, J. Péricart (MNHN); | ex., N’Kolbisson, 3.25N/11.00E, V1.1966, B. de Miré (MNHN); | ex., Plateau de Kounden, 5.42N/10.40E, VI11.1967, L. Matile (MNHN); | ex., Haute Nguemba, V111.1967, L. Matile (MNHN); | ex., Bafoussam, 5.31N/10.25E, 11.1969, N. Berti (MNHN); 5 ex., Logone-Ufer, 65 km S. F. Foureau, 11.30N/ 14.56E, V11.1978, Salix (CMa). — Central African Republic: | ex., Pays Mandjia, 5.46N/19.43E, Miss. Chari-Tchad, 1904, J. Decourse (MNHN); | ex., Bessou, amont Fort de Possel, 4.41N/ 17.33E, Miss. Chari-Tchad, V1.1904, J. Decourse (MNHN); 2 ex., Fort de Possel, 5.01N/19.15E, Miss. Chari-Tchad, IX.1904, J. De- course (MNHN); 2 ex., Babua, Obersanga, 5.48N/14.49E, 11.1914, Tessmann (ZMHB); 6 ex., Bosum, Uam-Gebiet, 6.19N/16.38E, 11.1914, Tessmann (ZMHB); 2 ex., Bambari, 4.13N/17.35E, 111.1964, G. Pierrad (MRAC). — Chad: 5 ex., Gory-Damraou, 10.06N/17.35E, Miss. Chari-Tchad, VI.1904, J. Decourse (MNHN); 3 ex., Fort Archambault, 9.08N/18.22E, Bahr el Azreg, Miss. Chari-Tchad, V.1904, J. Decorse (MNHN); | ex., N’Gouri, Kanem distr., 15.38N/15.22E, VIIL1958, P. Renaud (MRAC); | ex., Bas Chari, env. de Fort Foureau, 12.05N/14.56E, VI.1963, J. Pericart (MHNH); | ex., Bas Chari, env. de Fort Lamy, Farcha, 12.10N/14.59E, V11.1967, J. Péricart (MNHN); 9 ex., Bebedja, 8.41N/16.34E, V.1973, R. Linnavuori (MZHF); | ex., Farcha, 12.27N/15.13E, V.1973, R. Linnavuori (MZHF); | ex., Lamto, 13.24N/15.01E, V.1990, H. Perrin (MNHN). — Congo-Brazza- ville: 2 ex., Fort-Crampel, 7.00N/19.10E (ZMUH). — Congo: | ex. (NHRS); | ex., Dybowski (MNHN); | ex. Tavares (ZMHB); 3 ex., Chiloango, 5.12S/12.07E, Tschoffen (2 ex. IRSNB, 1 ex. MRAC); 3 ex., Matadi, 5.59S/13.27E Tschoffen (IRSNB); | ex., Ituri, La Moto, Madyu, 2.54N/29.37E, L. Burgeon (MRAC); 7 ex., Banana Boma, 6.01S/12.24E, 1891, M. Tschoffen, Jacoby coll., pulchella Klug (1 ex. BMNH, 4 ex. IRSNB, 2 ex. MRAC); | ex., SW Albert-Njansa, Undussuma, 1050 m, VII.1891, Stuhl- mann (ZMHB); | ex., SW Albert-Njansa, Buessa, VIII.1891, Stuhlmann (ZMHB); 4 ex., Dima, 3.29N/18.42E, IX.1908, A. Koller (MRAC); | ex., Matadi, , IX.1910, Dr. Bequaert (MRAC); 45 ex., Congo da Lemba, 5.42S/13.42E, V., X.—XIL1911, V., X.1912, 1.-111.1913, R. Mayné (MRAC); 4 ex., Elisabethville, 11.40S/27.28E, X.1911, Miss. Agric. (MRAC); 3 ex., Kapiri, 9.428/27.13E, X.1912, Miss. Agric. (MRAC); 1 ex., Banza, Manteka, 5.28S/13.47E, VI.1912, R. Mayné (MRAC); 2 ex., A- madi, 3.35N/26.47E, 11.1913, P. van den Plas (MRAC) | ex., Kiniati-Zobe, 5.07S/12.37E, X11.1915, R. Mayne (MRAC): | ex., Mayumba, Zobe, 5.07S/12.37E, 1.1916, R. Mayné (MRAC); 2 ex., Mayumbe, Lemba, 4.45S/14.15E, V11.1917, R. Mayne (MRAC): 8 ex., Nyangwe, 4.135/26.11E, ML-IV.1918, R. Mayne (MRAC): 1 ex., Albertville, 5.56S/29.12E, XI1.1918, R. Mayné (MRAC); | ex., Madyu, 1919, L. Burgeon (MNHN); 1 ex., Kisantu, 5.07S/15.05E, V,1919, P. Vanderijst (MRAC); 1 ex., Boma, VII.1920, H. Schouteden (MRAC); 3 ex., Haut-Uele, Watsa, 3.03N/29.32E, XI.1919, 1922, L. Burgeon (MRAC); 1 ex., Dungu, 3.37S/28.34E, 1V.1920, P. van den Plas (MRAC); 2 ex., Matadi, 5.49S/13.27E, X.1920, L. Burgeon (MRAC); 2 ex., Elisa- bethville, R. Lubumbashi, X1.1920, M. Bequaert (MRAC); 4 ex., Haut-Uele, Moto, 2.27N/26.25E, 1922, 1923, 1.1925, L. Burgeon (MRAC); | ex., Kwamouth, 3.10S/16.12E, 1.1922, H. Schouteden (MRAC); 5 ex., Haut-Uele, Abimva, 3.09N/29.50E, V.-V11.1925, L. Burgeon (MRAC); | ex., Katanga, Nyonga, V.1925, G. F. de Witte (MRAC); 4 ex., Haut-Uele, Yebo Moto, 1.-I1., 1X.1926, L. 271 Burgeon (MRAC); | ex., Kivu, Mokoto, 1.155/29.00E, 1927, G Babault (MNHN); 2 ex., 18 m SW of Elisabethville, 11,425 27.25E, 1927, Dr. H S. Evans (BMNH); 2 ex., alto Uelle, fl. Duru 3.45N/28.02E, 111.1927, F. S. Pratizi (MCSN); 3 cx., alto Uclle Yakuluku, 4.20N/28.48E, 1V.1927, F. S. Patrizi (MCSN); I ex alto Uelle, Kapili, 3.42S/27.53E, V.1927, F. S. Patrizi (MCSN); | ex., Uvira, 3.24S/29.08E, 1930, G. Babault (MNHN); | ex.,¿Lo- mami, Kambaye, 6.535/23.44, VI11.1930, P, Quarré (MRAC); | ex., Lulua, Sandoa, 9.41S/22.52E, X.1930, G. F. Overlact (MRAC); 18 ex., Kivu, Rwindi, 0.475/29.17E, 1000 m, X1.1931, X1.1934, G. F. de Witte (IRSNB); 1 ex., Ubangı, Dongo, 2.43N/18.24E, XII.1931, H. J. Bredo (MRAC); 1 ex., Lulua, Tshala, 7.56S/18.08E, V11.1932, G. F. Overlaet (MRAC); 3 ex., Lomami, Kaniama, 00.46N/24.16E, VII.1932, R. Massart (MRAC); 1 ex., Ruwenzori, Kalonge, 0.20N/29.48E, 2050 m, VIII.1932, L. Burgeon (MRAC); 1 ex., Beni, 0.30N/29.28E, V111.1932, L. Burgeon (MRAC); | ex., Kivu, Lulenga, 1.255 29.23E, IX.1932, L. Burgeon (MRAC); 3 ex., Kivu, Ngoma, 2.048/29.55E, X.1932, L. Burgeon (MRAC); | ex., Kivu, Katana, X1.1932, L. Burgeon (MRAC); 3 ex., Lulua, Kapanga, 8.215 22.35E, X11.1932, 111.1933, F. G. Overlaet (MRAC); | ex., Lu- vungi, 2.52S/29.02E, X11.1932, L. Burgeon (MRAC); | ex., Al- bertville, 1.1933, 1. Burgeon (MRAC); 38 ex., Kivu, Rutshuru, 1.115/29.27E, 1285 m, X11.1933, V1.1934, VI.-VII.1935, G. F. de Witte (25 ex. IRSNB, 12 ex. MRAC); 3 ex., Kilo, Dele, 1.49N/30.16E, 1V.1934, H. J. Brédo (MRAC); 2 ex., Ruhengeri, Source Kirii, 1.27S/19.00E, 1800 m, X.1934, G. F. de Witte (IRSNB, MRAC); 24 ex., P.N. Albert, Ndeko, 1.04N/20.04E, 1082 m, X1.1934, G. F. de Witte (12 ex. IRSNB, 12 ex. MRAC): 4 ex., P.N. Albert, May ya Moto, 0.535/29.21E, 950 m, X1.1934, G. F. de Witte (IRSNB); 2 ex., P.N. Albert, Camp Rwindi, 0.48S/29.18E, 1000 m, X1.1934, G. F. de Witte (MRAC); 2 ex.. P.N. Albert, Escarpment de Kabasha, 0.45S/29.13E, 1500 m, X11.1934, G. F. de Witte (IRSNB); 2 ex., Bunia, 1.34N/30.15E, 1V.1935, H. J. Brédo (MRAC); 4 ex., P.N. Albert, Nyasheke, Vole. Nyamuragira, 1.23S/29.19E, 1820 m, VI.1935, G. F. de Witte (IRSNB); 2 ex., P.N. Albert, Mayumbu, Vole. Nya- muragira, 1.25S/29.12E, 2100 m, VI.1935, G. F. de Witte (IRSNB); 2 ex., P.N. Albert, R. Ondo, affl. Rutshuru, VII.1935, H. Damas (MRAC); 3 ex., Kasenyi, 7.265/24.10E, VIL, 1X.1935, VI11.1937, H. J. Bredo (MRAC); 3 ex., Eala, 0.04N/18.17E, 1V.1936, J. Ghesquiere (MRAC): 1 ex., P.N. Albert, Bitshumbi, 0.39S/29.22E, IV.1936, L. Lippens (IRSNB); 10 ex., P.N. Albert, r. Rwindi, H.-1V.1936, L. Lippens (MRAC); 61 ex.. Rutshuru, V.1936, 1.-V.1937, L. Lippens / Ghesquiere (MRAC): 21 ex.. Bambesa, 3.28N/25.43E, V.1937, J. Vrijdagh (IRSNB): 1 ex.. Djugu, 1.55N/30.30E, V.1937, H. J. Bredo (MRAC); 11 ex.. Ni- oka, 7.078/26.37E,V11.1937, H. J. Brédo J. Ghesquiere (MRAC); 3 ex., Ituri, Bunia, 1.34N/30.15E, VII.1937, H. J. Brédo (1 ex. IRSNB, 2 ex. MRAC); I ex, Lac Albert, Mahagi, 2.18N 30.59E, VI1.1937, J. Ghesquiere (MRAC): 3 ex., Logo, 2.11N 30.56E, VII.1937, H. J. Brédo (MRAC): 1 ex., Lukolela, 1.035/17.12E, 1938, R. Massart (IRSNB); | ex., Ubangi, Budjala, 2.39N/19.42E, VI1.1938, C. Léontovitch (MRAC): | ex.. Elisa- bethville, 1.1939, H. J. Brédo (IRSNB): 1 ex., Mulungu. 4.54S/19.02E, 1939, Hendrickx (MRAC): 1 ex., Mayidi, 5.11S/15.09E, 1912, P. van Eyen (MRAC); 1 ex., Terr. Tan- ganika, Malonge, 10.24S/23.10E, V1.1943, H. J. Bredo (IRSNB): l ex., Terr. Tanganika, Katete, 10.545/27.54E, VII.1943, H. J. Brédo (IRSNB); 1 ex., P.N. Albert, Kiseguro, riv. Mwindi, IV.1945, G. F. de Witte (MRAC); 3 ex., P.N. Albert, Rumangabo, 1.20S/29.22E, IV.1945, G. F. de Witte (MRAC): 158 ex., P.N. Upemba, 8.30S/26.00E, several locations. VI.1945. IIL. X. X11.1947, 1.-VII., X.1948, 1.1949, G. F. de Witte (2 ex. IRSNB, 48 ex. MRAC); 3 ex., W. Ruwenzori, 0.23N/29.54E, 6000 ft., 272 Bonner zoologische Beitráge 53 (2004) V11.1945, v. Someren (NMK); 10 ex., Rwankwi, 1.32S/29.32E, IV.1946, IV.1948, X11.1947, VI, X1.1951, J. V. Leroy (MRAC); I ex., Bukima, IV.1948, J. V. Leroy (MRAC); 1 ex., Stanleyville, Yaolema, 1.54N/22.47E, IV.1949, R. Laurent (MRAC); 1 ex., P.N. Upemba, Lusinga, 8.56S/27.12E, 1760 m, V.1949, G. F. de Witte (IRSNB): 465 ex, P.N. Garamba, 3.40N/29.00E, several sub locations, X11.1949, L.—MI., V.,VIIL, X.-X1.1950, [.-XI1.1951, 1.- IML., VIL-V1.1952, Miss. H. de Saeger (65 ex. IRSNB, 400 ex. MRAC); 19 ex., Elisabethville, X1.1950/V1.1951, 1953, 1955, 1.1956, 1.1957, 1X.1958, V.1959, Ch. Seydel (MRAC); 4 ex., P.N. Albert, Bomboka, Kyandolire, 1650 m, X.1952, P. Vanschuyt- broeck & J. Kekenbosch (MRAC); 1 ex., Tanganika, Kamena, 7.258/28.46E, 1400 m, Riv. Kinga, 1.1953, H. Bomans (MRAC ); 10 ex., P.N. Albert, Kalonge, 1.18S/28.24E, 2060 m, 11.1953, P. Vanschuytbroeck & J. Kekenbosch (MRAC); 2 ex., P.N. Albert, Haut Lume, Nyamgaleka, 0.15N/29.48E, 2120 m, IV.1953, P. Vanschuytbroeck & J. Kekenbosch (MRAC); 1 ex., P.N. Albert, Moyenne Lume, Kiribata, 0.16N/29.48E, 1760 m, IV.1953, P. Vanschuytbroeck & J. Kekenbosch (MRAC); | ex., P.N. Albert, Migeri, riv. Kamukungu, 1700 m, IV.1953, P. Vanschuytbroeck & J. Kekenbosch (MRAC); 2 ex., Lubero, Mulo, 0.08S/29.16E, 1950 m, 11.1954, R. R. P. P. Celis-Bergmans (MRAC); 1 ex., Tanganika, Albertville, IV.-V.1955, H. Bomans (MRAC); | ex., Uvira, Itombwe, Hte. Kambekulu, 4.04S/28.40E, 2450 m, V1.1955, N. Leleup (MRAC); 1 ex., P.N. Albert, Mt. Musima, 2.195/29.17E, 2450 m, IV.1955, P. Vanschuytbroeck $ R. Fon- teyn (MRAC); 2 ex., P.N. Albert, Tschiaberimu, Mt. Buliwa, 2.048/28.55E, 2450 m, IV.1955, P. Vanschuytbroeck & R. Fon- teyn (MRAC); 14 ex., P.N. Albert, Mt. Hoyo, 1.13N/29.49E, 1280 m, V11.1955, P. Vanschuytbroeck (3 ex. IRSNB, 11 ex. MRAC); l ex., Kivu, Luvungi, 2.52S/29.02E, 111.1956, J. Decelle (MRAC); l ex., P.N. Albert, Kanyatsi, 0.355/29.18E, 912 m, VII.1956, P. Vanschuytbroeck & J. Kekenbosch (MRAC); 1 ex., P.N. Albert, Kasindi-Port, 2.20S/28.42E, 912 m, VII.1956, P. Vanschuyt- broeck (MRAC); 1 ex., P.N. Albert, Semliki, 0.085/29.36E, 912 m, VIIL 1956, P. Vanschuytbroeck (MRAC); | ex., P.N. Albert, Molidi, Watalinga, 0.40N/29.40E, 1210 m, IX.1956, P. Van- schuytbroeck (MRAC); 1 ex., P.N. Albert, Rugetsi, 0.13N/29.40E, 1100 m, X.1956, P. Vanschuytbroeck (MRAC); 2 ex., P.N. Al- bert, Ihumbia, 0.00/29.40E, 1050 m, X1.1956, VIII.1957, P. Van- schuytbroeck (MRAC); | ex., P.N. Albert, Téte de source Indray, 1840 m, X1.1956, P. Vanschuytbroeck (MRAC); 2 ex., P.N. Al- bert, Mati, Talya, 0.31N/29.20E, 1180 m, L, VII.1957, P. Van- schuytbroeck (MRAC); 1 ex., P.N. Albert, Mutsora, 0.19N/ 29.45E, 1340 m, 11.1957, P. Vanschuytbroeck (MRAC); 1 ex., P.N. Albert, Kanyatsi, Lac Edourd, 0.08S/29.37E, 912 m, V.1957, P. Vanschuytbroeck (MRAC) 1 ex., M’Paka, terr. Libenge, 4.46N/19.14E, VIL—VIII.1959, M. Pecheur (MRAC); 9 ex., Lualaba, Ruwe, 10.39S/25.30E, 11.1960, V. Allard (MRAC); 2 ex., Katanga, Kolwezi, 10.435/25.28E, X.1961, V. Allard (MRAC); | ex., Mongo, 0.02N/18.19E, 111.1975 (MZUF); 1 ex., Baraka, 4.06S/29.06E, VHI.1979, coll. Roggeman (CBe). — Eri- trea: | ex., Erythrée, coll. Clavareau (MRAC); 2 ex., Asmara, 15.20N/38.56E (BMNH, MNHN); 2 ex., Dint. Di Massaua, 15.30N/39.20E, 1884, Frasca (MCSN); 13 ex., Agordat, 15.33N/ 37.53E, 1.1906, D. Figini (MCSN); 10 ex., Agordat, 11.1930, E. Zavattarı (MCSN); 3 ex., Ghinda, 15.27N/39.03E, III.1906, D. Figini (MCSN); | ex., Decamere, 15.04N/39.03E, V.1963, R. Linnavuori (MZHF). — Ethiopia: | ex., Abyssinie, coll. Clavareau (MRAC); 3 ex., Abyss., Raffray, ex Mus. E. Allard (MNHN); 10 ex., Abyssinia, Vallis Erer, 9.10N/39.50E, Kovacs (HNHM); 1 ex., N. Dalla, 7.43N/40.55E, Jando (MNHN); 6 ex., Tigre, 1850, Schimper (MNHN); 1 ex., Abyssinie, Voy. A. Raffray, 1881 (MNHN); 1 ex., Arussi Galla, Ganale Gudda, 4.16N/42.01E, V.1893, V. Bottego (MCSN); 2 ex., Kashinuaha, Tshertsher, 1.1899, Dmitriev (ZISP); 1 ex., Buroma, Tshertsher, 1.1899, Dmitriev (ZISP); 1 ex., N. Galla, 11.30N/40.00E, X1.1900, v. Er- langer (ZMHB); | ex., Wallega, 9.00N/35.30E, VI.1904, Dmitriev (ZISP); 3 ex., Addis Abeba, 9.02N/38.42E, XII.1910, C. Citerni (MCSN); 3 ex., Between Addis Allem and Djem Djem, 9.02N/38.24E, 7000-8000 ft, IX.1926, H. Scott (BMNH); 1 ex., Plains NW of lake Zwai, 7.56N/38.43E, 5500-6000 ft, X.— X1.1926, H. Scott (BMNH); 5 ex., Mt. Chillälo, 7.55N/39.16E, 8500 ft, X1.1926, H. Scott (BMNH); 1 ex., Mt. Chillálo, 7000- 8000 ft, X1.1926, J. O. Cooper (BMNH); 5 ex., Addis Abeba, 1928, 1930, Schürhoff (ZMHB); 1 ex., Gondar, 12.36N/37.28E, 1932, Miss. M. Griaule (MNHN); 1 ex., Arero, 4.45N/38.49E, IV.1937, Miss. Zavattari nei Borana (Trieste); 1 ex., El Diee, V.1939, Miss. Zavattarı Sagan Omo (Trieste); 1 ex., Caschei nei Borana, 5.00N/39.00E, VII.1939, Miss. Zavattari (Trieste); 1 ex., V. Zuai, 8.00N/38.50E, 11.1954, Exped. Smeds (MZHF); 2 ex., Asella, 7.57N/39.08E, VIII.1954, Exped. Smeds (MZHF); 2 ex., Belleta Forest, 7.32N/36.31E, V1.1963, R. Linnavuori (MZHF); 1 ex., Shashamaanni, V1.1963, R. Linnavuori (MZHF); 4 ex., Addis Abeba, VII.1963, P. M. Schroeder (USNM); 1 ex., Shoa Prov., Wolisso, 8.32N/38.00E, VI.1971, R. O. S. Clarke (MRAC); 1 ex., Belleta Forest, 40 km SW Jimma, V.1971, R. O. S. Clarke (MRAC) 1 ex., Gibbie Riv., 175-185km SW Addis Abeba, V.1971, R.O.S. Clarke (MRAC); 1 ex., Gojeb river valley, 80 km SW Jimma, 7.20N/37.21E, 1650 m, VII.1971, R. O. S. Clarke (MRAC) | ex., Illubador Prov., 8 km N of Bedelle, 8.27N/ 36.20E, 1800 m, V.1972, R. O. S. Clarke (MRAC); 1 ex., Gemu Gofa Prov., Chencha, Gughé Mts., 6.12N/37.30E, 2800 m, IX.1972, R. O. S. Clarke (MRAC); 1 ex., Shoa, Awash Station, 6.58N/35.46E, IX.1973, G. de Rougemont (MRAC); 1 ex, Jimma, 7.40N/36.50E, 11.1974, H. Silfverberg (MZHF); 6 ex., Addis Abeba, 11.1974, H. Silfverberg (MZHF); 3 ex., Harage Alemaya, 9.23N/41.56E, 11.1974, H. Silfverberg (MZHF); 2 ex., Kaffa Shebe, 7.40N/36.35E, 11.1974, H. Silfverberg (MZHF); 4 ex., Arba Minch, 6.02N/37.33E, 111.1974, H. Silfverberg (MZHF); 6 ex., Lake Langano, 7.38N/38.42E, sweep-netted, X.1980, A. De- meter (HNHM); 7 ex., Shafartak Bridge, Blue Nile, Abbai Gorge, 10.05N/38.18E, sweep-netted, X.1980, A. Demeter (HNHM); 4 ex., Koka, 8.20N/38.40E, sweep-netted, X.1980, A. Demeter (HNHM); | ex., Sodere, 8.24N/39.22E, sweep-netted, X1.1980, A. Demeter (HNHM); 8 ex., Akaki river, 8.50N/38.43E, sweep- netted, X1.1980, A. Demeter (HNHM); | ex., Awash NP, 11.30N/ 41.40E, sweep-netted, X1.1980, A. Demeter (HNHM); 1 ex., env. Pastoria, 1X.1984, L. Medvedev (CMe); 2 ex., Gambella, 8.15N/34.35E, X1.1986, L. Medvedev (CMe); 1 ex., 30 km NW Gambella, X11.1986, L. Medvedev (CMe); 2 ex., pr. Mlubabur, 20 km E Abobo, 7.51N/34.40E, XII.1986, L. Medvedev (CMe): 7 ex., Addis-Abeba, 2700, VIII.1988, X.—X1.1990, L. Medvedev (CMe); | ex., Sabata, 2000 m, X11.1988, L. Medvedev (CMe); 2 ex., Shoa, Debre Zeyt, 10.35N/35.48E, V.1989, Werner, coll. Dóberl (CBe); 1 ex., Shewa, Rift Asella Valley, 7.57N/39.08E, MIL.-V.1989, Dedoch (CBz); 17 ex., Ambo, 11.15N/39.34E, X.1990, Acacia, L. Medvedev (CMe); 2 ex., Arsi, reg. Mondo genet, 7.30N/39.30E, 1850 m, X111.1990, Werner, coll. Dóberl (CBe); 2 ex., Shoa, Lake Langano, 1600 m, VI.1990, Werner (MZUF); 8 ex., Shewa, Debre Lib., 9.04N/38.05E, 2500 m, 111.1997, M. v. Tschirnhaus (ZFMK). — Gabon: | ex., Kankan, 0.02S/12.14E, VII1.1937, L. Berland (UNHN). — Gambia: 6 ex., Ht. Gambie, 13.15N/16.00W, IL—I 11.1901, E. Laglaize (MNHN); 3 ex., Kouroussa, 10.40N/9.52W, XI.1995, Lange (CBe). — Ghana: | ex., Gold Coast (BMNH). — Guinea: 2 ex., Dalaba, 10.47N/12.12W, XII.1910, A. Chevalier (MNHN ); 2 ex., Dalaba, 1200 m, VII.1945, H. Durand (MNHN); 4 ex., Seredou, 8.23N/9.23W, IV.1975, A. Zott, lux (ZMHB): 8 ex., Tabuna, 9.31N/12.26W, IX.1983, 111.1984, C. B. Myrzin (CMe); 4 ex., Thomas WAGNER: Revision of the vincta Species-group of Monolepta Chevrolat 273 Telimela, 11.1984, C. B. Myrzin (CMe); | ex., Kotento Yoko, 111.1984, C. B. Myrzin (CMe). — Guinea-Bissau: 6 ex., Bolama, 11.33N/15.37W, V1.-X11.1899, L. Fea (MCSN); 1 ex., Contubo, 12.04N/15.23W, 1.1938, coll. Laboissiére (MRAC); | ex., Missáo da Guiné, Aldeia de Cuor, 11.1946, Dr. E. Frade (IICT). — Ivory Coast: | ex., Ashante, 7.10N/1.40W, Baly coll. (BMNH); | ex., Dimbroko (MHNH); | ex., G. Melou (MNHN); 2 ex., Haut Sas- sandra, pays Youra, 4.50N/6.00W, V.1910, F. Fleury, A. Cheva- lier (MNHN); 3 ex., Haut Sassandra, pays Dyola, 4.40N/6.20W, 1910, F. Fleury, A. Chevalier (MNHN); 2 ex., Danané, 7.19N/8.06W, 1.1939, L. Chopard (MNHN); 4 ex., Adiopodoumé, IL, X., X11.1947, Ch. Primot (MNHN); | ex., Adiopodoumé, IX.1952, coll. P. Cachan (MNHN); | ex., Akandjé, 5.24N/3.54W, 1.1962, E. Lavabre (MNHN); 9 ex., Bingerville, 5.20N/3.52W, VI., IX-X.1962, 111.1964, J. Decelle (MRAC); | ex., Zepreghé, Daloa, 6.56N/6.28W, IX.1962, J. Decelle (MRAC); 6 ex., Koun- Abronso, 7.31N/3.15W, X1.1961, 1.1962, J. Decelle (MRAC); 2 ex., Nguessankro, Bougounanou, 7.57N/5.08W, X.1963, J. Decelle (MRAC); 1 ex., Ferkessedougou, 9.20N/5.00W, V.1964, J. De- celle (MRAC); 2 ex., Foro-Foro, 7.59N/5.03W, 1X.1973, R. Lin- navuori (MZHF); 2 ex., Man, X.1973, R. Linnavuori (MZHF); 16 ex., Comoe NP, 8.75N/3.80W, VI.-VI1.1997, C. Mody (ZFMK). — Kenya: 2 ex., Kenia, Varendorff, coll. Kraatz (DEI, ZMUH); | ex., Nairobi, Ngong Forest, 1.225/36.40E, 1900 m (IRSNB); 2 ex., Kibwezi, 2.25S/37.58E, X1.1887, G. Scheffler (ZMHB): 2 ex., Escarpment, 6500-9000 ft, IX.1900-IV.1901, W. Doherty (MNHN); 1 ex., Simba, 3350 ft, IV.1911, S. A. Neave (BMNH); 2 ex., Voi, 3.23S/38.35E, 11.1912, S. A. Neave (BMNH); | ex., Ramissi, X1.1915, Methner (ZMHB); | ex., Tana River, V1.1915, G. Babault (MNHN); 1 ex., Fort Hall, 0.45S8/37.00E, X11.1919, Patrizi (MCSN); 1 ex., Kisumu, 0.068/34.45E, 3756 ft, X1.1920, A. F. J. Gedye (BMNH); 1 ex., Nairobi, 1.17S/36.50E, X1.1920, A. F. J. Gedye (NMK); 3 ex., Chania Falls, Thika, 1.03S/37.05E, 5050ft., 1.1921, A. F. J. Gedye (NMK); 2 ex., Nairobi, 1.17S/ 36.50E, V.1921, V1.1937, A. F. J. Gedye (NMK); 1 ex., Kabete, 1.16S/36.43E, VIII.1921, H. E. Box (BMNH); | ex., Machakos, 2.00S/37.40E, X1.1923, J. W. Hunt (NMK); 1 ex., Ngong KC, 1.22S/36.40E, VII.1925, A. F. J. Gedye (NMK); 1 ex., Gasi, 4.258/39.30E, X1.1927, Dr. van Someren (NMK); | ex., Thika, 1.03S/37.05E, IV.1930, coffee (BMNH); | ex., Yala River pres Kisumu, 0.048/34.05E, X.1930, coll. G. Babault (MNHN); 2 ex., Kaimosi, 0.12N/34.57E, III.-IV.1932, A. Turner (NMK); 2 ex., Marakwet, Kigeyo Escarpment, 1.05N/35.37E, 1932-1933, Miss. de l’Omo, C. Arambourg et al. (MNHN); 3 ex., Elgon Saw mill, Mt. Elgon, ver est, 1.05N/34.40E, 2470 m, 1932-1933, Miss. de l’Omo, C. Arambourg et al. (MNHN); 1 ex., Thika Falls, Kukiyu, 1700 m, 1932-1933, Miss. de l'Omo, C. Arambourg et al. (MNHN); 1 ex., Suam fishing hut, Mt. Elgon vers est, 1.06N/34.33E, 2400 m, 1932-133, Miss. de l'Omo, C. Arambourg et al. (MNHN); | ex., Nairobi, 1660 m, 1932-1933, Miss. de l’Omo, C. Arambourg et al. (MNHN); | ex., Lower Tana, Sabaki, 3.098/40.07E, IV.-V.1932, Turner-Mac Arthur (NMK); 2 ex., Ki- sumu, IV.1936, H. J. A. Turner (NMK); 2 ex., Kanziko, 1.49S/38.15E, IX.1936, V.1937, Mac Arthur (NMK); | ex., Athi Falls, VII.1937, A. F. J. Gedye (NMK); 1 ex., Karakuta, 1.03S/36.57E, VIII.1937, R. E. Toker (NMK): 6 ex., Kitui, 1.22S/38.01E, X.1937, R. Toker (NMK); 2 ex., Lake Baringo, 0.38N/36.05E, 3400 ft, 1.1938, D. G. MacInnes (NMK); | ex., Stoneham, E. Surr. Estate, 11.1938 (NMK): 1 ex., Malakisi, 0.41N/34.25E, 1600 m, 11.1938, A. Holm (NHRS); 1 ex., Mt. El- gon, River Swam, 1.48N/35.09E, 2000 m, 11.1938, A. Holm (NHRS); 2 ex., Kaptega Estate, 1.11N/34.46E, 2050 m, IV.1938, A. Holm (NHRS); I ex., Thika Dist., 1.00S/37.20E, IV.1938, S. Blom Bjorner (NMK); 63 ex., Chyulu Hills, 2.355/37.50E, 5600 ft, IV.-VI1.1938, Coryndon Museum (62 ex. NMK, l ex. USNM): 3 ex., Thika, 1.045/37.02E, X.1937, X.1938, S. Blom Bjorner (| ex. BMNH, 2 ex. NMK); 2 ex., Kabete, 1.168/36.43E, 5800 ft, X11.1938, 1.1939, Mac Innes (BMNH, NMK); 2 ex., Malindi, 3.13S/40.07E, V.1940, G. W. Jeffery (BMNH, NMK); | ex., Ka- rura Forest, 1.30S/36.46E, 11.1941 (NMK); 2 ex., Nairobi, IV.1943, Meneghetti (NMK); | ex., Kaiti River, 1.455/37.42E, 11.1947, v. Someren (BMNH); | ex., Taita Hills, 3.255 38. 20E, V111.1947, van Someren (NMK); | ex., Mombasa, 4.04S/39.40E, IX.1947, T. H. E. Jackson (NMK); | ex., Mombasa, VIII.1948, A. F. J. Gedye (NMK); | ex., Kange-Ngomeni Road, 0.50S/38.25E, V.1951, sweep net attached to car 6 ft above ground (BMNH); 2 ex., Diani Beach, 4.18S/39.35E, VIL-VII.1951, N. L. H. Krauss (BMNH); | ex., Nairobi, 1.1953, E. Pinhey (NMK); | ex., Taveta, 15 km E, 3.23S/37.42E, VII.1968, N. Sanfilippo (MCSN); 8 ex., Nairobi, VIII.—X.1969, M. P. Cliften (NMK); | ex., Kisumu, X11.1970, A. E. Stubbs (BMNH); 1 ex., Tiwi Beach, 4.14S 39.35E, V.1971, B. D. Valentine coll. (BPBM): | ex., L. Na- ivasha, Crescent Island, 0.54S/36.37E, 6200 ft, 1.1972, C. F. Hug- gins (BMNH); 3 ex., Kora Rock, Tana River Exp., VII.1976, tilley lamp (NMK); 10 ex., Kaptega, 1980 m, 1.-11.1979, T. E. Leiler (NHRS); | ex., Naro Moru, 0.10S/37.01E, VIII.1979 (NNML); 4 ex., Stony Athi, 1.35S/37.00E, VI.1980, D. G. Furth (USNM); 5 ex., Langata Forest, 1.20S/36.47E, VI.1980, D. G. Furth (USNM); 2 ex., Nairobi, 1850 m, 111.1981, E. Räsänen, at light (MZHF); 3 ex., Kora NR, 0.38S/37.00E, Base Camp, VHI.1983, M. Clifton (NMK); 2 ex., Nairobi, 1X.1987, B. L. Parker (NMK); | ex., Kakamega Forest, 0.18N/34.53E, IV.1991, R. Copeland (NMK); 5 ex., Mt. Elgon NP, nr. Chepanyalli Cave, dry evergreen montane forest, 2500 m, 1.1992, O. Merkl & G. Varkonyi (HNHM); 2 ex., Arabuko Sokoke FR, 3.20S/39.52E, IX.-X.1992, L. Bartolozzi (MZUF); 1 ex., Taita Distr., Bura, 3.30S/38.18E, VI.1994, L. Bartolozzi, B. Cecchi, A. Sforzi (MZUF); 2 ex., Masai Mara, 1.25S/34.55E, 1600 m, VI.1994, L. Bartolozzi (MZUF); 1 ex., Ngong Hills, Nairobi Res. Stat., X1.1995, R. Copeland (NMK); 1 ex., Kitale, Kapenguria, 2300 m, XII.1995, M. Snizek (MIZT); 1 ex., Taita Hills, Rongo Forest, 3.21S/38.26E, X.—XI.1996, Malaise trap, ICIPE (NMK): 1 ex., Taita Hills, Wundanyi, X1I.1997, M. Snizek (MIZT): 1 ex., Nai- robi, Langata, 1.20S/36.46E, 1650 m, 1.1999, Th. Wagner, insec- ticidal fogging of Teclea simplicifolia (ZFMK); 1 ex., Taita Hills, n. Wundanyi, 3.24S/38.22E, 1660 m, I111.1999, Th. Wagner (ZFMK); 1 ex., Mwingi, Ngumi, 0.56S/38.04E, X1.1999, M. Snizek (MIZT); 21 ex., Kakamega Forest, Kaimosi, 0.12N 34.57E, IX.2001, Th. Wagner et al. (ZFMK): 106 ex., Kakamega Forest, nr. Buyangu, 0.13N/34.44E, IL, IV.2002, light trap (ZFMK); 5 ex., Chogoria, 0.14S/37.37E, 1800 m, X.2002, Co. & Th. Wagner (ZFMK). — Malawi: | ex., SW shore L Nyasa, be- tween Ft. Johnson & Monkey Bay, 1650 ft, 11.-11.1910, S. A. Neave (BMNH); | ex., W shore L Nyasa, between Domira Bay & Kotakota, 13.35S/34.27E, XI.1910, S. A. Neave (BMNH); 9 ex., Mlanje, 16.058/36.29E, IV.-VI1.1913, S. A. Neave (BMNH): 1 ex., Side of Mt. Mlanje, 16.00S/36.30E, 3000-4000 ft. XII.1913. S. A. Neave (BMNH); 3 ex., Zomdu Plateau, N. L. H. Krauss. B.M. 1957-3458 (BMNH); | ex., Mt. Mulanje, 15.58S/35.38E. 900 m, X11.1983, Bellamy (TMSA); | ex., Nyiaka Plateau, Chil- inda, 10.19S/33.48E, 2250 m, 1V.1991, D. Lachaise (MNHN). Mali: | ex., cercle Kandia, 14.46N/11.30W, 1935, P. Malzy (MNHN); 14 ex., IRCT, M'Pesoba, 12.40N/5.43W, IX.—X.1969, VII.-IX.1970, G. Pierrard (MRAC); 9 ex., Fana, 12.47N/6.57W, VIM.-IX.1969, VIII.1970, G. Pierrard (MRAC); | ex.. Kassorola, 14.08N/7.25W, X.1969, G. Pierrard (MRAC); 5 ex., Lutana, IX.1969, VIII.1970, G. Pierrard (MRAC). — Mogambique: | ex.. Mozamb. Nyassa, Thelwall (BMNH): | ex.. Zambese, 1882, Du- rand (MNHN); 1 ex., Beira, 19.49S/34.52E, 111.1903. P. A. Sheppard (BMNH); 1 ex., Vallée du Revoué, env. d’Andrada, 274 18.52S/32.53E, X1.1905, G. Vasse (MNHN); 2 ex., Chibababa, Lower Buzi R., X11.1906, C. F. M. Swynnerton (BMNH); 2 ex., Vallée du Pongoué, Guengére, 18.45S/33.40E, VI.1906, G. Vasse (MNHN); 2 ex., Prov. du Gorongoza, Tendo du Sungoue, 18.325/34.15E, 40 m, X.1907, G. Vasse (UN HN); | ex., Prov. du Gorongoza, Foret Dinhanoonde, 360 m, X.1907, G. Vasse (MNHN); 2 ex., Zambeze, Nova Choupanga pr. Chemba, 17.10S/35.00E, IV.1929, P. Lesne (MNHN); 1 ex., env. de Beira, Manga, 19.45S/34.50E, VII.1929, P. Lesne (MNHN); 1 ex., Ugano, Matengo-Hochland, wsw. V. Songen, 12.51S/35.48E, X11.1935, Zerny (HNHM); 1 ex., Missáo Zoologica, Mufo, 10.375/34.08E, VI1.1948 (ICT); 1 ex., Missáo Zoologica, Tica, 19.008/33.07E, V11.1948 (IICT); 3 ex., Missáo Zoologica, Gon- dola, 19.055/33.39E, VHI.1948 (ICT); 7 ex., Missao Zoologica, Bengo, 19.04S/33.37E, VIIL.1948 (ICT); 1 ex., Missáo Zo- ológica, Zina1, VHI.1948 (IICT); | ex., Missao Zoológica, Cosme, Vila Coutinho, 18.40S/34.24E, IX.1948 (IICT); 2 ex., Missao Zo- ológica, Tambara, 16.44S/34.15E, IX.1948 (IICT); 1 ex., Missäo Zoologica, Lifidzi, 14.33E/34.14E, IX.1948 (ICT). — Nigeria: | ex., Gomba, 12.55N/8.38E, Mataoro lakes, 1.1929, Dr. LI. Lloyd (BMNH); | ex., NW state, Kontagora River, 10.24N/5.29E, 3 mls. from Niger, VHI.1970, P. H. Ward, on semicultivated land (BMNH); 4 ex., SE State, Obudu Cattle Ranch, 6.38N/9.55E, V111.1974, R. Linnavuort (MZHF). — Rwanda: | ex., Bugarama, 2.27S/29.29E, 1.1926, H. Schouteden (MRAC); 1 ex., Gabiro, 1.535/30.23E, X.1932, L. Burgeon (MRAC); | ex., Bugarama, V.1951 (IRSNB); 2 ex., Kisenyi, 1.418/29.15E, VIII.1952, A. E. Bertrand (MRAC); 1 ex., Shangugu, 2.29S/28.54E, 1500 m, IV.1953, P. Basilewsky (MRAC); | ex., Kayove, terr. Kisenyi, 1.53S/29.21E, 2000 m, 11.1953, P. Basilewsky (MRAC); 1 ex., terr. Ruhengeri, Kagogo, 1.22S/29.46E, 1900 m, 1.1958, P. Basilewsky (MRAC); 3 ex., est Muhavura, 1.235/29.40E, 2100 m, 1.1958, P. Basilewsky (MRAC); 2 ex., env. Astrida, 2.36S/29.44E, 1954/1955, G. Foucart (MRAC); I ex., Nyarubuye, 1.585/29.59E, 111.1972, F. Cuypers (MRAC); 1 ex., Cyangugu, Nyakabuye, 2.10S/29.31E, X.-X1.1983, H. Mühle (CKi); 2 ex., Gako, Kibun- go, 2.128/30.27E, 1400 m, 1X.1984, H. Mühle (CKi); 4 ex., Kigali, 2.008/29.40E, X.1970, coll. Roggeman (CBe); | ex., Gi- furwe, 1.33S/29.48E, [V.1972, coll. Roggeman (CBe); 2 ex., Mt. Kigali, V.1972, coll. Roggeman (CBe); 2 ex., Icyanya, X.1973, coll. Roggeman (CBe); 2 ex., Gisenyi, 1.41S/29.15E, 1V.1973, coll. Roggeman (CBe); | ex., Tumba, 1.41S/29.57E, V.1973, coll. Roggeman (CBe); 3 ex., Nasho I, 2.025/30.43E, VIII.1988, E. Fischer (ZFMK); 41 ex., Rusumo, Ibanda Makera, 2.13S/30.42E, X.1993 / 1X.2002, Th. Wagner, partim insecticidal tree fogging of Lannea fulva, Teclea nobilis (ZFMK). — Senegal: | ex., Kati, 13.54N/14.57W, VII.1913, coll. Clavareau (MRAC); I ex., Sin- thiou-Maleme, 13.03N/15.18W, VIII.1939, J. Risbec (BMNH); 2 ex., Guede, 14.53N/15.52W, 11.1946 (BMNH); 7 ex., Sébikotane, 14.45N/17.05W, XI-XI1.1945, 111.1946, VII.1947, H. Durand (MNHN); 1 ex., Diakéne, 12.27N/16.37W, X.1960, Sala (MCTB); | ex., Sangalkam, 14.47N/17.13W, VIIL.1967, A. Des- carpentries et al. (MNHN); 1 ex., Sebikotane, VIIL.1971, A. Villiers (MNHN). — Sierra Leone: | ex. (BMNH); | ex., M. pul- chella Klug, Jac. coll. (BMNH); 1 ex., Baly coll. (BMNH); 1 ex., ex coll. J. Weise (ZMHB); 2 ex., Rhobomp, coll. Kraatz / coll. Ja- coby (BMNH, DEI); | ex., ex coll. J. Weise (ZMHB); 1 ex., V111.1892, Rev. D. Z. Morgan, duplicata Sahlberg (BMNH); 1 ex., Kaballa, 9.35N/11.33W, V.1912, J. J. Simpson (BMNH). — Somalia: 2 ex., Mogadiscio, 2.04N/45.22E, ex. coll. Breuning (MRAC); | ex., nr. Hargeissa, V1.1963, Linnavuori (MZHF); | ex., Mogadiscio, 111.1974 (CMa). — Sudan: | ex., Latuka, 4.39N/32.33E, 1918, J. R. Yardley (IRSNB); | ex., W. Darfur, N. Jebel Murra, 10.21N/23.24E, 5600 ft., VII.1932, M. Steele (BMNH); | ex., Upper Nile, Malakal, 9.32N/31.39E, 1.1963, Lin- Bonner zoologische Beitráge 53 (2004) navuori (MZHF); 1 ex., Equatoria, Lotti Forest, 4.00N/32.32E, 111.1963, R. Linnavuori (MZHF); 3 ex., Equatoria, Imatong Mts. nr. Gilo, 4.03N/32.51E, HI.1963, Linnavuori (MZHF); 1 ex., Equatoria, Kapoeta-Boma, 4.46N/33.35E, 11.1963, Linnavuori (MZHF); | ex., Jebel Marra, 12.55N/24.12E, Gallery forest, on dry rock in stream, 1X.1983, R. Moore (BMNH). — Tanzania: | ex., Usambara, Nguela, 4.45S/38.30E (IRSNB); 1 ex., Moshi, 3.20S/37.21E, coll. Hauser (IRSNB); 1 ex., Zanzibar, 6.10S/ 39.12E, Raffray (MNHN); 12 ex., Ukerewe Island, 2.098/32.52E, Conrads (4 ex. IRSNB, 8 ex. NMK); 1 ex., Arusha-Chini, Katona, 3.098/38.26E (NMK); 1 ex., inter Marti et Arusha, Katona (NMK); 1 ex., Kibosho, Katona (NMK); 2 ex., Montes Lettema, Katona (NMK); 1 ex. Kilimandjaro, 3.04S/37.22E, Katona (NMK); 1 ex., Kilimandjaro, Bornemissza (NMK); 1 ex., Kili- mandscharo, Dschagga-Land, Madchame, T. Paesler (ZMHB); | ex., “Kilimandjaro / ex. coll. Weise / Type / Monolepta sjóstedti m [invalid type] (ZMHB); | ex., “Kilimandjaro, Sjóstedt. 1905-6 / Kibonoto 00-1900 / ex. coll. Weise / Type [invalid type] (ZMHB); | ex., Kirumba b. Muansa, 2.31S/32.54E, Holtz (ZMHB); 4 ex., Parek, 2000 m, Ch. Schróder (ZMHB); 4 ex., Kikuiru, Mas- saisteppe, Ch. Schróders (ZMHB); 4 ex., Kilimandjaro, Sjóstedt (ZMHB); | ex., Kilimandjaro, Sjóstedt, Kibonoto, 1’-1200 m, IV. (NHRS); 1 ex., Kwai, 4.44S/38.21E, Paul W. (NHRS); 2 ex., Usambara, Derema, 4.45S/38.30E, 850 m, IX.-X.1891, Conradt (ZMHB); 2 ex., Langenburg, Ineteyanur nya a. Nyassa, 9.01S/ 33.39E, VI,1898, Fülleborn (ZMHB); 1 ex., Rungwe, 9.15S/ 33.40E, 11.1899, Stolz (ZMHB); 1 ex., W.-Pemba, 111.1903, Voeltzkow (ZMHB); | ex., Pemba, Chake-Chake, 5.15S/39.45E, IV.1903, Voeltzkow (ZMHB); 1 ex., Pemba, Fundu, 5.03S/ 39.39E, 1V.1903, Voeltzkow (ZMHB); 2 ex., Moshi, 3.20S/37.21E, 1905, Fl. Rau, Katona (NMK); 1 ex., Amani, 5.095/38.36E, X.-XH.1905, 900 m, Ch. Schröder (ZMHB); 7 ex., Arusha-Ju, 3.22S/36.38E, X1.1905, Katona (NMK); 2 ex., Papy- rus-Sumpf, süd 6. Kilimandjaro, 1.1906, Ch. Schröder (ZMHB); 2 ex., Panganisteppe, Mombo-Masinde, 1.1906, Ch. Schróder (ZMHB); 1 ex., Amani, 11.1906, Vosseler (ZMHB); 1 ex., D. Sambesi Geb., X.1906, F. Seiner (ZMHB); 1 ex., Shirati, 1.08S/33.59E, 11.1909, Katona (NMK); 6 ex., Lake Ippe, 3.35S/37.45E, 1910, Gotsch (NHMW); 1 ex., Meru, Ngare na Nyuki, 3.09S/36.51E, Sjöstedt (NHRS); 3 ex., Usangu Distr., 8.30S/34.15E, X1.—X11.1910, S. A. Neave (BMNH); 1 ex., Kili- mandscharo, 2500-3000 m, am Bismarckhügel oberh. Marangu, südl. Mawenzi am Fuße der Hochweiden, 11.1912, Ch. Schröder (ZMHB); | ex., Nord-Uluguru, 7.05S/37.35E, 1400-1900 m, 11.1914, Methner (ZMHB): 4 ex., Kirumba, X1.1914, 11.1915, Holtz (3 ex. ZMHB, | ex. ZMUH); 2 ex., Narobi bei Tanga, 5.058/39.02E, IV.1915, Methner (ZMHB); 2 ex., Muansa, VI- 11.1915, Holtz (ZMHB); | ex., K-D River Camp, Noisinak Bridge, 3500 ft, V.1916, W. A. Lamborn (BMNH); 5 ex., E. Tanganyika, Kigoma, 4.52S/29.36E, IX.1918. R. Mayné (MRAC); 1 ex., Nata, 2.00S/34.24E, 11.1960, Dr. Szunyoghy (NMK); 1 ex., Bunduki, Uluguru Mts., 7.058/37.35E, 6000 ft, X1.1948, J. G. Williams (NMK); 5 ex., Bunduki, Uluguru Mts., Moy. Mgeta, 7.02S/ 37.34E, 1300 m, IV.-V.1957, P. Basilewsky & N. Leleup (MRAC); 4 ex., Mt. Hanang, 2200-2500 m, 4.26S/35.24E, V.1957, P. Basilewsky & N. Leleup (MRAC); 1 ex., Ngorongoro Rest Camp, 3.11S/35.23E, 2400-2500 m, V1.1957, P. Basilewsky & N. Leleup (MRAC); 4 ex., Mt. Kilimandjaro, Marangu, 3.178/37.31E, 1600-1700m, VII.1971, P. Basilewsky & N. Leleup (MRAC); 2 ex., Mulukulazo, 111.1969, I. A. D. Robertson, sweep net (BMNH); | ex., Ngorongoro Crater, 1500 m, Berger et al. (MRAC); | ex., Morogoro, 6.49S/37.40E, 600 m, V.-VL1971, L. Berger et al. (MRAC); 1 ex., Uluguru Mts., Kiroka, 6.50S/37.49E, 725 m, V.1971, L. Berger et al(MRAC); 1 ex., Uluguru Mts., Kinola, 1500-1750 m, VI.1971, L. Berger et Thomas WAGNER: Revision of the vincta Species-group of Monolepta Chevrolat 27 al.(MRAC); | ex., Uluguru Mts., Lukwangule Plateau, 2400-2600 m, VII.1971, L. Berger et al. (MRAC); 5 ex., Uluguru Mts., Chenzema, 7.06S/37.35E, 1700 m, VII.1971, L. Berger et al.(MRAC); 2 ex., Morogoro, 1.1974, H. Silfverberg (MZHF); | ex., Uzungwa Mts., Mwanihana F. above Sanje, 8.20S/35.58E, 1000 m, VIII.1982, M. Scholtze & N. Scharff, pitfall trap (ZMUC); | ex., Uzungwa Mts., Chita FR, 8.31S/35.56E, 750 m, X.1984, M. Scholtze & G. Petersen (ZMUC); | ex., Morogoro, 6.49S/37.40E, University Campus, 1.1988, T. Pócs, at light (NMK); 1 ex., Uluguru Mts., 1V.1991, Werner (CD6); 2 ex., Pan- gani Falls Forest, 5.20S/38.40E, 1.-111.1993 (ZMUC); 11 ex., Mkomazi G. R., 4.10S/38.10E, Acacia tortilis, Acacia mellifera, Acacia recifiens, Acacia senegal, Lannea stuhlmanni, Melia volkensii, Terminalia sp.A canopy sampling, VII.—VIIL.1994, G. McGavin (BMNH); 150 ex., Mufindi Distr., Iringa, Ut’zungwa Scarp FR, 8.31S/35.54E, 750 m, 1111996, McKamey et al., can- opy UV and fog (ZMUC); 2 ex., 12 km O Iringa, Little Ruawa Riverside Camp, 7.48S/35.480, 1X.2002, U. Heinig (CHe). — Togo: 8 ex., Bismarckburg, 8.15N/0.55E, X11.1892, V.-V1.1893, L. Conradt (ZMHB); | ex., Misahóhe, 6.59N/0.40E, V1.1894, F. Baumann (ZMHB). — Uganda: | ex., Mulangu, 0.30N/32.02E, ex. Staudinger (MRAC); 2 ex., Bussu, Busoga, 0.45N/33.30E, 11.1909, 1910, E. Bayon (MCSN); | ex., Mt. Kokanjero, SW of Elgon, 6400 ft, VIII.1911, S. A. Neave (BMNH); | ex., Siroko R. nr W foot of Mt. Elgon, 1.29N/34.14E, 3600 ft, VIIL.1911, S. A. Neave (BMNH); 2 ex., between Seziwa R. & Kampala, VIII.1911, S. A. Neave (BMNH); | ex., Daro/Durro Forest, Toro, 4000-4500 ft., X.1911, S. A. Neave (BMNH); | ex., Mpanga For- est, Toro, 0.15N/32.05E, 4800 ft, X1.1911, S.A. Neave (BMNH); l ex., Tero Forest, SE Buddu, 0.50S/31.40E, 3800 ft, IX.1911, S. A. Neave (BMNH); 1 ex., Bugoma, Forest, Unyoro, 1.15N/ 30.53E, 3700 ft, XII.1911, S. A. Neave (BMNH); 2 ex., Kafu R. near Hoima, Kampala Rd., 1.25N/31.22E, 3500 ft, XI1.1911,S. A. Neave (BMNH); 1 ex., Between Mitiana & Entebbe, 0.24N/32.03E, 3800 ft., 1.1912, S. A. Neave (BMNH); 2 ex., N of L. Isolt, 3700 ft., 1.1912, S. A. Neave (BMNH): l ex., Shores of L. Isolt, Wamala, 0.04N/31.54E, 3800 ft, 1.1912, S. A. Neave (BMNH); | ex., S foot & slopes of Mt. Elgon, 5100-5800 ft, 1912 (BMNH); | ex., Hima Riv., 0.18N/30.10E, IV.1912, Dr. Bayer (MRAC); 1 ex., Tero, 0.50S/31.40E, V11.1912, C. C. Gowdey (ZMUH); 3 ex., Mujenje, VIII.1913, Katona (NMK); 9 ex., Kampala, 0.19N/32.35E, X.1920, IX.1947, IV.-VI.1948, XII.1925, A. F. J. Gedye (NMK); 2 ex., Jinja, 0.19N/33.59E, V111.1924, VIII.1926, A. F. J. Gedye (NMK); 1 ex., Buamba, 7000 ft, X.1926 (ZMUH); 9 ex., Kiryanga, 1.06N/31.04E, IV.1927, H. Hargeraves (5 ex. BMNH, 4 ex. ZMUH); | ex., Bu- songoro, 0.05N/30.00E, X11.1927, G. D. H. Carpenter (BMNH); | ex., Budongo, 1.45N/31.35E, 1X.1932, T. Jackson (BMNH); 1 ex., Ruwenzori Range, Kilembe, 0.12N/30.00E, 4500 ft., XI1.1934-1.1935, F. W. Edwards (BMNH); | ex., Budongo F., 1.45N/31.35E, X.1936, A. F. J. Gedye (NMK); 1 ex., Bubulu, 0.56N/34.16E, X.1938, T. H. C. Taylor (BMNH); | ex., Kyagwe, 0.25N/32.50E, V1.1938, T. H. C. Taylor (BMNH); l ex., Kam- pala, 0.19N/32.35E, VI.1940, A. F. J. Gedye (NMK); 3 ex., W of Ruwenzori, 6000 ft, VII.1945, van Someren (1 ex. BMNH, 2 ex. NMK); 2 ex., Bwamba, 0.43N/30.04E, VIL.-VII1.1946, V1.1948, v. Someren (NMK); 1 ex., Kampala, 3200 ft, XI1.1946, A. F. J. Gedye (USNM); 2 ex., Bwamba Forest, 2500 ft., 111.1948, J. G. Williams (NMK); 5 ex., Bwamba, Hakitengya, X1.1948, W. H.R. Lumsden, from fan trap (BMNH); 1 ex., Mafuga Forest, 1.03S/29.52E, IM.-VI.1951, T. H. E. Jackson (NMK); 1 ex., Ru- wenzori Range, Bugoye, 4500 ft, IX.1952, D. S. Fletcher (BMNH); I ex., Ruwenzori Range, Semliki F., 2850 ft, VIIL- IX.1952, D. S. Fletcher (BMNH); | ex., Kampala, X11.1952, A. F. Gedye (NMK); 1 ex., Katera, 0.428/32.00E, X1.1953, T. H. E. Un Jackson (NMK); | ex., Jinja, 0.19N/33.59E, X11.1954-11.1955, P S. Corbet, M.V. light trap (BMNH); 2 ex., Bugiri, 0.34N/33.45E, 1400 m, P. VII1.1957, Basilewsky & N. Leleup (MRAC); | ex., Kawanda, 0.01S/31.29E, 11.-111,1958, P. Whalley, M. V. light trap (BMNH); 2 ex., Entebbe, 0.05N/32.29E, 111.1972, Falk (MRAC) 315 ex., Kibale Forest, 0.50N/31,06E, VIL—VIIL, X.1983, IV.-V,, VII, IX.-X11.1984, I., HE-IV. X11.1985, 1991, 1X.199), M Nummelin, sweep (MZHF); 265 ex., Budongo Forest nr. Sonso, 1.45N/31.35E, VI.1995, insecticidal canopy fogging of Cy- nometra alexandri, Rinorea beniensis, Teclea nobilis, Trichilia rubescens, Th. Wagner (ZFMK); 3 ex., Semliki Forest, 0.48N/30.09E, 11.1997, U. Göllner (ZMHB); 15 ex., Semliki For- est, 11.1997, Th. Wagner, insecticidal fogging of Cynometra alex- andri, Elais guinensis, U. Göllner (1 ex. ZMHB, 14 ex. ZFMK); 5 ex., Nyabyeya, 1160 m, 1.40N/31.32E, 1X.1997, Th. Wagner (ZFMK); | ex., Kibale Forest, nr. MUBS, VIII.1997, M. Boppré (ZFMK); 15 ex., Kibale NP, U. Kanyawara, 0.50N/31.06E, 1600 m, VIII.1998, L. Schmidt (ZFMK); 1 ex., Kasese, Queen Elisa- beth NP, Mweya, 0.10N/30.05E, VIII.1998, I. Schmidt (ZFMK). Zambia: 3 ex., Niamadzi R., nr. Nawalia, VIII.1910, S. A. Neave (BMNH); 1 ex., Upper Luangwa R., 14.49S/19.06E, VIT.—VIII.1910, S. A. Neave (BMNH); 3 ex., Caia, Zambesi, 17.49S/35.23E, X.1911, H. Swale (BMNH); | ex., Namwala, 111.1913, H. C. Dolman (BMNH); 1 ex., Chilonga, 12.02S/31.21E, VII.1913, on Citrus trees infested with black aphids (BMNH); 10 ex., Mwengwa, 13.00S/27.66E, VII.—VIII.1914, H. C. Dollman (BMNH); 14 ex., Lukanga, 14,33S/27,48E, VII.1915, H. C. Dol- man (BMNH); | ex., Kashitu, 13.45S/28.40E, N. of Broken Hills. VII.1915, H. C. Dolman (BMNH); 1 ex., N’Changa, 13.295 30.27E, 1931, C. T. MacNamara (BMNH): | ex., Sesheke, 16.45S/24.20E, 950 m, HI.—VI.1991, W. Slobbe (NNML); 3 ex.. Kafue River, Rimo Marine Motel, 15.49S/28.12E, III.1993, U. Göllner (ZMHB). — Zimbabwe: | ex., Vumba Mts., Nebelwald. ca.1500 m, 19.06S/32.47E, 111.2000, U. Heinig (CHe); 1 ex.. Kariba, Mopani Bay Camp, 16.31S/28.49E, 111.2000, R. Beenen (CBe); 1 ex., Chimanimani NP, 19.48S/32.52E, XII.1998, S. Beévar (CBz); | ex. dto., A. Kudrna (CBe). Monolepta ronbeeneni sp. n. Description. Total length. 3.10-3.60 mm (mean: 3.25 mm; n = 10). Head. Yellow to yellowish-red, mouth-parts including palpi brownish (Fig. 27). Antennae pale yellow, last two to four antennomere brownish or black, all antenno- meres very elongated (Figs 27, 28), length of antenno- meres two to three 1.00—1.10 (mean: 1.04), length of an- tennomeres three to four 0.41—0.62 (mean: 0.46). Thorax. Prothorax pale yellow to yellowish (Fig. 27). Pronotal width 0.85—1.05 mm (mean: 0.96 mm), prono- tum broad, pronotal length to width 0.59-0.64 (mean: 0.61), very finely punctured. Elytra yellow to reddish- yellow, base with transverse black band, often undulate posteriorly (Fig. 27a), subapical usually with large ovate spot, which is often more brownish and not sharply de- limited. Elytral length 2.30-2.80 mm (mean: 2.42 mm). maximal width of both elytra 1.45-1.75 mm (mean: 1.65 mm), maximal width of both elytra to length of elytron 0.66-0.71 (mean: 0.69). Scutellum yellow. Meso- and metathorax yellow to yellowish-red, rarely brownish, legs yellow. 276 Bonner zoologische Beitrage 53 (2004) Figs 27-31: Monolepta ronbeeneni sp. n. 27: colour pattern; 28: basal antennal articles (a: Ú, 31a b C d b: Y); 29: three different sper- mathecae; 30: bursa-sclerites (a: dorsal, b: ventral); 31: median lobe (a: lateral, b: dorsal from Albertine Rift, c: dorsal from west- ern Central Africa, d: ventral, without endophallic structures). Abdomen. Yellow to yellowish-red. Female genitalia. Spermatheca with small spherical nodulus, comparatively broad middle part and long, strongly curved cornu (Fig. 29). Dorsal part of bursa sclerites small, elongated triangular (Fig. 30a), ventral part very broad (Fig. 30b). Male genitalia. Median lobe straight, very slender at apex, in specimens along the Albertine Rift apex slightly enlarged (Fig. 31b), specimens from western Central Africa apex parallel-sided (Fig. 31c). Tectum comparatively broad, ventral groove parallel-sided (Fig. 31d). Dorsal pair of median endophallic spiculae very strong, other median spiculae slender, ventral spiculae absent, lateral spiculae with large U-shaped apex (Fig. 31b), which is slenderer in most specimens from west- ern Central Africa (Fig. 31c). Distribution. Forests and savannas of northern Central Africa from northern and the Central African Republic to Uganda and Rwanda (Fig. 32). Diagnosis. Monolepta ronbeeneni sp. n. is one of the smallest afrotropical Monolepta species. It is equal in size and antennal characters to M. wittei Laboissiere, 1940, a species which differs in having throughout yel- lowish-red elytra, a black apical abdominal segment, very different male genital characters, and is restricted to Lake Kivu region at the Albertine Rift (WAGNER 2002). Monolepta buquetii can be easily distinguished by the short basal antennomeres, the short antennae, broader pronotum, it has no reddish-yellow elytral col- oration and does not occur along the Albertine Rift. Only a very few specimens of M. vincta and M. sharo- nae sp. n. are as small as M. ronbeeneni sp. n., which Thomas WAGNER: Revision of the vincta Species-group of Monolepta Chevrolat 2 meen M. sharonae sp. n. IA RAS M. ronbeeneni sp. n. o 1-4 @ 5-9 (A) 10-16 (@) 17-42 Fig. 32: Distribution of Monolepta ronbeeneni sp. n., and M. sharonae sp. n. can be distinguished by more slender antennae and dif- ferent elytral coloration. Etymology. Dedicated to the Chrysomelidologist Ron Beenen, Nieuwegein, The Netherlands. Type material. Holotype: & “Congo Belge, P.N.G., Aka, 19.V.1952, H. de Saeger, 3484” (MRAC); type locality: Congo, Pare National d'Garamba, 3.40N/29.00E. Paratypes: Central African Republic: | ex., Dar-Banda, Merid., Krébédjé, Fort Sibut, 5.46N/19.06E, Miss. Chari-Tchad, X.1904, J. Decorse (MNHN); | ex., Uamgebiet, Bosum, 6.19N/16.38E, V1.1914, Tessmann (ZMHB). — Congo: 2ex., Ituri, La Moto, Madyu, 2,27N/26.25E, |. Burgeon (MRAC); | ex., Viag. Ruwenzori, S. A. R. D. Abruzzi (MIZT); 5 ex., Haut-Uelé, Moto, 2.27N/26.25E, 1920, V.1923, L. Burgeon (MRAC) 3 ex., Haut-Uelé, Abimva, 3.09N/29.50E, V.1925, L. Burgeon (MRAC); 2 ex., alto Uelle, Yaku- luku, 4,20N/28.48E, IV.1927, F. S. Patrizi (MCSN); 4 ex., alto Uelle, Kapili, 3.428/27.53E, V.1927, F. S. Patrizi (MCSN); | ex., Ituri, Ni- oka, 2.10N/30.39E, V1.1934, J. V. Leroy (MRAC); 2 ex. Rutshuru, 1.115/29.27E, X11.1937, J. Ghesquiere (MRAC); 1 ex., Mongbwalu, 1.57N/30.02E, VI.1938, Scheitz (MRAC); P.N.Garamba, 3.40N/29.00E: 1 ex., I/a/3, 11.1950, H. de Saeger (MRAC); 2 ex., A- kam, V.1950, H. de Saeger (IRSNB, MRAC); 4 ex., Vo/l & Vo/2, V.1950, H. de Saeger (MRAC); 1 ex., km 31, V.1950, H. de Saeger (MRAC); 11 ex., Vo/l & 1/0/2, X.1950, Miss. H. de Saeger, G. De- moulin (MRAC); 2 ex., Il/ad/9, X1.1951, H. de Saeger (MRAC): 4 ex., Garamba/2, source, V1.1952, H. de Saeger (MRAC): | ex., PFSK, 22/8, V1.1952, H. de Saeger (MRAC); 1 ex., PFNK, 12/9, V11.1932, H. de Saeger (MRAC); | ex., Mabanga, 4.22N/29.47E, 11.1952, H. de Saeger (IRSNB); 8 ex., Pidigala, 4.27N/29.28E, IV.1952, H. de Sae- ger (1 IRSNB, 7 MRAC); 1 ex., Mt. Embe, 4.40N/29.33E, IV.1952, H. de Saeger (MRAC); 42 ex., Aka2, 3.50N/28.57E, V.1952, H. de Saeger (12 ex. IRSNB, 30 ex. MRAC); 16 ex., Inimvua, 4.35N/29.43E, V.1952, H. de Saeger (2 ex., IRSNB, 14 ex. MRAC): | ex., Iso, 4.18N/29.56E, III, IX.1952, H. de Saeger (MRAC); | ex.. Mt. Moyo, 1.13N/29.49E, IX.1952, H. de Saeger (MRAC); P.N.A., Massif Ruwenzori, 0.20N/29.50E: | ex., Kyandolire, 1700 m, X.1952, P. Vanschuytbroeck (MRAC); 5 ex., Mt. Degio, 2200 m, VII.1954, P Vanschuytbroeck & H. Synave (MRAC); | ex., Litongo, 1575 m, VII.1954, P. Vanschuytbroeck (MRAC); 1 ex.. Kombo, 1550 m, 278 (MRAC); 1 ex., Kombo, 1550 m, V11.1954, P. Vanschuytbroeck (MRAC): 1 ex., Secteur Nord, Muramba, 1050 m, X1.1956, P. Van- schuytbroeck (MRAC); | ex., Katibombo, camp des gardes, 900 m, X.1957, P. Vanschuytbroeck (MRAC). — Rwanda: | ex., Kibungu, 2.10S/30.32E, X.—X11.1937, R. Verhulst (MRAC); 30 ex., Rubona, 2.13S/29.43E, “s/haricots”, X.1966, LS.A.R. (MRAC); 1 ex., Rubona, V.1963, G. Pierrard (MRAC); 2 ex., Icyanya, X.1972, coll. Roggeman (CBe); 4 ex., Mayaga, 2.06S/29.59E, V.1973, coll. Roggeman (CBe). — Tanzania: | ex., Mujenje, VII.1913, Kittenberger (ZMHB). — Uganda: 1 ex., Koki Country, SW Buddu, 0.25S/31.40E, 4100 ft, X.1911, S. A. Neave (BMNH); 1 ex., SE Ankole, 0.45S/30.45E, 4400-400 ft, X.1911, S. A. Neave (BMNH); 1 ex., Kalwa, IX.1932, T. Jackson (BMNH); 1 ex., Budongo, 1.45N/31.35E, IX.1932, T. Jackson (BMNH); | ex., Kawanda, I1.-111.1958, at light trap, P. Whal- ley (BMNH). Monolepta sharonae sp. n. Description. Total length. 3.40-4.20 mm (mean: 3.75 mm; n= 8). Head. Including mouth parts pale red to yellowish-red, with deeply incised transverse post antennal suture (Fig. 33). Antennae pale yellow, usually only tip of last an- tennomere dark brown, antennomeres 4 to 10 short, less than 4 times long than broad at apex. Length of anten- nomeres two to three 0.85-0.95 (mean: 0.90), length of antennomeres three to four 0.43—0.50 (mean: 0.47) (Fig. 34). Thorax. Prothorax entirely pale yellow. Pronotal width 1.00-1.15 mm (mean: 1.08 mm), pronotal length to width 0.60-0.63 (mean: 0.61), finely punctured, prono- tum usually significantly narrowed at base. Anterior quarter of elytra black, transverse black spots in the api- cal third of each elytron (Fig. 33). Elytral length 2.70- 3.20 mm (mean: 2.85 mm), maximal width of both elytra 1.80-2.15 mm (mean: 2.04 mm), maximal width of both elytra to length of elytron 0.68-0.73 (mean: 0.71). Scutellum yellowish-red. Meso- and metathorax, coxa, trochanter an basal two third of femur reddish- yellow, outer parts of legs yellow. Abdomen. Pale yellow. Female genitalia. Spermatheca small, with small nodulus, slender middle part and slender cornu (Fig. 35). Dorsal part of bursa sclerites broad at base strongly narrowed towards apex (Fig. 36a), ventral part slender, outer margin finely serrate (Fig. 36b). Male genitalia. Apical part of median lobe strongly nar- rowed, conical and dorsally bent towards apex (Fig. 37a), basal three quarter broad, parallel-sided (Fig. 37b). Tectum very broad, ventral groove of median lobe wide, narrowed towards the orifice (Fig. 37c). One median endophallic spiculum large and usually bent towards the right in dorsal view, another rpart of median spiculae Bonner zoologische Beitráge 53 (2004) short and twisted, ventral spiculae strongly bent, lateral spiculae complex curved, with strong, medially oriented spine at apex (Fig. 37b). Distribution. This species is known from Nigeria through the Congo Basin to western Uganda and Rwanda, southwards to Central Mocambique and An- gola (Fig. 32) and can be expected also in Rwanda, western Tanzania and Zambia. Diagnosis. Monolepta sharonae sp. n. is very similar to specimens of M. vincta, which has the same coloration type (Fig. 18c). Since it is sympatrically distributed with the very polymorphic M. vincta, both species can be clearly distinguished by examination of the male genita- lia only (Figs 24, 24, 25, 37), while doubtless identifica- tion of females is not possible. M. sharonae sp. n. is on average smaller (total length up to 4.2 mm, M. vincta up to 4.7 mm), the basal antennomeres are longer (anten- nomere 3 to 4: 0.43-0.50, M. vincta: 0.28—0.37), and the post antennal suture is deeply incised in all known specimens of M. sharonae sp. n., but rarely found in M. vincta. Since only about ten percent of all available males of both species were examined in this study, there might be some more specimens of the new species be- yond the material identified as M. vincta. Etymology. Dedicated to the Chrysomelidologist Sharon Shute, The Natural History Museum, London, UK. Type material. Holotype. © “Angola, (A26), Salazar, 1.1.A.A., 9.- 15.11.1972 / at light / Southern African Exp. B. M. 1772-I (BMNH); 9.18S/14.55E Paratypes. Cameroon: 1 EX. Bafoussam, 5.31N/ 10.25E, 11.1969, N. Berti (MNHN). — Congo: | ex., Ituri, La Moto, Madyu, 2.27N/26.25E, L. Burgeon (MRAC); 1 ex., Beni a Lesse, 0.30N/29.28E, VIL1911, Dr. Murtula (MRAC); 1 ex., Amadi, 3.35N/26.47E, IV.1913, P. van den Plas (MRAC); 1 ex., Rutshuru, 1.11S/29.27E, V.1937, J. Ghesquiere (MRAC); 1 ex., P. N. Garamba, 3.40N/29.00E, l/o, V.1950, H. de Saeger (MRAC); 1 ex., P. N. Garamba, I/c, VIII.1950, G. Demoulin (MRAC); lex., P. N. Garamba, Vo/2, X.1950, H. de Saeger (MRAC); 2 ex., P. N. Garamba, II/gd, V.1951, H. de Saeger (MRAC); 3 ex., P. N. Garamba, PpK/60/d, XII.1951, H. de Saeger (MRAC); 2 ex., P. N. Garamba, IVfd, X11.1951, H. de Saeger (MRAC); 1 ex., P. N. Garamba, Tori, 111.1952, H. de Saeger (MRAC); 1 ex., P. N. Garamba, Ndelele, 4.22N/29.47E, VI.1952, H. de Saeger (MRAC); 1 ex., P. N. Garamba, PFSK, VI.1952, H. de Saeger (MRAC); 1 ex., P. N. Garamba, Ante, VII.1952, H. de Saeger (MRAC); 2 ex., P. N. Garamba, IVjd/9, 1., VIII.1952, H. de Saeger (MRAC). — Mocambique: | ex., Beira, 19.498/34.52E, V1.1900, G. A. K. Marshall (BMNH). — Nigeria: | ex., Yengre, 10.14N/8.48E, VIII.1973, R. Linnavuori (MZHF). — Rwanda: | ex., Ruhengeri, source Kirti, 1.355/29.40E, 1800-1825 m, X.1934, G. F. de Witte (MRAC); 1 ex., Kagogo, 1.22S/29.46E, 1990 m, 1.1953, P. Basilewsky (MRAC). — Uganda: | ex., Daro For- est, Toro, 1.05S/31.00E, 4000-4500 ft, X.1911, S. A. Neave (BMNH). Thomas WAGNER: Revision of the vincta Species-group of Monolepta Chevrolat Figs 33-37: Monolepta sharonae sp. n. 33: colour pattern; 34: basal antennal articles (a: ©, b: ); 35: three different spermathe- cae; 36: bursa-sclerites (a: dorsal, b: ventral); 37: median lobe (a: lateral, b: dorsal, c: ventral, without endophallic structures). Monolepta naumanni sp. n. Description. Total length: 3.60-4.80 mm (mean: 4.23 mm; n = 10). Head. Yellow, some specimens with pale brownish ver- tex (Fig. 38b), labrum, mandibles, maxillary and labial palpi brown. Antennae yellow, usually only last anten- nomere brown, rarely last two to three antennomeres brown (Fig. 38c). Antennae long, antennomeres 4 to 10 slender, elongate. Length of antennomeres two to three 1.00-1.05 (mean: 1.01), length of antennomeres three to four 0.37-0.43 (mean: 0.39). Thorax. Prothorax entirely pale yellow. Pronotal width 1.10-1.45 mm (mean: 1.26 mm), pronotal length to width 0.58-0.61 (mean: 0.59), very finely and irregu- larly punctured. Anterior third of elytra black or dark brown, in 20 % of specimens examined reddish brown, middle pale brownish-yellow, each elytron with black (10 %; Fig. 38b), brown (70 %; Fig. 38c) or without (20 %; Fig. 38a) subapical spot, this spot with reddish outer margins (Figs Figs 38b, c). Elytral length 2.80-3.50 mm (mean: 3.18 mm), maximal width of both elytra 1.90 2.50 mm (mean: 2.31 mm), maximal width of both elytra to length of elytron 0.71-0.75 (mean: 0.73). Scutellum yellow or yellowish-red. Meso- and metatho- rax and legs brownish-yellow, apex of femur, tibia and tarsi paler yellow. Abdomen. Brownish-yellow. Female genitalia. Spermatheca small, with small nodulus, slender middle part and long and slender cornu (Fig. 40). Dorsal part of bursa sclerites slender, with small spines (Fig. 41a), ventral part slender, outer mar- gin finely serrate (Fig. 41b). Male genitalia. Apical part of median lobe narrow, conical towards apex, basal three quarter parallel-sided (Fig. 42). Tectum broad, conical (Fig. 42b), ventral groove of median lobe wide, strongly narrowed towards the orifice (Fig. 42c). Median endophallic spiculae slen- der, ventral spiculae strong, hook-like, lateral spiculae broad, short, plate-like medially with spur (Fig. 42b). Distribution. Known from wet tropical forests from eastern Congo and Rwanda to southern Uganda, West- ern Kenya (Kakamega Forest) and north-west Tanzania (Ukerewe Island in Lake Victoria; Fig. 17). Diagnosis. Monolepta naumanni sp. n. can be distin- guished from all other species of the Monolepta vincta- group by the elytral coloration where subapical elytral spots have red outer margins. A very similar coloration ata Labois- is found in some specimens of M. quadriz )] siere, 1940, which occurs syntopely with M. naumann sp. n. at some locations, but ıs much larger (total length } 5.50-6.40 mm), elytra are very slender, and genital characters of both sexes are very different (WAGNER 280 Bonner zoologische Beitráge 53 (2004) Figs 38-42: Monolepta naumanni sp. n. 38: colour pattern; 39: basal antennal articles (a: 42a e 7, b: 2); 40: three different spermathe- cae; 41: bursa-sclerites (a: dorsal, b: ventral); 42: median lobe (a: lateral, b: dorsal, c: ventral, without endophallic structures). 2003b). Specimens without apical spot and reddish- brown elytral base are very similar to some M. umbro- basalis Laboissiere, 1940, which also occurs syntopely with M. naumanni sp. n. along the Albertine Rift and is very similar in size (total length, 3.30-4.10 mm) and ex- ternal characters, but has very different genital charac- ters including a broad and apically depressed median lobe (WAGNER 2002). Etymology. Dedicated to the late Clas Michael Naumann, my unforgotten mentor. Type material. Holotype: © “Uganda, District Masindi, Budongo For- est n. Sonso, 1°45’N, 31°35’E, 1.-10.VIL9S, Th. Wag- nerleg. / T.r- 15/2” (ZFMK) Paratypes: Congo: | ex., Malela, 4.22S/26.08E, X11.1913, L. Bur- geon (MRAC); | ex., Alto Uelle, fl. Ouru, 2.07N/31.14E, 11.1927, F. S. Patrizi (MCSN); 1 ex., Mongbwalu, Kilo, 1.49N/30.06E, 1938, Mme Scheitz (MRAC); 1 ex., Stanleyville, Ongoka, riv. Lowa, 1.238/26.02E, IV.-IX.1952, J. Pantos (MRAC); 1 ex., P. N. Garam- ba, Mt. Embe, 4.40N/29.33E, IV.1952, H. de Saeger (MRAC); 4 Thomas WAGNER: Revision of the vincfa Species-group of Monolepta Chevrolat 28 (MRAC); 4 ex., P. N. Garamba, Aka, 3.50N/28.57E, V.1952, H. de Saeger (MRAC); 3 ex., P. N. Garamba, Dedegwa, 4.35N/29.43E, IV., V.1952, H. de Saeger (MRAC); 1 ex., Kivu Sud, Irangi, 1.545/28.27E, 900 m, X.1993, insecticidal tree fogging of Carapa grandiflora (Meliaceae), Th. Wagner (ZFMK). — Kenya: 8 ex., Kakamega Forest near Buyangu, 0.18N/34,53E, 1X.2001, X.2002, some by insecticidal tree fogging of Teclea nobilis, Th. Wagner (ZFMK). — Rwanda: | ex., Gisakura,2.00S/29.50E, V1.1972, coll. Roggeman (CBe). — Tanzania: 14 ex., Ukerewe Island, 2.098/32,52E, ex. coll. Conrads (NMK). — Uganda: 2 ex., Kam- pala, 0.19N/32.35E, V1.1940, A. F. J. Gedye (BMNH); | ex., Kibale Forest, 0.50N/31.06E, 1983, M. Nummelin (MZHF); 4 ex., same label as holotype, insecticidal fogging of Trichilia rubescens (Meli- aceae) and Cynometra alexandri (Caesalpiniaceae) (ZFMK); 2 ex., Semliki Forest, 0.48N/30.08E, 11.1997, Th. Wagner (ZFMK). Key to species Afrotropical Monolepta-species, which are charac- terized by a black transverse band at the elytral base, a further transverse band or isolated black spots in the apical third of elytra, where other parts of elytra are yel- low or yellowish-red, can be distinguished after the fol- lowing key. It can be used for specimens up to 5.60 mm total length. Larger specimens (up to 6.70 mm) with the colour pattern mentioned above belong to M. quadri- zonata Laboissiere, 1940, which was revised elsewhere (WAGNER 2003b). Excluded from this key are also Monolepta-species which have a mid-elytral transverse black band, black outer elytral margins and often a red elytral suture like the abundant and widely distributed M. elegans Chevrolat 1837, M. vinosa Gerstaecker, 1871, M. cruciata Guerin-Meneville, 1858, and some other species, which will be revised in the near future. l Pronotum very broad (prontal length to width: 0.54-0.58; Fig. 7); second antennomere particu- larly in males very short (length of second to third antennomere: 1.00-1.40, length of third to fourth antennomere: 0.20-0.36; Fig. 8); small (total length: 3.00-4.20 mm); spermatheca of peculiar shape (Fig. 9); apical half of median lobe very slender (Fig. 11); known from savannas of West Africa towards north-eastern Congo ..................000.. DM ee skanensenstene Monolepta buquetii Chevrolat, 1837 — Pronotum less broad (prontal length to width: 0.58—0.67); second antennomere more elongated (length of second to third antennomere: 0.75—1.10, length of third to fourth antennomere: 0.28-0.62); often larger than 4.20 mm; spermatheca of differ- ent shape; median lobe less narrowed in the apical half, exceptions are M. ronbeeneni sp. n. and very few specimens of M. vincta, which differ in endo- puallicrarmatune (Biss 25, Sl sees sseceeeses 2 D Basal antennomeres elongated and slender, second antennomere of same length or longer than third (length of second to third antennomere: 1.00-1.10, Figs 28, 39); elytra usually with large isolated spot n in the apical third, instead of a transverse band, other parts of elytra more reddish-yellow (Figs 27, 38): re- stricted to Guineo-Congolian forests in northern Central and western East Africa (Figs 17, 32) ......... 3 Basal antennomeres less elongated, second anjen- nomere same length or shorter than third (length of second to third antennomere: 0.75-1.00, Figs 2, 13, 19, 34); elytra either with smaller isolated spot or transverse band in the apical third (Figs 1, 12, 16, 33), other parts pale yellow to yellow; distrib- uted throughout tropical Africa; check of male genitalic structures is necessary in most specimens to ensure a correct identification, single females can sometimes not be allocated to species without Very small (total length: 3.10-3.60 mm); elytra slender (width of both elytra to length of elytron: 0.59-0.64, Fig. 27); third antennomere very long (length of third to fourth antennomere: 0.41-0.62, Fig. 28); Central African Republic towards western Uganda and Rwanda (Fig. 17)...............................- tio Monolepta ronbeeneni sp. n. Larger (total length: 3.60-4.80 mm); elytra broad (width of both elytra to length of elytron: 0.71 0.75, Fig. 38); third antennomere less elongated (length of third to fourth antennomere: 0.37-0.43): known from north-eastern Congo towards western Kenya (Kakamega Forest), Rwanda and north- western Tanzania (Ukerewe Island)......................... AO Monolepta naumanni sp. n. Third antennomere significantly longer than second (length of second to third antennomere: 0.75—0.88, Fig. 13), and about half as long as fourth antenno- mere (length of third to fourth antennomere: 0.46- 0.54); pronotum comparatively slender (prontal length to width: 0.63-0.67); known from Eritrea, northern Somalia, Arabian Peninsula, Israel and Jordan (Eis I 2 ee RE O OEA Monolepta lepida Reiche, 1858 Third antennomere shorter (length of second to third antennomere: 0.86-1.00: length of third to fourth antennomere: 0.30-0.50): pronotum broader (prontal length to width: 0.58-0.65): tropical At- A ener A 5 On average larger (total length: 3.80-5.60 mm): median lobe homogeneously conical and slightly widened at apex (Fig. 5): north-eastern Congo and Uganda towards the Cape, most abundant and widely distributed in southern Africa (Fig. 6): specimens which occur syntopely with M. vincta are significantely larger than those........................... ee Monolepta melanogaster (Wiedemann, 1823) 282 Bonner zoologische Beitráge 53 (2004) — On average smaller (total length: 3.25-4.75 mm); median lobe narrowed and parallel-sided at apex (Figs 23, 24, 37); throughout tropical Africa but not known from the Republic of South Africa, and rare in other countzries of southern Africa ............ 6 6 Third antennomere more elongated (length of third to fourth antennomere: 0.43-0.50, Fig. 34); trans- verse post antennal suture deeply incised, prono- tum pale yellow (Fig. 33); lateral endophallic spi- culae strong, curved inwards, hooked at apex (Fig. 37); Nigeria through Congo basin towards western Tanzania, Zambia......... Monolepta sharonae sp. n. — Third antennomere shorter (length of third to fourth antennomere: 0.28—0.37, Fig. 19); transverse post antennal suture usually not incised, pronotum often red, rarely black; endophallic armature different (Figs 23, 24, 25); throughout tropical Africa with exception of South Africa Postal Monolepta vincta Gerstaecker, 1871 REFERENCES CHAPUIS, F. (1879): Phytophages Abyssiniens du musée civique d histoire naturelle de Génes. Annali di Museo Civico Storia Naturale Genova 15: 5-31. CHEVROLAT, L. A. A. (1837): P. 407 Monolepta, in: DE- JEAN (ed.) Catalogue des Coléopteres de la collection de M. le Compte Dejean. 3rd. edition, revue, corrigee et augmentee, fasc. 5. Paris. GERSTAECKER, A. (1871): Beitrag zur Insektenfauna von Zanzibar. Archiv fiir Naturgeschichte 37(1): 42-86. JOANNIS, L. de (1866): Gallerucides, tribu de la famille des Phytophages, ou Chrysomelines. L’Abeille — Mé- moires d’Entomologie 3: 1-168. LABOISSIERE, V. (1920a): Diagnoses de Galerucini nou- veaux d’Afrique (Col. Chrysomelidae). Bulletin de la Société entomologique de France 1920: 50-53. LABOISSIERE, V. (1920b): Diagnoses de Galerucini nou- veaux d’Afrique (Col. Chrysomelidae). Bulletin de la Société entomologique de France 1920: 98-101. LABOISSIERE, V. (1940): Galerucinae. Exploration du Parc National Albert 31: 1-98. Institut des Parcs Nationaux du Congo Belge, Bruxelles. REICHE, L. (1858): Especes nouvelles ou peu connues de Coléopteres, recueilles par M. F. de Saulcy. Annales de la Société entomologique de France 6: 1-60. WAGNER, Th. (2000a): New Monolepta species (Coleop- tera: Chrysomelidae) from Eastern Africa. Entomolo- gische Zeitschrift 110: 34-40. WAGNER, Th. (2000b): Revision of afrotropical Monolepta species (Coleoptera, Chrysomelidae, Galerucinae). Part I: Species with red and black coloured elytra, pronotum and head, with description of new species. Entomologische Zeitschrift 110: 226-237. WAGNER, Th. (2001): Revision of Afrotropical Monolepta Chevrolat, 1837 (Coleoptera: Chrysomelidae, Galerucinae). Part Il: Species with red elytra, prono- tum and elytra, with descriptions of new species. Bon- ner Zoologische Beiträge 50: 49-65. WAGNER, Th. (2002): Revision of Afrotropical Monolepta species (Coleoptera, Chrysomelidae, Galerucinae). Part III: Species with red elytra and yellow prothorax, including description of new species. Deutsche Ento- mologische Zeitschrift 49: 27-45. WAGNER, Th. (2003a): Present status of a taxonomic revi- sion of afrotropical Monolepta and related groups (Galerucinae). Pp. 133-146 in: FURTH, D. G. (ed.) Special Topics in Leaf Beetle Biology. Proceedings V. International Symposium on the Chrysomelidae, Foz do Iguacu 2000. Pensoft. WAGNER, Th. (2003b): Revision of afrotropical Monolepta Chevrolat, 1837 (Coleoptera, Chrysomelidae, Galerucinae). — Part IV: Species with red head and thorax and black elytra or black elytra with red apex, with description of new species. Annales Sciences Zo- ologiques, Miscellanea 49: 37-89. WAGNER, Th. (2004): Phylogeny of Afrotropical Mono- lepta and related taxa (Galerucinae). Pp. 75-84 in: JOLIVET, P., SANTIAGO-BLAY, J. A. & M. SCHMITT (eds) New Developments on the Biology of Chry- somelidae. Academic Publishing bv. WEISE, J. (1903): Afrikanische Chrysomeliden. Archiv für Naturgeschichte 69: 197-226. WEISE, J. (1909): 7. Coleoptera, 12. Chrysomelidae und Coccinellidae. Pp. 153-267 in: SJOSTEDT, Y. (ed.) Wissenschaftliche Ergebnisse der schwedischen zo- ologischen Expedition nach dem Kilimandscharo, dem Meru und den umgebenden Massaisteppen Deutsch- Ostafrikas 1905-1906. Palmquist Aktiebolag, Stock- holm. WIEDEMANN, C. R. W. (1823): Zweihundert neue Káfer von Java, Bengalen und dem Vorgebirge der guten Hoffnung. Zoologisches Magazin 2(1): 3-153. WILCOX, J. A. (1973): Chrysomelidae: Galerucinae: Luperini: Luperina. Pp. 433-664 in: JUNK, W. (ed.) Coleopterorum Catalogus Suppl. 78 (3). 's-Gravenhage, Junk. Author’s address: Dr. Thomas WAGNER, Universitat Koblenz-Landau, Institut fiir Integrierte Naturwissen- schaften — Biologie, Universitátsstr. 1, D-56070 Ko- blenz; e-mail: thwagner@uni-koblenz.de Bonner zoologische Beitráge Band 53 (2004) Heft 1/2 Seiten 283-289 Bonn, Juni ’ Universität Koblenz-Landau, Institut für Ingegrierte Naturwissenschaften Diversity Patterns of Plants and Phytophagous Beetles in Sub-Saharan Afric: Wolfgang KÚPER”, Thomas WAGNER” & Wilhelm BARTHLOTT” 2 Universität Bonn, NEES Institute for Biodiversity of Plants, Bonn, Germany Abstract. Species distribution of flowering plants (Angiospermae) and the phytophagous beetle group Monolepta (Chrysomelidae) in Sub-Saharan Africa (south of 17° N) are compared based on species numbers per square degree grid. The beetle data comprise all 89 valid species for Africa (21,000 specimens of Afrotropical Monolepta have been currently revised). The plant diversity data are based on 6,269 species with some 330,000 distributional records (10-15 % of all African angiosperm species). Shared centres of species richness of both taxa are geodiverse, montane forests, namely the Albertine Rift, Eastern Arc Mountains, isolated East African volcanoes, montane areas in Cameroon and northeastern parts of a the Republic of South Africa. However, the Cape and the Upper Guinea Region show diverging patterns: plant species richness is higher than the richness of Chrysomelidae. Actual diversity patterns versus sampling artefacts are discussed. Certain mechanisms, like allopatric speciation processes, contribute to a diverse flora and fauna in areas of high geodiversity. Thus, if there is a general and taxon-independent positive relationship between geodiver- 200 Biologie, Koblenz, Germany sity and species richness, these areas are of explicit value for the conservation of terrestrial biodiversity. Key words. Angiosperms, Chrysomelidae, biodiversity, biogeography, distribution patterns, conservation 1. Introduction The understanding of the spatial distribution of biodi- versity is a fundamental requirement for its conservation and sustainable use. However, current biogeographic knowledge is restricted to a tiny proportion of terrestrial biodiversity. While there is comparatively detailed in- formation on the global distribution of vertebrate diver- sity (RODRIGUEZ et al. 2004), there is only little infor- mation about the spatial distribution of the remaining 99 % of animal species, particularly for such a megadi- verse group like insects. More problematic, the repre- sentativeness of patterns of vertebrate diversity, e.g. pat- terns of insects, might be limited. In contrast, plants in their function as the foundation of food-webs may serve as a surrogate to assess patterns of overall terrestrial biodiversity (BARTHLOTT et al. 1996, 1999). Flowering plants and beetles, in particular the most spe- ciose phytophagous beetle taxa, Chrysomeloidea and Curculionoidea, represent one of the oldest and largest radiation of plant-animal co-evolution (FARRELL 1998). The diversity of flowering plants (BARTHLOTT et al. 1996, 1999) and the extent to which phytophagous bee- tles are specialized on their food-plants, were the basis for estimations which dramatically changed our idea of terrestrial biodiversity (ERWIN 1982). Despite the ex- tend of specialisation of phytophagous beetles in tropi- cal biomes and that numbers have been clearly overes- timated on the basis of conclusions from temperate I In commemoration of Clas Michael Naumann zu Königsbrück (26.06.1939 — 15.02.2004) regions (MAY 1990; VDEGAARD 2000; WAGNER 2000a; NOVOTNY et al. 2002), there is no doubt that the number of existing species is several times higher than the num- ber of yet described species. This is particularly true for the approximately 150,000 species of leaf beetles and weevils (LAWRENCE & BRITTON 1991). Both beetle groups illustrate the most prominent prob- lem of biodiversity research. The more speciose taxa are and the smaller the mean range size of their species is, the less complete is our knowledge on their large scale biogeographic patterns. While the global distribution patterns; e.g. of the vertebrates, which have on average a large range size, are apparently known on a 0.5° de- gree resolution (RODRIGUES et al. 2004), the situation for plants (SCHATZ 2002) and particularly arthropods is completely different. While plants are at least relatively well known from a taxonomic perspective, there is a strong need for taxonomic research on arthropods. This includes the phytophagous beetles, which have an key ecological position in the food-web as the most species rich group of first order consumers. Thus, chrysomelids provide a good example for studying speciation proc- esses, including co-evolution of angiosperms and their consumers in tropical systems. Since a detailed assessment of arthropod diversity pat- terns at the global scale is impossible, we rely on data for selected groups and selected areas. Based on the most comprehensive databases on the Sub-Saharan dis- tributions of plants and the chrysomelid group of Monolepta, we exemplarily compare centres of species richness of plants and phytophagous beetles. We discuss 284 consequences of the quality of available biogeographic data for our knowledge on the biodiversity of both groups and for potential application of these data for conservation biology. 2. DATA AND METHODS 2.1. Taxonomy and diversity data for Monolepta (Chrysomelidae) Afrotropical species of Monolepta Chevrolat, 1837 were recently revised (WAGNER 2003a). These beetles form the largest group of the Galerucinae (Chrysomelidae), having worldwide about 600 nominal species. Many of the 160 species described from Africa were found to be synonyms or needed to be transferred to other genera due to their phylogenetic position. However, 50 valid species remained in Monolepta, and there is about the same number of new species, most of them presently described (WAGNER 2000b, 2001b, 2002, 2003b; Wag- ner this issue) or awaiting description. Unfortunately, food-plants of most Monolepta species are still un- known. There are only very few data available from la- bels of dried specimens, with direct observations in the field and literature (JOLIVET & HAWKESWOOD 1995) in- dicating that some species feed on Citrus sp. and some others on other Rutaceae. The revision of the 89 valid species of Afrotropical Monolepta is based on 21,000 specimens. Five hundred specimens are recently col- lected mainly in eastern Africa, while most are dried specimens from all major collections housing African insects, principally museums in Berlin, Brussels, Lon- don, Nairobi, Paris, Pretoria, Tervuren and Windhoek. After this comprehensive revision of material, detailed data on the distribution of Afrotropical Monolepta spe- cies are now available. Only four Monolepta species are distributed throughout tropical Africa, and known from a variety of biomes such as tropical forests, savannas and even deserts, from coastal regions to montane areas. All other species show more restricted geographical and ecological distributions. Forest-dwelling species are of- ten more restricted and a high degree of endemism is found amongst species from montane areas (WAGNER 2001a). Coordinates are derived from label data on dis- tribution using a gazetteer compiled by Ugo Dall’ Asta (Africa-Museum, Tervuren) for locations from Congo (Zaire) and generally using the Alexandria Digital Li- brary Gazetteer Server. Data of all Monolepta species is based on 2320 localities in subsaharan Africa. 2.2. Diversity data for Plants Earlier versions of the plant dataset have been described and analysed in previous publications (LINDER 2001; LOVETT et al. 2000; LINDER et al. 2005; KUPER et al. 2004; Lovett et al. 2004). Since 2003, an international Bonner zoologische Beitráge 53 (2004) group of research institutions has contributed data on the distribution of African plants to the Biogeographic Information System on African Plant Diversity, which is hosted and curated by the BIOMAPS Project within the BIOLOG-BIOTA framework (www.biota-africa.org). The spatial precision of the distribution data varies be- tween exact localities mainly from herbarium collec- tions with georeferenced localities, to 1? resolution data from digitised maps. Additional information on the ori- gin of this dataset is documented in BURGESS et al. (in press, see also Acknowledgements). The current database covers African-wide distribution records for 6,269 species (status as of March 2004), all of which had been taxonomically revised. This com- prises between 10 and 15 % of the African angiosperm species (LEBRUN & STORK 1991-1997; BEENTIE et al. 1994). There are currently 330,000 distribution records in the form of confirmed collection localities available. 2.3. Data preprocessing and analysis Data are organized in MS Access databases and have been plotted and analysed using ArcView 3.2a GIS software. In order to achieve maximum comparability with previous analyses on Sub-Saharan diversity (BALMFORD et al. 2001; BROOKS et al. 2001; BURGESS et al. 2002, in press; DE KLERK et al. 2004), all distribu- tion data were rescaled to a 1° grid resolution within a base map of 1,713 one-degree latitude-longitude grid cells covering mainland Sub-Saharan Africa south of 17° North. By restricting the geographic coverage to Af- rica south of the Sahara and excluding those species only found on offshore islands, a database with 5,985 plant species and 89 species of Monolepta remains for further analyses. For the scatterplot, we only used those 580 grid cells in which at least one species of both groups occurred. 3. RESULTS 3.1. Shared centres of diversity of plants and Monolepta There is an overlap of centres of high plant and Monolepta diversity in montane areas, namely the Al- bertine Rift system including the Virunga Volcanoes, the Ruwenzori, Bwindi Impenetrable Forest and the Ka- huzi Biega National Park at the south-western extreme; the Eastern Arc, in particular East Usambara; the iso- lated East African Mountains such as Mt. Elgon, Mt. Kenya and Mt. Kilimanjaro; southern Katanga (Mi- tumba and Kundelungu Mountains, major collections in Upemba/ Kundelungu National Park and close to Lubumbashi); and to a lesser extend afromontane re- gions in Ethiopia, the northeastern parts of the Republic of South Africa, and lower Guinea (Fig. la/b). Wolfgang KUPER, Thomas WAGNER & Wilhelm BARTHLOTT: Diversity Patterns of Plants and Phytophagous Bectles 285 Fig. 1: Species richness per one-degree grid cell in Sub-Saharan Africa south of 18th degree latitude. — A. plants, n (species) 5985, B. Monolepta, n (species) = 89). Richness values are rescaled to percentages of the maximum richness per grid of each taxon (Monolepta: max = 20; for plants: max = 682). The scale ranges from high species richness (dark red) to low species rich- ness (bright yellow). In grid cells with no present species, the grey background colour represents elevation above sea level (dark grey: high elevation, bright grey: low elevation). Black foreground lines indicate national boundaries. 0 Garamba NP 0 Virunga ENZO gUsambara SS O1 Kampala area ® Mt. Elgon Upemba NP ee NY Kahuzi Biega NP ubumbashi area 0 000 9 0 0 e... 0 0 0.00 00 0090 000 00 O 00 0 0 oe 0600 00 ® Abidjan area 00 0. 940090 @ 00 0 WD 0 00 0000 0000 “m BER CM OCS 00@0 8 © Ge A 00 @ @ 000 00 MIA 69 0 0 W of Douala 0 0 m. © BEN ae .0 o Western Cape Western cape 400 0 Mt. Cameroon al Mt. Nimba 0 # of species of Monolepta O I I # of plant species Fig. 2 : Species richness of Monolepta and plants in 580 one-degree grid cells in Sub-Saharan Africa. Only grids where both taxa were present with at least one species are considered. Outliers were labelled according to their geographic position (NP: National Park). 286 3.2. Divergent diversity patterns of plants and Monolepta Even though some centres of species richness of both groups do coincide, the correlation between the patterns of species richness of plants and Monolepta are very low (Fig. 2). The correlation can probably be explained merely by the fact that the majority of the Monolepta species occur in forest habitats, and that forests are comparatively phytodiverse. The most obvious differ- ence between the diversity patterns of plants and beetles is the outstanding plant diversity of the Cape Floristic Kingdom. This observation is contrary to all animal taxa yet mapped such as vertebrates (BROOKS et al. 2001) but apparently also insects (GILIOMEE 2003). There is no doubt that this pattern 1s based on ecological and his- torical characteristics of this landscape, which seem to favour certain plant taxa more strongly than any animal taxon. The same is true for the Succulent Karroo. Few grids, all in regions with savannah habitats, have an exceptionally high number of Monolepta species, but comparatively low number of plant species. These are Garamba National Park in north-eastern Congo, Upemba National Park in south-eastern Congo (Ka- tanga), the region west of Katanga along the Lulua River and the region around Kampala in Uganda. These areas belong to the best-collected regions for these bee- tles (and generally for insect groups). Some extended expeditions have been carried out in national parks in the former Belgian Congo over the years, and about 18 % of all known specimens of Afrotropical Monolepta are from Garamba National Park alone. Hence, high di- versity in these particular grids is biased by high sam- pling effort. The Upper Guinea region, characterized by high plant diversity, has low beetle diversity. Since West Africa is less well sampled for Monolepta than all other African regions, this 1s most likely also a sampling artefact. However, this region lacks high mountains as typical for the rift systems in Central and East Africa, and Monolepta may be generally less diverse in West Africa. Due to the strong influence of sampling intensity on the Monolepta dataset, there are no significant correlations of the derived species richness patterns of these beetles (or the residuals between the species richness of Monolepta and plants) with abiotic factors such as hy- drothermic parameters (e.g. waterbalance, number of dry months), and topodiversity. Correlations between species richness of Monolepta and either plants or abiotic parameters did not improve when tested on the larger spatial scale. Bonner zoologische Beitráge 53 (2004) 4. DISCUSSION 4.1. Afrotropical Mountains: centres of biodiversity On a global scale, many centres of plant diversity are located in tropical montane areas such as the South American Andes, the Eastern Arc, Crystal Mountains, Papua New Guinea, Himalaya or the Mesoamerican Cordillera Central (BARTHLOTT et al. 1999; MUTKE & BARTHLOTT 2005). In Africa, the montane regions Al- bertine Rift, the Eastern Arc, the Southern Rift extend- ing into Chimanimani-Nyanga, the northeastern part of South Africa extending from Sekhukhuneland and Soutpansberg into the Maputaland Centre, and the West African Mt. Cameroon are well known centres of rich- ness and endemism for a variety of ecologically very different taxa such as birds, mammals, snakes, amphibi- ans (BROOKS et al. 2001) and plants (MUTKE et al. 2001). Our study gives evidence that this also is the case for phytophagous beetles. Most Monolepta species in Cen- tral and East Africa have a restricted distribution. Twelve species are endemic to montane forests along the Albertine Rift, ten are restricted to East African mountains in Kenya and Tanzania, mainly to Mt. Kenya, Mt. Kilimanjaro and to the Eastern Arc Moun- tain mosaic, and a further nine species are restricted to montane areas in Ethiopia and Eritrea (WAGNER 2001a). To a lesser extent, montane regions in Cameroon also have higher species diversity and some endemic species. The diversification of many Monolepta species is obvi- ously strongly effected by geographical speciation in isolated montane forests. Many of the forest-dwelling Monolepta species appear to be young species in evolutionary terms. The Quater- nary extension of the forest biome may be crucial in ex- plaining the distribution pattern. In the late Pleistocene, the climate, particularly in central and East Africa was much cooler and drier (HAMILTON 1981). The timber- line was about 1000 m lower than at present (e.g. BON- NEFILLE et al. 1990; LOVETT 1993) and most lowland areas, including the Congo Basin, were too dry for for- est vegetation and were covered by savannahs. The pro- posed forest refuge core areas coincides well with the highest degree of endemism and diversity of Monolepta in montane areas of Cameroon, the Albertine Rift and higher mountains in Kenya and Tanzania, including the Eastern Arc Mountain mosaic. Other Monolepta spe- cies, which are restricted to dry forest and savannahs, have a wider range. In the early Holocene, the climate became warmer and more humid. Wet tropical forests expanded to a presumed maximum about 6,000 years ago (HAMILTON 1981). During that time the Sudano- Zambesian savannah zone was presumably separated into a north-western and a south-eastern part, leading to a corresponding distribution pattern also found in other Wolfgang KUPER, Thomas WAGNER & Wilhelm BARTHLOTT: Diversity Patterns of Plants and Phytophagous Bectles 2 insects like butterflies and paussid beetles (CARCASSON 1964; NAGEL 1987). Combined analyses on large scale biodiversity, in par- ticular those comparing such different taxa like beetles and plants, have been heavily criticised as paradoxical, since they tend to neglect the diversity of ecological characteristics at the species level (HUSTON 2001). In- deed, the influence of abiotic factors or historical proc- esses depends on the characteristics of each taxon. For example, steep topography or climatic fluctuations can lead to reduced gene flow between populations, but this strongly depends on the mobility of the animals or seed vagility in plants. The high degree of endemism in the Cape flora is partly interpreted as a consequence of the low dispersability of a high proportion of the species; e.g. of about 1000 ant-dispersed species in the Fa- baceae, Proteaceae, Restionaceae, Rhamnaceae, and Ru- taceae (GOLDBLATT & MANNING 2002). For the most speciose and endemic plant taxon in the South Ameri- can Andes, the Orchidaceae, it is not dispersal but lim- ited mobility of highly specialized arthropod pollinators that may be a key to understanding the extremely high number of local endemic species (KUPER et al. 2004). Nevertheless, if considering biodiversity not exclusively a result of the spatial distribution of abiotic factors (pat- tern diversity) but as a result of a variety of mechanisms (process diversity), it is clear that geodiverse, montane areas are an ideal place where a high variety of mecha- nisms in parallel can promote high levels of diversity: the steep orography can promote allopatric speciation processes, whereas orographic rainfall and horizontal mist interception in mountains may facilitate survival of species during long-term climate dynamics (FJELDSA & LOVETT 1997) and opportunistic colonialisation by widespread species. Habitat suitability but also habitat diversity as a consequence of geodiversity, provide suit- able conditions for a speciose and structurally rich vege- tation which in turn promotes consumer diversity due to diverse habitats and a high net primary production. The afromontane areas, the Cape and parts of the West African upper Guinea Centre of plant diversity, cover more than 80 % of the Sub-Saharan flora on less than 10 % of its total area (KUPER et al. 2004). Preliminary analyses suggest that the proportion of the Sub-Saharan vertebrate fauna covered by these areas might be even higher than that for plants (BROOKS et al. 2001), and the situation for phytophagous insects may be similar. Due to the provided habitat diversity, one explanation for species richness of the Arotropical mountains might be the high number of overlapping ranges contributed by species of taxa which have their actual centre of diversity at different places. However, a significant number of endemic species and species with small dis- tribution ranges of different taxa are restricted to Afro- tropical mountains. The occurrence of similar diversity stimulating mechanisms in very different taxa highlights the importance of Afrotropical mountains as overall centres of terrestrial biodiversity. 4.2. Consequences of fragmentary biodiversity information J Our plant data are the most comprehensive ever assem- bled for the study area, but inevitably have limitations. There are certain areas in Ethiopia, the Sudan, the Cen- tral African Republic, the Republic of the Congo, the Democratic Republic of the Congo and Angola, whose plant diversity is not adequately documented (KUPER, unpubl. data), and the scientific exploration of these ar- eas is a most important challenge for the future. Unsur- prisingly, the situation is even more problematic for phytophagous beetles. While the low diversity in Upper Guinea may be partly explained by the relative scarcity of montane areas; e.g., the Crystal Mountains in Gabon and Equatorial Guinea, are clearly undersampled. The concentration of the present records to urban areas or such with a comparatively good infrastructure (e.g. Ad- dis Ababa, Mbandaka, and several National Parks) is symptomatic for sampling artefacts. Currently it is not possible to quantify how much of the congruence in richness patterns of beetles and plants in our database is due to the fact that the same areas tend to be better col- lected for both taxa, resulting in a comparatively high number of species of both groups in these cells. The more speciose taxa are, and the smaller the mean range size of their species is, the less complete is our knowledge on their large scale biogeographic patterns. At the same time, the restricted knowledge concerning the distribution of any taxon limits its consideration in spatial priority setting approaches for nature conservation. Current conservation approaches in Africa are mainly based on patterns of vertebrate diversity (BURGESS et al. 2004; FJELDSA et al. 2004; DE KLERK et al. 2004). This may mainly result from their charisma, making them at- tractive for funding as well as for non-scientific but valuable field survey by lay people, and also from the history of Africa’s protected areas (FJELDSA et al. 2004). Whereas about 95 % of the vertebrates in Sub- Saharan Africa are covered by the existing set of IUCN protected areas, the proportion for plants covered by the same set is at 74 % (BURGESS et al. in press). Only in a few cases are large scale patterns of plant diversity more appropriately considered, e.g. for the Republic of South Africa (COWLING et al. 2003). Though an amount of about 140 million plant specimens have been collected globally (BGCI 6/2004), estimations on the basis of the TROPICOS database at the Missouri Botanical Garden revealed that for about 80 % of all taxa which are repre- sented in the form of collections, there are less than ten collection localities worldwide (SCHATZ 2002). 288 Bonner zoologische Beitráge 53 (2004) A large attraction of currently available numerical algo- rithms for selecting conservation areas is that they can go beyond subjective preferences for the cute and the cuddly, but if biogeographic data and phylogenies are available only for “attractive” groups, we are no further forward (MACE et al. 2003). For arthropods, most likely less than 20 % of the existing species have been de- scribed so far. The risk of this situation is that we are actually considering only the tip of the iceberg of prob- lems in nature conservation. The bad message from our study is that even after spending several ten thousand hours of manpower on the collection of the distribution of African plant species and checking label data of about 21,000 beetle speci- mens, we are far from a satisfying documentation and understanding of the large-scale diversity patterns of these taxa. The good news might be, if such different taxa like ver- tebrates, plants and phytophagous beetles do have cen- tres of biodiversity in common, it may be possible to protect a large proportion of species, in relatively well defined and small areas. These areas possibly comprise species that are not yet described. Unfortunately, in many cases exactly these areas are also most attractive for human settling (CINCOTTA et al. 2000; BALMFORD et al. 2001). Hence, more comprehensive evidence show- ing taxon-crossing relevance of these areas could at least be a solid argument in favour of the sustainable use and protection of its biodiversity. Ackowledgements. We thank the numerous experts who have assisted to compile the plant and beetle data used in this paper. The establishment and co-ordination of the Bio- geographic Information System on African Plant Diversity at the Nees Institute for the Biodiversity of Plants in Bonn is funded by the German Federal Ministry of Education and Research (BMBF-BIOLOG Programme) and the Akademie der Wissenschaften und Literatur, Mainz. We thank Henk Beentje (Kew), Cyrille Chatelain and Laurent Gautier (Geneva), Tony Rebelo (Kirstenbosch), Jerome Degreef (Meise), Peter Frankenberg (Stuttgart), Norbert Júrgens (Hamburg), Don Kirkup (Kew), Sigrid Liede (Bayreuth), Peter Linder (Zúrich), Roger Polhill (Kew), Adjima Thiombiano (Ouagadougou), Mauricio Velayos and Fernando Casas (Madrid), Jan Wieringa and Mare So- sef (Wageningen) and Georg Zizka (Frankfurt) for valuable contributions to the plant distribution dataset. We thank Henning Sommer for fruitful discussions and Jens Mutke for his efforts to establish and maintain the BIOMAPS working group in Bonn. Juliane Dame assisted with the manuscript. REFERENCES BALMFORD, A., MOORE, J. L., BROOKS, T., BURGESS, N., HANSEN, L. A., WILLIAMS, P. & RAHBEK, C. (2001): Integrating biodiversity priorities with ‘conflicting socio-economic values in the Guinean-Congolian for- est region. Science 291: 2616-2619. BARTHLOTT, W., LAUER, W. & PLACKE, A. (1996): Global distribution of species diversity in vascular plants: to- wards a world map of phytodiversity. Erdkunde 50: 317-327. BARTHLOTT, W., BIEDINGER, N., BRAUN, G., FEIG, F., KIER, G. MUTKE, J. (1999): Therminological and Methodo- logical Aspects of the Mapping and Analysis of the Global Biodiversity. Acta Botanica Fennica 162: 103— 110. BEENTIJE, H. J., ADAMS, B., DAVIS, S. D. & HAMILTON, A. C. (1994): Regional Overview: Africa. Pp. 101-148 in: DAVIS, S. D., HEYWOOD, V. H. & A. C. HAMILTON (eds.) Centres of Plant Diversity. A guide and strategy for their conservation. Vol. 1: Europe, Africa and the Middle East. IUCN Publications Unit, Cambridge, UK. BONNEFILLE, R., ROELAND, J. C. & GUIOT, J. (1990): Tem- perature and rainfall estimates for the past 40,000 years in equatorial Africa. Nature 346: 347-349. Brooks, T., BALMFORD, A., BURGESS, N., FJELSDA, J., HANSEN, L. A., MOORE, J., RAHBEK, C. & WILLIAMS, P. (2001): Toward a Blueprint for Conservation in Af- rica. BioScience 51: 613-624. BURGESS, N., RAHBEK, C., LARSEN, F. W., WILLIAMS, P. & BALMFORD, A. (2002): How much of the vertebrate di- versity of Sub-Saharan Africa is catered or by recent conservation proposals? Biological Conservation 107: 327-339, BURGESS, N. D., BROWN, J., WESTAWAY, S., TURPIE, S., MESHACK, C., TAPLIN, J., MCCLEAN, C., KUPER, W., MUTKE, J. & LOVETT, J. C.: Major gaps in the distribu- tion of protected areas for threatened and narrow range Atrotropical plants. Biodiversity & Conservation (in press). CARCASSON, R. H. 1964. A preliminary survey of the zo- ogeography of African butterflies. East African Wild- life Journal 2: 122-157. CINCOTTA, R., WISNEWSKI, J. & ENGELMAN, R. (2000): Human population in the biodiversity hotspots. Nature 404: 990-992. COWLING, R. M., PRESSEY R. L., ROUGET M. & LOMBARD, A. (2003): A conservation plan for a global biodiver- sity hotspot — The Cape Floristic Region, South Af- rica. Biological Conservation 112: 191-216. DE KLERK, H. M., FJELDSA, J., BLYTH, S. & BURGESS, N. D. (2004): Gaps in the protected area network for threat- ened Afrotropical birds. Biological Conservation 117: 529-537. ERWIN, T. L. (1982): Tropical forests: Their richness in Coleoptera and other arthropod species. Coleopterists Bulletin 36: 74-75. FARRELL, B. D. (1998): “Inordinate fondness” explained: Why are there so many beetles? Science 281: 555— 339. FJELDSA, J. & LOVETT, J. C. (1997): Geographical patterns of young and old species in African forest biota: the significance of specific montane areas as evolutionary centres. Special Issue: Biodiversity and Conservation 6: 325-347. FJELDSA J., BURGESS, N. D. & DE KLERK, H. M. (2004): Will Africa's protected areas network conserve Af- rica’s threatened mammals? Oryx 38: 17-25. Wolfgang KUPER, Thomas WAGNER & Wilhelm BARTHLOTT: Diversity Patterns of Plants and Phytophagous Beetles 289 GILIOMEE, J. H. (2003): Insect Diversity in the Cape Floris- tic Region. African Journal of Ecology 41: 3. GOLDBLATT, P. & MANNING, J. C. (2002): Plant diversity of the Cape region of Southern Africa. Annals of the Missouri Botanical Garden 89: 281-302. HAMILTON, A. C. (1981): The quaternary history of Afri- can forests: its relevance for conservation. Journal of African Ecology 19: 1-6. HUSTON, M. (2001): People and Biodiversity in Africa. Science 293: 1591-1592. JOLIVET, P. & HAWKESWOOD, T. J. (1995): Host-plants of Chrysomelidae of the world. Backhuys, Leiden. KUPER, W., SOMMER, J. H., LOVETT, J. C., MUTKE, J., LINDER, H., BEENTJE, H., VAN ROMPAEY, R. A. S. R., CHATELAIN, C., SOSEF, M. & BARTHLOTT, W. (2004): Africa's hotspots of biodiversity redefined. Annals of the Missouri Botanical Garden 91: 525-536. LAWRENCE, J. F. & BRITTON, E. B. (1991): Coleoptera. Pp. 543-683 in: CSIRO, Division of Entomology (ed.) The Insects of Australia. University College of Lon- don Press, London. LEBRUN, J.-P. & STORK, A. (1991-1997): Enumeration des plantes a fleurs d'Afrique tropicale. 4 Vols. Editions des Conservatoire et Jardin botaniques de la Ville de Geneve, Geneva, Switzerland. LINDER, H. P. (2001): Plant diversity in Sub-Saharan tropi- cal Africa. Journal of Biogeography 28: 169-182. LINDER, H. P., LOVETT, J. C., MUTKE, J., BARTHLOTT, W., JURGENS, N., REBELO, T. & KUPER, W. (2005): A nu- merical re-evaluation of the Sub-Saharan phytochoria of mainland Africa. Biologiske Skripter 55: 229-252. LOVETT, J. C. (1993). Climatic history and forest distribu- tion in eastern Africa. Pp. 23-29 in: Lovett, J. C. & WASSER. S. K. (eds.) Biogeography and ecology of the rain forest of Eastern Africa. Cambridge University Press, Cambridge. LOVETT, J. C., RUDD, S., TAPLIN, J. & FRIMODT-MOLLER, C. (2000): Patterns of plant diversity in Africa south of the Sahara and their implications for conservation management. Biodiversity and Conservation 9: 33-42. LOVETT, J. C., MARCHANT, R., TAPLIN, J. & KUPER, W. (2004): The oldest rainforests in Africa: stability or re- silience for survival and diversity? Pp. 198-229 in: PURVIS, A., GITTLEMAN, G., BROOKS, T. (eds.) Phy- logeny and Conservation. Cambridge University Press, Cambrigde. MACE, G. M., GITTLEMAN, J.L. & PURVIS, A. (2003): Pre- serving the Tree of Life. Science 300: 1707-1709. May, R. M. (1990): How many species? Philosophical Transactions of the Royal Society of London, Series B 330: 293-304. MUTKE, J. & BARTHLOTT, W. (2005): Patterns of vascular plant diversity at continental to global scales. Biolo- giske Skripter 55: 521-531. MUTKE, J., KIER, G., BRAUN, G., SCHULTZ, C. & BARTH- LOTT, W. (2001): Patterns of African vascular plant di- versity - a GIS based analysis. Systematics and Geog- raphy of Plants 71: 1125-1136. NAGEL, P. 1987. Arealsystemanalyse afrikanischer Fúhlerkáfer (Coleoptera, Carabidae, Paussinae). Ein Beitrag zur Rekonstruktion der Landschaftsgenese. Erdwissenschaftliche Forschung 21: 1-233. Franz Steiner Verlag, Wiesbaden. NOVOTNY, V., BASSET, Y., MILLER, S. E., WEIBLEN, G. D., BREMER, B., CIZEK, L. & DROZD, P. (2002): Low host specificity of herbivorous insects in a tropical forest Nature 416: 841-844. MDEGAARD, F. (2000): How many species of arthropods? Erwin's estimate revised. Biological Journal of the Linnean Society 71: 583-597. J RODRIGUES, A. S. L., ANDELMAN, S. J., BAKARR, M. I., BOITANI, L., BROOKS, T. M., COWLING, R. M., FISH POOL, L. D. C., DA FONSECA, G. A. B., GASTON, K. J. & HOFFMANN, M. (2004): Nature 428: 640-643. SCHATZ, G. E. (2002): Taxonomy and herbaria in service of plant conservation lessons from madagascar's en- demic families. Annals of the Missouri Botanical Gar- den 89: 145-152. WAGNER, Th. (2000a): Diversity and distribution patterns of beetles in different forest types in the Budongo For- est Reserve, Uganda. Biotropica 32: 502-514. WAGNER, Th. (2000b): Revision of afrotropical Monolepta species (Coleoptera, Chrysomelidae, Galerucinae). Part I: Species with red and black coloured elytra, pronotum and head, with description of new species. Entomologische Zeitschrift 110: 226-237. WAGNER, Th. (2001a): Biogeographical and evolutionary aspects of aftrotropical Monolepta Chevrolat, Afro- maculepta Hasenkamp & Wagner and Bonesioides Laboissiere (Coleoptera: Chrysomelidae, Galerucinae). Cimbebasia 17: 237-244. WAGNER, Th. (2001b): Revision of Afrotropical Monolepta Chevrolat, 1837 (Coleoptera: Chrysomeli- dae, Galerucinae). Part II: Species with red elytra, pronotum and elytra, with descriptions of new species. Bonner Zoologische Beitráge 50: 49-65. WAGNER, Th. 2002. Revision of Afrotropical Monolepta species (Coleoptera, Chrysomelidae, Galerucinae). Part III: Species with red elytra and yellow prothorax, including description of new species. Deutsche Ento- mologische Zeitschrift 49: 27-45. WAGNER, Th. 2003a. Present status of a taxonomic revi- sion of afrotropical Monolepta and related groups (Galerucinae). Pp. 133-146 in: FURTH, D. G. (ed.) Special Topics in Leaf Beetle Biology. Proceedings V. International Symposium on the Chrysomelidae, Foz do Iguacu 2000. Pensoft, Sofia. WAGNER, Th. 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Are contents and length of the paper well balanced? Are there any important references to the subject that have been considered? Are all tables and illustrations necessary? Are the tables and figures sufficient? not ] 7 | o ; o . ] y : A EN . . rn ] a. | k y a o a oe i j ‘ A o 7 7 o A A AS 0 1 AS Ñ 1 ae j Ar en) : . = ' e ‘i » : ee Ry Ne 2 N a, 7 . ' 1, ya 4 m ' 7 f j ; y 5 f er o 7 Ñ ral I} 1a ly | Hi i i " \ o " \ f 3 ' ' t al y N o . q A e ' s ny) oe , o 7 , . \ i f . er N . ME 4 y j . . nn O n : n ' 3 o DA y ™ : 2 . “3 A SCHUNKE, Anja €. & HUTTERER, Rainer: 169 The Variance of Variation: Geographic Patterns of Coat Colouration in Aromalurops and Anomalurus (Mammalia, Rodentia, Anomaluridac) SCOBLE, Malcom J.: 18 Issues in Delimiting Genera in Invertebrates: an Example from the Lepidoptera (Macariini: Geometridae: Ennominac) SINCLAIR, Bradley J. & SAIGUSA, Toyohei: 193 Revision of the Trichockinocera dasysentellum Group from East Asia (Diptera: Empididae: Clinocerinac) SONNENBERG, Rainer « BLUM, Thomas: 211 Aphyosemion (Mesoaphyosemion) etsamense (Cyprinodontiformes: Aplocheiloidei: Nothobranchiidae), a New Species from the Monts de Cristal, Northwestern Gabon SPEIDEL, Wolfgang « STUNING, Dieter: 221 Ambia naumanni sp. n., a New Species of Musotiminae from Yunnan (Lepidoptera, Crambidae) SPEIDEL, Wolfgang, BUSCHBAUM, Ulf & MILLER, Michael A.: 227 A New Paracymoriza Species from Lombok (Indonesia) (Lepidoptera, Crambidae) VANE-WRIGHT, Richard I. « BOPPRE, Michael: 235 Adult Morphology and the Higher Classification of Bra Hubner (Lepidoptera: Nymphalidae) WAGNER, Thomas: 255 | Revision of the zincta Species-group of Monolepta Chevrolat, 1837 from Africa, Arabia and the Near East (Coleoptera: Chrysomelidae, Galerucinae) Kürer, Wolfgang, WAGNER, Thomas & BARTHLOTT, Wilhelm: 283 | Diversity Patterns of Plants and Phytophagous Beetles in Sub-Saharan Africa Photo by M. Schmitt, A. Hofmann & O, Niehuis ITHSONIAN | Im 088 AT SCHMITT, Michael: 1 Clas M. Naumann (26.06.1939 — 15.02.2004) — in memoriam AsPOCK, Horst & ASPOCK, Ulrike: 13 Neuropterologische Beitrage in den Werken von Wilhelm Friedrich VON GLEICHEN, genannt Russwurm (1717-1783) BISCHOFF, Inge, ECKELT, Esther & KUHLMANN, Michael: al On the Biology of the Ivy-Bee Colletes hederae Schmidt € Westrich, 1993 (Hymenoptera, Apidae) GASSMANN, Dirk: am The Phylogeny of Southeast Asian and Indo-Pacific Calicnemiinae (Odonata, Platycnemididae) HOFMANN, Axel: 81 Neue Zygaena-Taxa aus Südosteuropa, Vorder- und Zentralasien (Lepidoptera: Zygaenidae) Huper, Bernhard A., PEREZ G., Abel & BAPTISTA, Renner L. C.: 99 Leptopholcus (Araneae: Pholcidae) in Continental America: Rare Relicts in Low Precipitation Areas HUNDSDÖRFER, Anna K.: 109 First Record of Hy/es dahl (Geyer, 1827) (Lepidoptera: Sphingidae) from the African Mainland LOURENCO, Wilson R.: 111 A New Genus and Species of Scorpion from Afghanistan (Scorpiones, Buthidae) MEY, Wolfram & SPEIDEL, Wolfgang 115 Two New Species of Eoophyla Swinhoe, 1900 from Continental South East Asia (Lepidoptera: Crambidae, Acentropinae) MIsoF, Bernhard, KLUTSCH, Cornelya E C., NIEHUIS, Oliver & PATT, Alexandra: 121 Of Phenotypes and Genotypes: Two Sides of one Coin in Taxonomy? RÖSLER, Herbert, ZIEGLER, Thomas, Vu, Ngoc Thanh, HERRMANN, Hans-Werner & BÖHME, Wolfgang: 135 A New Lizard of the Genus Gekko Laurenti, 1768 (Squamata: Sauria: Gekkonidae) from the Phong Nha — Ke Bang National Park, Quang Binh Province, Vietnam SAUER, Klaus Peter & KULLMANN, Harald: 149 Analyse der biologisch-ókologischen Ursachen der Evolution der gastrcneuralen Metazoa — Testen einer phylogenetischen Hypothese SCHINTLMEISTER, Alexander: 165 Peridea clasnaumanni spec. nov. (Lepidoptera: Notodontidae) aus China (Continued on inside back cover) museum : KOENIG Herausgegeben vom Zoologischen Bonner zoologische Beiträge / Forschungsmuseum Alexander Koenig, Bonn Band DO Heft 3/4 2005 (2004) +f Leibniz Gemeinschaft Bonner zoologische Beitrage werden publiziert 1m Eigenverlag Zoologisches Forschungsmuseum Alexander Koenig, Adenauerallee 160, D-53113 Bonn (Germany) Die Zeitschrift erscheint mit vier Heften 1m Jahr, zum Preis von 11,50 € je Heft bzw. 46,- € je Band incl. Versand. 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University, WTAMU Box 60808, Canyon, Texas 79016, USA, E-mail: dsıssom@wtamu.edu/ Dr. Miguel VENCES, Technische Universitat, Zoologisches Institut, Abteilung Evolutionsbiologie, Spielmannstr. 8, D-38106 Braunschweig, Germany, Tel. +49 531 391 3237, Fax +49 531 391 8198, E-mail: m.vences(@tu-braunschweig.de PD Dr. Heike WÄGELE, Rheinische Friedrich-Wilhelms- Universität, Institut für Evolutionsbiologie und Okologie, D-53121 Bonn, Tel.: +49 228-73 5159, Fax: +49 234-322 4114, E-mail: hwaegele(@evolution.uni-bonn. de Dr. Erich WEBER, Eberhard-Karls-Universitat, Zoologische Schausammlung, Sigwartstr. 3, D-72076 Tubingen, Germany, E-mail: erich. weber(@uni-tuebingen. de Bonner zoologische Beitráge Band 53 (2004) | Heft 3/4 | Seiten 293-295 Bonn, Dec. 2005 Do Endings of Adjective Flectible Species Names Affect Stability? A Final Note on the Gender of Podarcis Wagler, 1830 / (Reptilia, Lacertidae) Wolfgang BOHME & Jórn KOHLER Zoologisches Forschungsmuseum Alexander Koenig, Bonn, Germany Abstract. We here reply to the arguments put forward by ARNOLD (2000) and LANZA & BOSCHERINI (2000) concerning the gender of the genus Podarcis. ARNOLD’s argument that a change of endings of adjective flectible species names threatens stability is rejected by clearly following the articles of the current edition of the Code (ICZN 1999). We finally conclude that the gender of Podarcis must be masculine for the following reasons: (1) WAGLER (1830) did not deter- mine the gender by combination with any species name nor by statement, (2) the name Podarcis is of common variable gender, (3) the Principle of First Reviser does not apply to determination of gender of names, (4) the Code clearly states that a name of common variable gender is to be treated as masculine. Key words. Grammatical gender, nomenclature, stability of names. In a previous note, BOHME (1997) had argued that the lacertid lizard genus Podarcis was not of feminine (as generally used before) but of masculine gender. This view provoked strong criticism (MAYER 1998; ARNOLD 2000; LANZA & BOSCHERINI 2000). Whereas BOHME (1998) replied already to the critics of MAYER (1998), we feel it also necessary to answer on the views put forward by ARNOLD (2000) and by LANZA & BOSCHE- RINI (2000). BOHME’s (1997) main argument for the “masculiniza- tion’ of Podarcis was not the Homerian use of the Greek adjective podarkes attributed to the ancient hero Achilleus (Ayi//evg); this circumstance was only a first hint to the problem, which lead to question the feminine gender of this lacertid genus name. The point was rather the assumption that FITZINGER (1843) was the first to fix a type species for Podarcis Wagler, 1830, viz. Seps mu- ralis Laurenti, 1768. Furthermore, although being aware that muralis is again an ambiguous adjective as to its gender (masculine or feminine), BÖHME (1997) re- garded the genus name Seps in LAURENTI's (1768) use as masculine because this author himself used it clearly in this way by describing for instance Seps argus and Seps ruber (= Lacerta agilis Linnaeus, 1758) in the same work. LAURENTI's (1768) practice is apparent in other 18th century names, e.g. Seps stellatus Schrank, 1798 (likewise a synonym of L. agilis). Some 19th cen- tury names, however, implied Seps also to be a feminine noun, combining it with species names such as Seps vit- tata Leuckart, 1828 or Seps quadrilineata Metaxa, 1833, but this does not affect the original combination and therefore masculine determination of the genus’ gender by LAURENTI (1768). From these grounds, we think that the viewpoint to regard Seps Laurenti as masculine is still justified. In contrast, BOHME’s (1997) assumption that FITZINGER (1843) was the first to have ‘masculinized’ Podarcis Wagler by fixing a (masculine) type species, cannot withstand the arguments put forward by ARNOLD (2000) who justifiably states that Podarcis was earlier treated as feminine by BONAPARTE (1836), since this author used the combinations Podarcis taurica, P. oxycephala and P. muralis sicula and conformed this usage subse- quently (BONAPARTE 1839). However, the discussion about the first revising author according to article 24.2. (ICZN 1999) does not apply to the determination of gender of names, as already stated correctly by other authors (KWET 2001, CARAMASCHI 2004). The Code does not allow the adscription of the gender by any subsequent author and all the sources for gender identification must be found in the original pub- lication. Therefore, the arguments of BOHME (1997) and ARNOLD (2000) concerning first reviser’s action are both not applicable to clarify the grammatical gender of Podarcis. LANZA & BOSCHERINI's (2000) arguments for a femini- ne gender, however, are very weak. They do restrict the problem to the philological “side of the medal”. They regard BOHME’s (1997) view an “unverifiable hypothe- sis”, but assume instead that “almost surely the etymol- ogy of Podarcis is not from the Greek adjective “po- darkes’ but from the Latin proper name Podarce (....), of course of Greek derivation, meaning ‘Swiftfoot’ and mentioned by Homer as one Harpy....”. This assumption that WAGLER's (1830) name Podarcis is not at all de- 294 rived from an adjective, but from a “Latin proper name” (why should Homer use Latin names in his “Iliad’”?) is likewise an unverifiable hypothesis and cannot be taken as “evidence” for a proposed continuing use of Podarcis as feminine. We therefore agree with ARNOLD (2000) in that Podar- cis derived from the Greek (subsequently Latinized) ad- jective “podarcis” with originally undetermined gender. We disagree, however, with ARNOLD's (2000) further reasoning that a change of a genus' gender and a subse- quent adaptation of the endings of species names (as re- quired by Article 34.2. of the Code) would “contravene the spirit of the International Code of Zoological No- menclature (International Trust of Zoological Nomen- clature, 1999) which promotes the stability of names, for example in the Preamble (p. 2) and in Articles 23.2 and 81)”. Our point is here, that a change of the ending of an adjective flectible species name does not touch the problem of stability and universality at all. A changed ending of a given name is not a new or a different, but absolutely still the same — and therefore stable — name. We think that the change of an ending of a specific name as an adaptation to the gender of a changed, dif- ferent genus should be regarded as a normal process, because the change of generic names is due to zoologi- cal arguments and thus has nothing to do with nomen- clatural instability. A good and rather recent herpeto- logical example is that of the spiny-tailed agamas, Uromastyx, a genus which was treated as masculine for decades until LANZA (1983) justifiably claimed that there is hardly a more feminine word to be found than the Greek word “mastix”, wherefore Uromastyx 1s clearly and in a philologically correct way to be consid- ered as feminine. No problems arose since then to change the naming of Uromastyx acanthinurus, U. ae- gyptius, U. ocellatus, U. ornatus, etc. into U. acanthin- ura, U. aegyptia, U. ornata and so on, just because the names have not at all been changed, and the “virtues of stability” (ARNOLD 2000) have been respected. ARNOLD's (2000) further argument that the change of a species name's ending could cause confusion in non- specialist users 1s also not convincing to us. To stay with the example of Uromastyx, even non-taxonomist working in applied disciplines such as ecology, etc. could not really suspect U. loricata to be a different tax- on from U. loricatus. A similar case (out of hundreds more) is the Madagascan geckonid genus Paroedura, which had to be split off from the paraphyletic Phyllo- dactylus (that means for zoological and not for nomen- clatural reasons) (DIXON & KROLL 1974). But unfortu- nately, specific names such as Phyllodactylus pictus remained unchanged as Paroedura pictus for some time, certainly not because of an intended “virtue of stability” of the respective authors, but much more likely because of a lack of Greek and Latin knowledge which lead to Bonner zoologische Beitráge 53 (2004) the ignorance of Art. 34.2. of the Code; or the amend- ment of endings has simply been forgotten in this and other cases. Fortunately, NUSSBAUM & RAXWORTHY (2000) provided the correct spellings in their recent re- vision of the genus. If a changing ending of a name is thought to threaten stability and universality, the Code (ICZN 1999) would have to be altered in that the endings of adjective, flect- ible specific names would not have to follow the gender of the genera any more, 1.e. Article 31.2. would have to be deleted. But this would at the same time destroy the philological basis on which the whole system of bio- logical nomenclature is built, viz. the Greek and Latin languages. Our view is furthermore supported by Article 30.1.4.2. of the current edition of the Code (ICZN 1999), which clearly states that “a genus-group name that is or ends in a word of common variable gender (masculine or feminine) is to be treated as masculine unless its author, when establishing the name, stated that it is feminine or treated it as feminine in combination with an adjectival species-group name”. This article probably applies to several other cases where the genus’ gender was not determined in the original description and therefore more changes of adjectival species-group names from feminine to masculine are to be expected. If such actions threaten stability, the Code would be self- contradictory — we think it is not. A last point which we should like to address here is ARNOLD's (2000) final recommendation: “However, .... changes, for instance when a group within a known clade is separated as a new genus, can be avoided by using subgenera”. This proposal considers genera simply as operational units and is contra productive to modern ef- forts for a new, less arbitrary generic concept (see e.g. DUBOIS 1988), which includes biological reality and takes into account that the genus, in spite of all subsequent higher categories, is the classificatory level above the species where reticulate evolution just begins to stop. In conclusion, by considering: (1) that WAGLER (1830) did not combine the generic name Podarcis with any species name and therefore did not demonstrate the in- tended gender of the genus; (2) that the name Podarcis is most probably derived from the Greek adjective ‘po- darkes’ (Latinized: ‘podarcis’) and hence is of variable gender (masculine or feminine); (3) that the Principle of First Reviser does not apply to determination of gender of names; and (4) that the Code (ICZN 1999) clearly states that a name of common variable gender is to be treated as masculine, we hereby reassert that the gender of Podarcis must be considered masculine. Acknowledgements. We are indebted to two anonymous reviewers for useful comments and to Anthea Gentry (ICZN) for further discussion. Wolfgang BOHME & Jórn KOHLER: Gender of Podarcis — Stability of Names 295 REFERENCES ARNOLD, E. N. 2000. The gender of Podarcis and the vir- tues of stability, a reply to W. Böhme. Bonner zoolo- gische Beitráge 49( 1-4): 71-74. BOHME, W. 1997. A note on the gender of Podarcis (Sauria: Lacertidae). Bonner zoologische Beiträge 47(1-2): 187-188. BOHME, W. 1998. Podarcis siculus, -a, -um? — Entgegnung auf eine Entgegnung. Die Eidechse 8: 101-102. BONAPARTE, C. L. J. L. 1836. Iconographia della Fauna Ita- lica per le quattro classi degli Animali Vertebrati. Ro- ma: Salviucci. BONAPARTE, C. L. J. L. 1839. Amphibia Europaea ad systena nostrum vertebratorum ordinata. Memorie del- la Accademia della Scienze di Torino (Series 2) 2: 385-415. CARAMASCHI, U. 2004. The gender of the genus Scinax Wagler, 1830 (Anura, Hylidae). Herpetological Re- view 35(1): 27-31. Dixon, J. R. & KROLL, J. C. 1974. Resurrection of the ge- neric name name Paroedura for the phyllodactyline geckos of Madagascar, and description of a new spe- cies. Copeia 1974(1): 24-30. Dusots, A. 1988. The genus in zoology: a contribution to the theory of evolutionary systematics. Mémoires du Museum national de l'Histoire naturelle Paris (Zool.) 140: 1-122. FITZINGER, L. 1843. Systema reptilium. Vindobonae: Brau- miiller et Seidel, 106 + 1x pp. ICZN — INTERNATIONAL COMMISSION ON ZOOLOGICAL NOMENCLATURE. 1999. International Code of Zoo- logical Nomenclature adopted by the International Un- ion of Biological Sciences. Fourth Edition. Interna- tional Trust for Zoological Nomenclature, London, 306 + xxix pp. KwET, A. 2001. Siidbrasilianische Laubfrösche der Gat- tung Scinax mit Bemerkungen zum Geschlecht des Gattungsnamens und zum taxonomischen Status von Hyla granulata Peters, 1871. Salamandra 37(4): 211- 238. LANZA, B. 1983. A list of the Somali amphibians and rep- tiles. Monitore zoologico Italiano 1983: 193-247. LANZA, B. & BOSCHERINI, S. 2000. The gender of the gen- era Podarcis Wagler 1830 (Lacertidae), Pelamis Daudin 1803 (Hydrophiidae) and Uropeltis Cuvier 1829 (Uropeltidae). Tropical Zoology 13: 327-329. LAURENTI, J. N. 1768. Synopsin reptilium. Viennae, Joan. Thom. Nob. de Trattnern, 214 pp. MAYER, W. 1998. Das Geschlecht der Gattung Podarcis: Eine Entgegnung. Die Eidechse 8: 99-100. NUSSBAUM, R. A. & RAXWORTHY, C. J. 2000. Systematic revision of the genus Paroedura Günther (Reptilia: Squamata: Gekkonidae), with the description of five new species. Miscellaneous Publications Museum of Zoology University of Michigan 189: 1-26. WAGLER, J. G. 1830. Natiirliches System der Amphibien, mit vorangehender Classification der Sáugethiere und Vögel. München, Stuttgart, Tübingen, Cotta'sche Buchhandlung, 354 + vi pp. Authors’ address: Prof. Dr. Wolfgang BOHME (cor- responding author), Dr. Jórn KOHLER, Zoologisches For- schungsmuseum Alexander Koenig, Adenauerallee 160, 53113 Bonn, Germany. E-mail: w.boehme.zfmk@uni- bonn.de, j.koehler.zfmk@hlmd.de Received: 13.06.2002 Accepted: 09.01.2003 Revised: 21.06.2004 Corresponding editor: M. Schmitt Bonner zoologische Beitrage | Band 53 (2004) Heft 3/4 Seiten 297-301 Bonn, Dec. 2005 Morphology, Genitalia, and Natural History Notes. on the Enigmatic Tiger Beetle, Mantica horni ; Kolbe, 1896 (Coleoptera, Cicindelidae) Michael FRANZEN” & Walter HEINZ”? 7 Oberneuching, Germany Schwanfeld, Germany Abstract. The morphology of the extremely rarely collected tiger beetle, Mantica horni, is described on the basis of a large series from a new locality near Maltahóhe, southern Namibia. Male genitalia and female external genitalia are de- scribed for the first time and compared with those of the presumed sister genus, Manticora. Additionally, we firstly re- port on habitat and behaviour of the species. Key words. Manticora, habitat, Namibia. 1. INTRODUCTION The cicindelid tribe Manticorini consists of two genera, Mantica Kolbe, 1896 and Manticora Fabricius, 1792 and contains very large, crespuscular to nocturnal, ex- clusively southern African species. In contrast to the comparably well known genus Manticora (e.g., KLUG 1849; ROER 1984; MARES 1976, 1995, 1997, 2002; LEFFLER 1980; WERNER & WIESNER 1994, 1995; WERNER 2000), the only species of the genus Mantica, the southern Namibian M. horni Kolbe, 1896, is re- garded as extremely rare and its biology is virtually un- known. Since the original description, which is based on two males (KOLBE 1896), only a very few additional in- dividuals have been collected, most of them presumably incidentally (MARES 1976, 2002; WERNER & WIESNER 1994, 1995; WERNER 2000). Comprehensive informa- tion on the morphology of the species is only given in the original description, where KOLBE (1896) presents a fairly good description of the two male syntypes (for an English translation of the original description see PERINGUEY 1896). Subsequently, there were almost no data added, with the exception of a few comparative features mentioned by HORN (1910) and some basic habitat data (“Sand plateau on top of the Fish River Canyon without almost no vegetation”, “long grass and other scattered plants near Karasburg”) by MARES (2002). Most recently, good photographs of almost all collected specimens including one of the syntypes (not “holotype” as mentioned by MARES 2002) were pub- lished by WERNER & WIESNER (1994), WERNER (2000), and especially MARES (2002). However, until today al- most any detailed information on habitat, biology and even female morphology are lacking. In February 2003, O. and W. Heinz discovered a mass aggregation of Mantica horni. This gave us the oppor- tunity to report firstly on certain aspects of the mor- phology, genitalia, habitat, behaviour, and prey of this enigmatic species. 2. MATERIAL AND METHODS In total 45 specimens (24 males, 21 females) were ex- amined. The series was collected on 5 February 2003, approximately 3-5 km W of Maltahóhe (Namibia, Har- dap district) at the upper course of the Hudup River (a tributary of the Fish River), 24°50'S, 16°54'E, 1300 m a.s.l. All specimens are currently in the authors' collec- tions. MARES (1995) found no distinguishing characters among male genitalia of Manticora species. Consequently, we compared Mantica genitalia only with one male of Man- ticora livingstoni from 30 km E of Rundu, Namibia, and one female of Manticora cf. latipennis from an un- known locality in the Republic of South Africa (both in the collection of M. Franzen). Measurements taken are: total length (without labrum), body width (across the widest points of the elytrae), and elytral length. The internal sacs of two male aedeagi were gradually blown out with water. Toothpaste was than injected into the fully blown out sacs to keep them in form; finally they were fixed and stored in 5% for- maldehyde solution. Female genitalia were removed from two individuals, which had their ovipositors everted. The nomenclature of the male genitalia follows ISHIKAWA (1978) and MATALIN (1998), and those of the female genitalia FREITAG (1972). 298 Bonner zoologische Beitráge 53 (2004) 3. RESULTS 3.1. Description 3.1.1. Morphology. Coloration entirely shiny black. Head. Antennal scape and antennal segments 2 to 4 with scattered semierect setae additionally to sensories (Fig. la). Left mandible (Fig. 1b, c) slightly longer than right in males; in a very few individuals mandible lengths appear almost equal; mandible lengths equal in all females; outer basal half of mandibles with long e- rect setae. Labrum (Fig. 1d) much wider than long, with 6-10 (mostly 8) submarginal setae; indistinctly quadri- dentate, “teeth” appear as rounded bulges. Clypeus with some 12-15 long erect setae. Frons and vertex smooth (except for setigerous punctures), shiny; frons with me- dian field of long erect setae, extending anteriorly to ey- es; a conspicuous ridge present from above insertation of antennae to middle of eye; row of four supraorbital setae extending posteriorly to irregular row of 10-12 se- tae submarginally across head. Eyes small. Genae with scattered, long, erect, thin setae. Prothorax. Pronotal shape globose, with very deeply impressed anterior and posterior transverse sulci, latter forming two somewhat rounded-triangular, posteriorly reaching lobes; a minute bulge at posterior angle; median longitudinal sulcus in- \ distinct if visible; a short lateral bead extends from shortly after anterior transverse sulcus to posterior third of pronotum; pronotal surface shiny, with scattered long, erect setae along anterior, posterior, and lateral margins; anterior setose margin reaching to anterior transverse sulcus, lateral margin reaching onto disc; centre of disc almost glabrous except some single short setae. Proepisterna shiny with smooth surface. Proster- num with transverse ridge bearing row of short erect se- tae. Pterothorax. Mesepisterna glabrous, females lack coupling sulcus. Elytra. Shape of elytrae ovoid, apically pointed, shoulders rounded; surface of elytrae and epipleura of elytrae with numerose large carinate tuber- cles (largest ending in a cylindrical cone), each one bea- ring a fine, long seta at its posterior base; tubercles have their highest point posteriorly; tubercles very small and almost lacking on anterior elytrae along suture and densest and highest within apical third of elytrae; api- cally largest tubercles form two oblique, serrated ridges, starting at posterior central third and extending to apical tip of suture. Abdomen. Sterna with transverse rows of fine, long semierect setae. Legs. Coxae with fine long setae, densest posteriorly; trochanters adpressed setose; femora, tibiae, and tarsomeres with dense cover of thick, black, erect setae; tarsi not dilated in males. Fig. 1: Mantica horni. a) first five antennal segments, male; b) left mandible, female; e) left mandible, male; d) labrum, male. Scale bar: 2 mm. Michael FRANZEN & Walter HEINZ: Morphology and Natural History of Mantica horni 3.1.2. Male genitalia. Shape of aedeagus moderately elongate and evenly curved, with considerably thick- ened basal part and very tiny apical knob (Fig. 2c); it lacks any structures like concavities, lateral flanges or lobes; dorsal ostium (apical orifice) very long, forming about 40% of aedeagal length, and basally divided by narrow, sclerotized thorn (Fig. 2c). Internal sac (Fig. 2a, b) strongly elongate, with large and protuding, thumb- like basal bladder; dorsally a long sclerite which is pro- bably identical with ligula sensu ISHIKAWA (1978). Truncal tube extremely elongate, formed by basi-lateral right bladder. Sclerites and bladders on apical portions only indistinctly discernible in our specimen. The iden- tifiable stuctures include a prominent praeputial pad, ve- ry long flagellum, relatively small medial tooth, upper limitator, and shield (Fig. 2a, b). Fig. 2: Male genitalia of Mantica horni. a) internal sac, right lateral view; b) apex of internal sac, left lateral view; c) adeagus. Sclerites: fl = flagellum, li = ligula, mt = median tooth, sh = shield, ul = upper limitator; bladders: B = basal bladder, BLR = basi-lateral right bladder, PP = praeputial pad. Scale bars: 3 mm. The aedeagus and the general internal sac proportions and structures resemble those of Manticora livingstoni almost completely. The only exception is the lack of a 299 praeputial pad in Manticora. However, it may be that this structure has been incompletely blown in the latter. 3.1.3. Female external genitalia. In two females ovi- positors everted with a length of 32-34 mm (oviduct + ovipositor). Sternum eight (Fig. 3a, b) apically broadly rounded, with narrow, triangular medial emargination: apex with numerous short, fine setae. Second gonocoxa with no ventral groove or notch, inner apical sides with some scattered very fine short setae (Fig. 3a). Second gonapophyses short and broad; lateral portions some- what triangular and distinctly shorter than medial, both portions curved dorso-medially (Fig. 3a, b). Syntergum 9&10 appearing somewhat ovoid with lateral portions slightly pointed at apex (Fig. 3c); apical two-thirds of lateral portions with numerous setae. Female external genitalia of Manticora resemble those of Mantica horni almost completely, with the exception of less broadened medial and lateral portions of second gonapophyses, less pointed lateral portions of synter- gum 9&10, and the overall larger size in Manticora. 3.1.4. Sexual dimorphism. Besides different mandible lengths and a tendency to have the left mandible longer than the right in males (see above), there are several morphometric characters which separate males from females: males have slightly shorter total lengths than females and possess shorter and narrower elytra (see Table 1.2 EL/TL,.BW/TL, BW/EL): Table 1: Morphometric values of Mantica horni. TL = total length; EL = elytral length; BW = body width. TL EL/TL BW/TL BW/EL Males ( n= 24) mean 28.0 0.60 0.39 0.66 standard deviation 1.08 0.01 0.01 0.03 range 26.0-30.0 0.58-0.62 0.37-0.41 0.61-0.71 Females (n = 21) mean 30.0 0.62 0.42 0.69 standard deviation 1:11 0.01 0.01 0.02 range 27.5-32.0 0.59-0.65 0.40-0.44 0.66-0.73 3.2. Habitat and natural history observations All specimens were collected from a gravel road leading between the Hudup River and the south-facing slopes of the directly adjoining hilly terrain. The steep slopes (in- clination about 30°) were rocky and had about 40% rocks, stones and gravel on the surface, with inter- spersed fine loamy substrate. However, loamy areas dominated near to the river. The vegetation consisted of grasses and shrubs, with interspersed thorn bushes. Vegetation cover was approximately 50%, with bushes making about 15%. 300 On February 5, the locality was visited two times. Dur- ing the first visit, about two hours before dusk, no bee- tles were observed. Returning one hour later, the mass aggregation was observed. From that time activity con- tinued until dusk and then markedly decreased. During the observations the weather was clear and air tempera- ture was around 30°C. On the previous day, a heavy thunderstorm brought the first heavy rains after a long dry season and there was much humidity around with puddles and small rivulets along the road. The locality was revisited in early morning of the next day, but without finding any more Mantica. Fig. 3: Female external genitalia of Mantica horni. a) apex of sternum, ventral view; b) apex of sternum, lateral view; ec) syntergum 9&10, dorsal view. s8 = sternum eight, sgp = second gonapophysis, sgx = second gonocoxa, t9&10 = syntergum 9 & 10. Scale bar: 2 mm. It appeared that during the beginning of the observations beetles' movements were directed downhill towards the road/river. Later, individuals stayed on the road. Cou- Bonner zoologische Beiträge 53 (2004) pling pairs comprised about 30% of the beetles ob- served. In addition, two females were presumably taken during oviposition because they had their ovipositors everted. Most individuals (including mating couples) were found actively foraging on swarming termites. Movements were fast and if disturbed, individuals tried to escape by running even more quickly into dark, shaded areas (e.g., under the car). Single individuals or couples were observed every 10 to 100 m along the road. Although no individuals of the series were teneral, three males possessed comparably soft aedeagi and it appears that hardening after emergence may not have been fully completed in these individuals. Thus, at least a part of the series may have been taken only several days after emergence (see WILLIS 1967: 189-190 for a timetable of hardening after emergence in some North American Cicindela). 4. DISCUSSION The new locality extends the known range of the species some 190 km to the north (measured from the formerly known northernmost precise locality (“Bethanien [Ha- nam Plateau]”: WERNER & WIESNER 1994). Despite this considerable range extension, we could not find any ob- vious morphological differences among our new mate- rial and the previously collected specimens (compared with the photographs in MARES 2002; WERNER 2000; WERNER & WIESNER 1994). We found only one exter- nal character mentioned by KOLBE (1896), which is dif- fering in our new material: KOLBE (1896) states that right and left mandible lengths are equal in males. Al- though indistinctly visible in some individuals, most of our males possess a slightly longer left mandible. The close relationship of Mantica and Manticora is stressed by several unique characters (see also MARES 2002): both genera possess unequal mandible lengths in males (but character heterogenous in Mantica), lack di- lated tarsi in males, possess ciliated apical palpal seg- ments and long setae on the outer mandible sides, have similar organized dorsal elytral tubercles which form two oblique ridges apically, and have rather similar male and female genitalia. By contrast, both genera are well separated by the longer, asymmetrically formed mandibles of Manticora, more labral teeth in Manticora (6 vs. 4), the generally much more “robust” habitus of all Manticora species, which is basically expressed by broader heads and elytrae, and pronounced bulged ely- tral shoulders. The observations at Maltahóhe indicate that short sur- face activity of Mantica is linked with rare and strongly localized rains in its arid habitats. WERNER (2000) men- tioned several observations that Manticora species were Michael FRANZEN & Walter HEINZ: Morphology and Natural History of Mantica horni 301 found abundantly under the same conditions, 1.e. hunt- ing for myriapods and tenebrionids after rainfall. Prey items have never been reported for Mantica, but we pre- sume that the species is an opportunistic feeder, pre- dominantly preying on such invertebrates, which share their seasonal surface activity with the tiger beetle. Acknowledgements. Many thanks to Jürgen Wiesner (Wolfsburg), who provided — once again — important litera- ture and commented a previous draft of the manuscript. We are also grateful to two anonymous reviewers for their helpful comments. REFERENCES FREITAG, R. 1972. Female genitalia of the North American species of the Cicindela maritima group (Coleoptera, Cicindelidae). The Canadian Entomologist 104: 1277- 1306. Horn, W. 1910. Coleoptera Adephaga, Fam. Carabidae, Subfam. Cicindelinae. Pp. 105-208 in: WYTSMAN, P. (ed.), Genera Insectorum. Pars 82B. ISHIKAWA, R. 1978. A revision of the higher taxa of the subtribe Carabina (Coleoptera, Carabidae). Bulletin of the National Science Museum, Tokyo, Ser. A (Zool- ogy) 4(1): 45-68. KLUG, F. 1849. Die Arten der Gattung Manticora F. Lin- naea Entomologica 4: 417-424, pls. 1-2. KOLBE, H. J. 1896. Afrikanische Coleoptera des Kónig- lichen Museums für Naturkunde zu Berlin. II. Theil. Entomologische Nachrichten 22(1): 5-7. LEFFLER, S. R. 1980. The larva of Mantichora Fabricius. Cicindela 12: 1-12. MARES, J. 1976. New species of Mantichora Fabr. (Col- eoptera, Cicindelidae, Mantichorini). Journal of Ento- mological Society of Iran 3(1/2): 137-141. MARES, J. 1995. Determination key to the males the genus Manticora Fabricius 1792 (Coleoptera, Cicindelidae). Acta Coleopterologica 11(2): 47-54. MARES, J. 1997. Nové poznatky ze Zivota mantikor. ZIVA, Casopis pro biologickou praci, 1979/2: 80-82. MARES, J. 2002. Manticora. A monograph of the genus. 205 pp., Taita Publishers, Hradec Kralove. MATALIN, A. V. 1998. The tiger beetles of “hybrida”- species group (Coleoptera, Carabidae, Cicindelinae). Il. A taxonomic review of the Iberian Cicindela, la- gunensis Gautier, 1872 complex. Grasiella 54: 75-96. PERINGUEY, L. 1896. Descriptive catalogue of the Coleop- tera of South Africa. Family Cicindelidae. Supple- ment. Transactions of the South African Philosophical Society 1896: 99-121. ROER, H. 1984. Zum Vorkommen und Beutefangverhalten des Sandlaufkáfers Mantichora latipennis Waterh. (Col.: Cicindelidae) in Súdwestafrika/Namibia. Jour- nal Stidwestafrikanische Wissenschaftliche Gesell- schaft 38: 87-93. WERNER, K. 2000. The tiger beetles of Africa (Coleoptera: Cicindelidae). Volume 1. 191 pp., Taita Publishers, Hradec Králové. WERNER, K. & WIESNER, J. 1994. Die Cicindelidae von Namibia (Coleoptera). Lambillionea XCIV(1): 49-80. WERNER, K. 8 WIESNER, J. 1995. Nachtrag zu ,,Die Cicin- delidae von Namibia” (Coleoptera). Lambillionea XCV(4): 603-610. WILLIS, H. L. 1967. Bionomics and zoogeography of tiger beetles of saline habitats in the Central United States (Coleoptera, Cicindelidae). University of Kansas Sci- ence Bulletin 47(5): 145-313. Authors’s addresses: Michael FRANZEN (correspond- ing author): Hauptstrasse la, 85467, Oberneuching, Germany; Walter HEINZ: Kembachstrasse 46, 97523 Schwanfeld, Germany. Received: 09.12.2003 Revised: 09.06.2005 Accepted: 10.06.2005 Corresponding editor: M. Schmitt Bonner zoologische Beitráge Band 53 (2004) | Heft 3/4 | Seiten 303-310 Bonn, Dec. 2005 On the Taxonomy and Biogeography of Stenus (s. str.) erythrocnemus Eppelsheim and Related Species / (Insecta: Coleoptera: Staphylinidae) Volker ASSING, Hannover, Germany Abstract. Material previously identified as Stenus erythrocnemus Eppelsheim, S. anatolicus Puthz, and S. maculiger Weise is revised with special reference to the internal structures of the aedeagus. Three new species of Stenus s. str. are described, illustrated, and distinguished from similar congeners: S. cypriacus sp. n. (Cyprus), S. distortus sp. n. (Anatolia), and $. prominens sp. n. (Lebanon, Israel). The distributions of $. cypriacus, S. distortus, S. erythrocnemus, S. anato- licus, and S. maculiger are mapped. The morphological variation of the male sexual characters of 5. ervthrocnemus and S. maculiger is illustrated and discussed. Key words. Palaearctic — Europe — Turkey — Cyprus 1. INTRODUCTION Among the Palaearctic species of the subgenus Stenus Latreille with spotted elytra, a species group can be identified which is characterized by a conspicuous — evidently synapomorphic — sclerotized basal structure in the internal sac of the aedeagus. In the Western Palae- arctic, this group comprises Stenus erythrocnemus Ep- pelsheim, S. maculiger Weise, S. anatolicus Puthz, and S. sacrimontis Puthz. The sclerotized internal structures of S. anatolicus, S. erythrocnemus, and S. sacrimontis were figured by PUTHZ (1970a, b), but their intra- and interspecific variation has never been studied. When comparing recently collected material of S. ana- tolicus from southern Anatolia with males previously identified as S. anatolicus and recorded as such from Cyprus (ASSING & WUNDERLE 2001), I became aware of some evident differences, not only in the size and shape of the median lobe, but especially in the morphol- ogy of the basal internal structure of the aedeagus. A subsequent more systematic study of the internal struc- tures of males previously identified as S. erythrocnemus and S. anatolicus from several public and private collec- tions then revealed the existence of three undescribed species, one from Cyprus, one from Anatolia, and one from Lebanon and Israel. 2. MATERIAL AND MEASUREMENTS The material treated in this study is deposited in the fol- lowing public and private collections: FMNH Field Museum of Natural History, Chicago (A. Newton) Lebanon — new species — distribution MHNG Muséum d'Histoire Naturelle, Geneve (G. Cuccodoro) NHMW Naturhistorisches Museum Wien (H. Schill- hammer) UH University of Haifa cAss author's private collection cFel private collection B. Feldmann, Miinster cPut private collection V. Puthz, Schlitz cRos private collection A. Rose, Oldenburg cSch private collection M. Schülke, Berlin cWun private collection P. Wunderle, Mönchenglad- bach The measurements are given in mm and abbreviated as follows: HW: head width across (and including) eyes; PW: maximal width of pronotum; PL: length of prono- tum along median line; EL: length of elytra from apex of scutellum to elytral hind margin; EW: width of elytra; ML: length of median lobe of aedeagus. 3. STENUS (S. STR.) ERYTHROCNEMUS EPPELSHEIM AND RELATED SPECIES 3.1. Stenus anatolicus Puthz (Fig. 1, Map 1) Types examined: Paratype Í: Anatolia mer., Korge & Heinz leg. / Pisid. Taurus, Bozburun, 1600-2000m, 20.VII.65 / Ö Paratype / Stenus anatolicus n. sp. det. V. Puthz 1968 (cPut). Two examined paratypes from Maras are not conspecific with the paratype from Bozburun, which was collected together with the holotype, and are now listed as paratypes of S. distortus sp. n. (see below). 304 Bonner zoologische Beitráge 53 (2004) Additional material examined: Turkey: Mugla: 14, Dalaman, 27.V.1991, leg. Schónmann & Schillhammer (cPut). Antalya: 14, Fa- kircal1, 22 km NE Demirtas, 600-700m, 4.V.1997, leg. Schulz, Vock & Sanetra (cAss); 13, 12, E Kumluca, 36°21°50N, 30°22'27E, 385m, stream bank, 3.1V.2002, leg. Assing & Wunderle (cAss); 12, SSW Antalya, Ciralı, 36°25N, 30°28E, 20-50m, stream bank, 4.1V.2002, leg. Assing & Wunderle (cAss); 14, Kemer, 3.-15.V11.1989, leg. Schmid (cPut); 18, 19, Bey Dagları, V.1968, leg. Fagel (cPut); 12, 3 km W Ciralı, 36°25 27N, 30°26 41E, 71m, stream bank, 28.111.2001, leg. Rose (cRos). Isparta: 14, S Egridir, Dedegöl Dag, N-slope, 1300- 1800m, 28.-29.VII.1971, leg. Heinz (cPut). Mersin: 734, 27%, Hacıiskakli, 27.1V.1978, leg. Be- suchet & Löbl (MHNG, cPut, cAss). Mersin or Karaman: 14, S Karaman, Sertavul pass, ca. 1600m, 13.V.1970, leg. Zwick (cPut). Locality ambiguous: 14: “Yenikóy, Toros”, 30. VIII.1947 (cPut); 12, same data, but 29.-31.V11.1947 (cPut). Measurements (mm) and ratios (range; n=7): HW: 0.91-0.98; PW: 0.68-0.72; PL: 0.76-0.83; EL: 0.92-1.03; EW: 1.01-1.10; ML: 0.86-0.92; HW/PW: 1.30-1.35; PL/PW: 1.08-1.15; EL/PL: 1.22-1.29; EL/EW: 0.90- 0.94. Comments: This species has been confounded with various other congeners, especially with the following three undescribed species and with S. erythrocnemus. Even the paratypes from Maras are not conspecific with the holotype (see types of S. distortus sp. n.). Among the closely related species, S. anatolicus is identified by the shape of the aedeagus and by the untwisted basal in- ternal structure of the median lobe (Fig. 1). For addi- tional, but less reliable characters see the remarks below the following species. A detailed description of the ex- ternal characters 1s given by PUTHZ (1970a). Distribution: The distribution of Stenus anatolicus is apparently confined to southwestern Turkey from Mugla in the west to central Mersin in the east (Map 1), a distribution pattern not uncommon among Turkish Staphylinidae. Records from other regions refer to one of the following species. 3.2. Stenus (Ss. str.) cypriacus sp. n. (Figs. 2, 7, Map 1) Types: Holotype 4: CYPRUS - S Platres, Moniatis, Bachtal, Bachufer, 700m, 12.1V.1995, Assing / Holoty- pus Í Stenus cypriacus sp. n. det. V. Assing 2002 (cAss). Paratypes: 52 Y: same data as holotype (cAss); ld: same data as holotype, but 07.1V.1995 (cAss); 248, 322: Cyprus, Commanderia, B8 b. Moniatis, 750m, Bachschlucht, 2.-12.4.95, P. Wunderle (cWun); 14, 19: Chypre, 700m, Kalopanayiotis, 17.V11.1977, C. Besuchet (MHNG); 14: CIPRO Val. Cedri, m. 1300m, 25.4.63, Henrot-Cerruti (cPut); 14: Cypr. Stavros, 18.- Map 1: Distributions of Stenus anatolicus Puthz (open circles), S. cypriacus sp. n. (filled circles), and S. distortus sp. n. (black squares) in Turkey and Cyprus, based on examined material. Michael FRANZEN & Walter HEINZ: Morphology and Natural History of Mantica horni 305 19.7.39, Hakan Lindb. / Field Mus. Nat. Hist. 1972, Ludwig Benick Colln. Acc. Z-14, 869 (FMNH); 19: ZYPERN, Mesopotamos, 32°54°/34°53’, 5.5.74, MA- LICKY (cPut); 14: Troodos, 11.6.1963, leg. T-E. Leiler (cWun). Description: Measurements (mm) and ratios (range; n=7): HW: 0.86 - 0.94; PW: 0.65 - 0.69; PL: 0.71 - 0.76; EL: 0.91 - 0.98; EW: 0.98 - 1.06; ML: 0.72 - 0.76; ENW/PW:1.33 - 1.39; PL/PW: 1.09 =:1.16; EL/PL: 1.22 - 1.30; EL/EW: 0.91 - 0.94. External characters as in Stenus anatolicus (Fig. 7), but of smaller average size (see measurements) and antero- lateral tubercles on pronotum mostly obsolete. (In S. anatolicus the tubercles are usually distinct.) Bases of femora (as in S. anatolicus) usually distinctly lighter (yellowish to reddish brown) than apical halves. 3: secondary sexual characters similar to those in S. anatolicus: sternite VI slightly flattened and with weakly concave posterior margin; sternite VII with rela- tively long and dense testaceous pubescence, shallowly, but extensively impressed in posterior half, posterior margin broadly and distinctly concave; sternite VIII weakly impressed before the broadly concave posterior margin; sternite IX similar to that of S. anatolicus; aedeagus of similar general morphology as in S. anato- licus, but distinctly smaller (see measurements and Fig. 2), apically more abruptly narrowed, and basal internal structures of completely different shape (Fig. 2). Etymology: The name is an adjective and refers to the fact that the species is apparently endemic to Cyprus. Comparative notes: In the related species with a simi- larly shaped aedeagus (S. anatolicus, S. distortus, S. prominens), the median lobe of the aedeagus is dis- tinctly larger (see measurements and figures), the basal internal structure of the internal sac is of completely dif- ferent shape (S. anatolicus) or smaller and more slender (S. prominens, S. distortus), the antero-lateral tubercles on the pronotum are usually more distinct (S. anato- licus, S. distortus), the size is on average larger (see measurements), and the femoral bases are not distinctly lighter than the apices (S. distortus). Distribution: Stenus cypriacus is presumably endemic to Cyprus (Map 1). It is known from several localities, but according to personal observations it is not abun- dant. The records of S. guttula Miiller in BAUDI DI SELVE (1870) and of S. anatolicus in ASSING & WUNDERLE (2001) refer to this species. 3.3. Stenus (s. str.) distortus sp. n. (Fig. 3, Map 1) Stenus anatolicus Puthz, 1970a (PUTHZ 1970a: 22ff) partim. Types: Holotype 4: TR — bor., Torul, 9.6.1994, leg. Scoupy / Holotypus ¢ Stenus distortus sp. n. det. V. Assing 2002 (cAss). Paratypes: 15, 49 9: same data as holotype (cFel, cAss); 14: Turkey, 2.8.86, Macka -> Sumela, leg. Feldmann (cFel); 14, Quelle am Tunnel vor Siirgu, Taurus / 13.V.1970, Zwick leg. / SO- Anatolien, Ost-Taurus bei Sürgü, 13.V.1970, Zwick (cPut); 2¢¢, 399: Anatolia mer., Heinz leg. / Höhle an z.T. subterranem Fluß, ca. 61 km südl. v. Bingöl, 9.VI1.1974 (cPut, cAss); 14: Türkei, Taurus, Bolkar Daglari, Umg. Ciftehan (Flußufer, ca. 1500 mNN), 26.V1.1990, leg. Zeuner (cPut); 14: ca. 80km NW Maras, Tiirkei, Zwick leg. / 16.V.1970, Bach a. d. Schneegr. / ANATOLIEN, 10.-22.5.1970, leg. Zwick (cPut); 14, Türkei, 17.5.1969, Maras, leg. Wewalka (cPut); 19, 329: TURKEY: 14.-19.vii.2000, Adiyaman prov.; Nemrut Dagi Mts., Karadut env., I. Smatana leg. (cSch, cAss); 14, 19: Maras, 17.5.69, leg. Wewalka / Paratype / Stenus anatolicus n. sp. det. V. Puthz 1970 (MHNG). Figs. 1-6: Aedeagus of Stenus anatolicus Puthz (1), S. cypria- cus sp. n. (2), S. distortus sp. n. (3), S. prominens sp. n. (4), S. erythrocnemus Eppelsheim (5), and S. maculiger Weise (6). Scale: 0.2 mm. Description: Measurements (mm) and ratios (range; n=7): HW: 0.91 - 1.00; PW: 0.71 - 0.79; PL: 0.77 - 0.89; ER 0.95. 21209 We 1206 = 1:16: MIL: 0.83 = 0.94: 306 Bonner zoologische Beitráge 53 (2004) HW/PW: 1.25 = 1,35; PL/PW: 1.06 - 1.17; EL/PL: 1.19 - 1.29; EL/EW: 0.88 - 0.94. Externally highly similar to S. anatolicus, but femoral bases usually not or only indistinctly lighter than apıcal halves and pronotum on average wider (see measure- ments). Í: aedeagus of similar shape and size, but basal internal structure twisted, of completely different shape (Fig. 3). Etymology: The name (Lat., adj.: twisted) refers to the shape of the basal internal structure of the aedeagus, which readily separates this species from S. anatolicus, with which it was previously confused. Comparative notes: For separation from S. anatolicus see description. S. cypriacus has a smaller aedeagus with a smaller and more slender basal internal structure; in addition, the body is on average smaller (little over- lap, see measurements). In S. prominens, the apex of the median lobe of the aedeagus clearly extends beyond the parameres. In both S. cypriacus and S. prominens the femora are basally lighter and the antero-lateral tuber- cles on the pronotum are indistinct or obsolete. Distribution: Stenus distortus is known from northern, central southern, and eastern Anatolia (Giimiishane, Trabzon, Malatya, Bingól or Diyarbakir, Nigde); its known distribution does not overlap with that of S. ana- tolicus and $. erythrocnemus (Map 1). 3.4. Stenus (Ss. str.) prominens sp. n. (Fig. 4) Types: Holotype ©: LIBANON, Touaite / Holotypus © Stenus prominens sp. n. det. V. Assing 2002 (cPut). Pa- ratypes: 19: Libanon, V1.1995, Ehden, 1500m, Horch Ehden, Bach, leg. C. Reuter (cFel); 14: ISRAEL, Ba- nias, 1.V11.1986, A. Shlagman / G. Haghebaert coll. & det., Stenus s. str. guttula Miill. (UH). External morphology as in S. anatolicus, but with indis- tinct or obsolete antero-lateral pronotal tubercles and with basal halves of femora testaceous to rufous. @: median lobe of aedeagus large, apically distintly extend- ing beyond parameral apices; basal internal structure twisted (Fig. 4). Description: Measurements (mm) and ratios (holotype, 3 paratype): HW: 1.04, 1.01; PW: 0.77, 0.76; PL: 0.85, 0.85; EL: 0.98, 1.00; EW: 1.07, 1.00; ML: 0.94, 0.94; HW/PW: 1.35, 1.34; PL/PW: 1.10, 1.12; EL/PL: 1.16, 1.18; EL/EW: 0.92, 0.92. Etymology: The name (Lat., adj.: protruding, exceed- ing, standing out) refers to the long apex of the median lobe of the aedeagus, a character distinguishing this species from its closely related congeners. Comparative notes and discussion: From the other Eastern Mediterranean representatives of the S. eryth- rocnemus goup, S. prominens is distinguished especially by the apically long median lobe of the aedeagus, and by the shape of the basal internal structure. In addition, it is separated from S. anatolicus, S. distortus, and S. ervthrocnemus by the indistinct or obsolete antero- lateral pronotal tubercles, from S. distortus and S. eryth- rocnemus by much lighter femoral bases, and from S. cypriacus by larger size. Distribution: Stenus prominens is known from three localities in Lebanon and Israel. Fig. 7: Stenus cypriacus sp. n. 3.5. Stenus erythrocnemus Eppelsheim (Fig. 5, Map 2) Type examined: Syntype Y: Lenkoran, Leder (Reitter) / 15. / 10. / c. Epplsh. Steind. d. / erythrocnemus Epp. Typ. / TYPUS (NHMW). Additional material examined: Turkey: Artvin: 534, 729, Artvin-Savsat, 4.V1.1989, leg. Schónmann & Schillhammer (NHMW, cPut, cAss); 13, Borcka, 27.V1.1970, leg. Schubert (NHMW); 384, Michael FRANZEN & Walter HEINZ: Morphology and Natural History of Mantica horni 12, Borcka, 1700m, 18.-27.V1.1970, leg. Schubert (NHMW, cPut); 644, Borcka, VII.1971, leg. Schubert (NHMW); 54, 429, Borcka, VII.1974, leg. Schubert (NHMW, cPut, cAss); 1¢, pass between Murgul and Arhavi, 1000-1200m, 17.VII.1972, leg. Heinz (cPut). Rize: 1°, Ardesen, Ilica, 20.V.1970, leg. Zwick (cPut); 346, 329, Firtina valley, 1300m, leg. Vit (MHNG, cPut, cAss). Locality not identified: 13, Karckal — Ot- ingo, 1200-1600m, 10.-12.V11.1983, leg. Heinz (cPut). Georgia: 14, Tbilisi, leg. Leder (Reitter) (cPut); 244, 62°, Zchneti near Tbilisi, 20.VII.1985, leg. Wrase (cPut, cSch). Abkhazia: 1¢, Avadkhara, 2000-2800m, 8.-13.V11.1982, leg. Wrase & Schüler (cSch). Armenia: 1¢, 12, Gegemskij Chr., Geghard, 2000m, 3.V11.1989, leg. Pütz (cPut). Iran: 938, 1029, Ilam, Tchaharmelleh, 33°57N, 46°17E, 28.V1.1974, leg. Senglet (MHNG, cPut, cAss); 13, Teheran, above Shemshale, 2400-2800m, 18.V111.1967, leg. Heinz (cPut); 14, 12, Kerman prov., 1964, leg. Löffler (cPut); 1¢, Kermanshah, Kenesht/ Kermänshäh, 34°29N, 47°09E, 3.V111.1973, leg. Senglet (cPut); 274, Fars, ca. 50 km W Shiraz, Dasht-e-Arzhan, 1900m, 21.1X.1997, leg. Schódl (NHMW); 14, 19, Fars, 30 km N Nurabad, 25.1V.1996, leg. Wewalka (NHMW ); 14, Fars, Dasht-e-Arjan, 55km NSSK W Shiraz, 22.1V.1993, leg. Pour (cPut): 14, 12, Iam, Saräb Eyvän, 33°45N, 46°22E, 26.V1.1974, leg. Senglet (cPut); 434, 429, Buyer Ahmad o Kuhgiluye, 30km S Yasug, spring, 25.1V.1996, leg Wewalka (NHMW, cPut, cAss); 14, Qars-1-Shirin, “Fiume ad W terr. nudo limo”, 7.1V.1956, leg. Soika (NHMW); 14, SW-Iran, Hoseiniyeh, 28km NNW Andimeshk, 360m, 12.-13.1V.1977 (cAss); 19, Gi- lan, Lunak, SW Langrad, 700m, 2.V11.1998, leg. Heinz (cPut); 14, 12, SE-Iran, Transhar [?] (NHMW, cPut). Pakistan: 244, Chitral, Bumburet, 2200-2350m, 24.V.1983, leg. Besuchet & Lóbl (cPut). Afghanistan: 1¢, Nurestan, SW Peé-Dara, 1200m, 9.V11.1971, leg. Kabakov (cPut). Measurements (mm) and ratios (range, arithmetic mean; n=16): HW: 0.89 - 0.98, 0.93: PW: 0.66 - 0.74, OTIESPE: 0:74 - 0.86,. 0.79; ML: 0.66 - 0.77, 0.72; HW/PW: 1.25 - 1.38, 1.32; PL/PW: 1.04 - 1.20, 1.12 Comments: Stenus erythrocnemus is readily distin- guished from all the preceding species by the differently shaped median lobe of the aedeagus, which is apically much more abruptly narrowed (Fig. 5). In addition, the elytra are often shorter and narrower, the hind wings sometimes of reduced length (less than twice the length of elytra), and the femoral bases are mostly dark (in this respect similar to S. distortus). The Iranian material dif- fers from the specimens seen from northeastern Anato- lia and the Caucasus region by the longer apex of the median lobe and the somewhat more slender and more 307 abruptly bent apex of the basal internal structure of the aedeagus. Slight differences were also observed in the material from Pakistan. However, until the distributions of these morphs are clarified, these differences are here tentatively interpreted as an expression of clinal intra- specific variation. 3 Distribution: The species is distributed from northeast- ern Anatolia (where its range borders on that of S. dis- tortus) and the Caucasus region to Iran, Afghanistan, and Pakistan (Map 2). 3.6. Stenus maculiger Weise (Figs. 6, 8, Map 3) Material examined: Austria: 1 ex., Niederösterreich, Oberndorf, Pfoisau, 3.V1.1972, leg. Rausch (cPut); 14, Kärnten, Zell, Huda- jamagraben, 25.V1.-4.V11.1987, leg. Wunderle (cWun); 14, Kärnten, Eisenkappel, 1.VIII.1942, leg. Scheerpeltz (MHNG); 12, Eisenkappel, 600m, 9.VII.1941, leg. Scheerpeltz (MHNG). Italy: 5¢¢, Camia, Rio Tagliezzo, 8.&29.X1.1953, 4.V11.1954, 19.11.1957, leg. Springer (cPut); 19, Car- nia, Rio Squalon, 15.V111.1954, leg. Springer (cPut). Slovakia: 17, Nova Sedlica, 15.VII.1961, leg. Smetana (cPut). Hungary: 14, Mecsek range, leg. Gebhardt (cPut). Croatia: 324, Dubrovnik, Ombla spring, 8.VIII.1958, leg. Endrödy-Younga (cPut); 894, 19, 1 ex., SE Du- brovnik, Mlini, 27.IX.-4.X.1968, leg. Benick (MHNG, cPut); 12, Dubrovnik (cPut). Bosnia-Herzovina: 324, Jablanica (cPut); 14, Foca, leg. Apfelbeck (Put); 12, Kladanj, 1909, leg. Leonhard (MHNG); 19, "Herzegowina", leg. Reitter (MHNG). Yugoslavia: Montenegro: 17, Mrtvica near Manastir Moraéa (cPut); 729, Manastir env., Morata (cPut); 234, Milocet, 4.V.1965, leg. Leiler (cPut); 19, 10km E Tresnjevik pass, near Andrijevica, 25.V.1979, leg. Zwick (cPut). Bulgaria: 19, Nesebar, Vlas, 17.-26.VII.1983, leg. Wrase & Schülke (cSch); 17, Eminska planina, Vlas, 12.V.1987, leg. Behne & Heinig (cSch); 12, SW- Bulgaria, Sandanski, 27.-28.1V.1985, leg. Wrase (cSch); 244, SW-Bulgaria, Samokov, 4.V.1966, leg. Löbl (cPut); 14, Rila, Bodrovec, 31.V111.1966, leg. Löbl (cPut); 14, SW-Bulgaria, Gospodinci, 3.V.1966, leg. Lóbl (cPut); 244, 19, Pirin, leg. Weirather (MHNG); 14, Sliven, X.1952, leg. Roubal (MHNG); 19, Sliven, 27.V1.1908, leg. Rambousek (MHNG); 14, Rhodopi, Pamporovo, 1500m, 28.V.1964, leg. Schulze (cPut): 19, Rhodopi, 5km N Schitoka lake, 41°42N, 24°36E, 22.V1.1980, leg. Malicky (cPut); 19, Katundere, 42°04N, 27°27E, 15.V1.1980, leg. Malicky (cPut). 308 Bonner zoologische Beitráge 53 (2004) Map 2: Distribution of Stenus erythrocnemus Eppelsheim, based on examined material. The easternmost records are in Pakistan and Afghanistan. Albania: 244, Lumi i Bences, Tepelene, V.1931, leg. Winkler (cPut); 14, Dukati (cPut); 1¢, Mirdita, leg. Winneguth (cPut); 24, Latif, Orosi (cPut); 14, Latif, Cukali (cPut). Greece: mainland: 2°99, ns 39°50N, 21°01E, 5.VI.1975, leg. Malicky (cPut); 13, Ladıkon. > 21°40E, 23.V.1987, leg. Malicky oo 238, Y, Serres, Hagiai Ioannis, moss, 17.VIII.1964, leg. ae (cPut); 19, Penday, 38°35N, 22°04E, 3.V1.1975, leg. Malicky (cPut); 14, Iti, below Peristeri, 38°51N, 22°10E, 500m, 11.V1.1987, leg. Malicky (cPut); 14, Timfristos, 2000m, 8.VII.1982, leg. Osella (cPut); 14 Voiotia, Levadia, moss, 4.1X.1964, leg. Puthz (cPut); 12, Pelion, 25.11.1981, leg. Wunderle (cWun); 14, Pangéo, 26.V11.1988, leg. Jách (cPut). Pelopónnisos: 344, 29°, Erimanthos, above Kalen- dzi, 3737.02N, 21°46" 34E. 1200m, stream bank, 27.11.1997, leg. Wunderle, Assing (cWun, cAss); 390, 27%, 8km NE Kalavrita, bank of Vouraikos river, 38°04 34N, 22°09'43E, 700m, 30.111.1997, leg. Assing (cAss); 473, 729, Parnon range, 1100m, spring, wet moss, 13. VI. 1996, leg. Wunderle (cWun, cAss); 230, Lakonía, Parnon, Polidroso, NE Sparti, 37°10°29N, 22°36'02E, 990m, waterfall, wet moss, 19.1V.1998, leg. Zerche (cAss). Levkäs: 1, Vafkeri, 450m, 26.1X.1993, leg. Assing (cAss). Andros: 13, 19, above Sarisa, 37°SIN, 24°54E, 20.1V.1984, leg. Malicky (cPut). Thassos: 14, Potamias, 13.1X.1991, leg. Jäch (cPut). Andros: 19, N Andros, 37°51N, 24°56E, 12.V1.1979, leg. Malicky (cPut). Samothráki: 444, 629, near Kremniotissa church, 40°25N, 25°34E, 24.V.1987, leg. Malicky (cPut); 19, 3 km SE Tharma, 14.V1.1993, leg. Jách (cPut). Samos: 12, W Karlovasi, 37°48N, 26°41E, 27.V.1979, leg. Malicky (cPut); 19, 2km-NE Kalithea, 37°45N, 26°36E, 26.V.1979, leg. Malicky (cPut). Sky- ros: 12, Ikm ih EN 28.1X.1992, leg. Jäch (cPut). Ikaria: 334, 599, W Chrisostomos, 37°35N, 26°13E, 2.VLI979; leg: Malicky (cPut, cAss); 222, W Mileo- pon, 37°39N, 26°14E, 2.V1.1979, leg. Malicky (cPut). Rhödos: 14, Petaloudes, 17.V.1966, leg. Leiler (cPut); 24, Farfalle valley, 12.V.1970, leg. Cerruti (cPut); 14, Apollona, [V.1966 (cPut); 1%, Salakos, 15.1V.1977, leg. Besuchet (MHNG ); 19, Epta Pigai, 9.1V.1977, leg. Besuchet (MHNG); 234, 2 exs., Kolimbia, 24.1V.1973, leg. Besuchet (MHNG). Ukraine: Crimea: 744, 59 Y, Crimeas reserve, upper part of Katcha a an 10.V.2000, leg. Ivanov (cPut, cSch, cAss); , Baidar, leg. Knirsch (cPut); 338, 229, laila ate leg. Winkler (MHNG, cPut); 19, sinferopal (MHNG). Turkey: Istanbul: 744, 19, Istranca Dag near Demirköy, pass, 700m, 12.VII.1972, leg. Heinz (cPut); 13, Beykoz, 1902, leg. Cameron (cPut); 14, Sile, 19.V.1987, leg. Schönmann & Schillhammer (cPut); 14, 19, Kilyos (FKumkoy), 27.V11.1969, leg. Besuchet (MHNG). Bolu: 14, forest between Bolu and Yigilca, 1100m, 18.V1.1987, leg. Heinz (cPut) 538, 799, Konuralp-Akcakoca, 400m, 15.V.1976, leg. Besuchet & Lóbl (MHNG, cPut); 14, Omerler near Bolu, 800m, 21.V.1976, leg. Besuchet & Löbl (MHNG). Gü- müshane: 14, Torul, 9.V1.1994, leg. Scoupy (cFel). Trabzon: 14, Macka, 1000m, VII.1971, leg. Schubert (cPut). Adiyaman: 534, Nemrut Dagi, Karadut, 14.- 19.V11.2000, leg. Smatana (cSch, cPut, cAss). Diyar- bakir: 323, 329, 6lkm S Bingól, cave near partly subterranean river, 9.VII.1974, leg. Heinz (cPut, cAss). Bitlis: 13, S Tatvan, 1700-2000m, 21.V.-18.V1.73, leg. Schubert (cPut). Hakkari: 17, N Semdinli, 3.V1.1987, leg. Schönmann & Schillhammer (cPut). Mersin: 19, Haciiskakli, 27.1V.1978, leg. Besuchet & Löbl Michael FRANZEN & Walter HEINZ: Morphology and Natural History of Mantica horni 309 (MHNG). Isparta: 17, Anamasdagi, leg. Weirather (cPut). Burdur: 244, 19, Celtikci, 850m, 5.V.1975, leg. Besuchet & Löbl (MHNG). Antalya: 13, 399, 30km N Finike, W Catallar, 23.5.1991, leg. Schónmann & Schillhammer (cPut); 19,29 9, SSW Antalya, Ciralı, 36°25 N, 30°28’E, stream bank, 04.1V.2002, leg. As- sing & Wunderle (cAss, cWun); 19, 3 km W Cirah, 36252 7N, 30°26’°41E, 71m, stream bank, 28.111.2001, leg. Rose (cRos) 19, E Kumluca, 36°21’50N, 30°22°27E, 385m, stream bank, 3.1V.2002, leg. Assing & Wunderle (cAss); 12, same data, but 25.111.2002 (cAss); 13, N Kumluca, Altinyaka, 24.5.1991, leg. Schónmann & Schillhammer (cPut); 14, E Antalya, S Akseki, Topraktepe, 30.V.1983, leg. Brachat (cPut); 238, 19, Alanya, Kuzyaka, 180m, stream bank, 12.111.2000, leg. Rose (cRos); 13, 2km NW Antalya, 363123N, 30°00°43E, 795m, moist locality with gravel, under bridge, 26.1112001, leg. Rose (cAss); 238, 299, Catallar, 36%29"13N, 30%04'56E, 370m, stream bank, 26.111.2001, leg. Rose (cRos); 14, 19, Skm E Büyükalan, 36°43’29N, 30°21°13E, 1352m, roadside, near stream, 27.11.2001, leg. Rose (cRos, cAss) 1, lkm N Ségiitcumavi, 36°41’51N, 30°22°30E, 1463m, stream bank, 29.111.2001, leg. Rose (cRos); 12, N Sagırın, Koprülü Canyon, 37%0437N, 31°13’56E, 30m, stream bank, 17.11.2002, leg. Rose (cRos); 12, 5km NW Kemer, 36°36’03N, 30°29’04E, 197m, river bank, 30.111.2001, leg. Rose (cRos). Mugla: ld, Baba Dag ["Salbakos Gbg."], leg. Weirather (MHNG). Izmir: 1%, Izmir (MHNG). Comparative notes: Stenus maculiger can be distin- guished from S. anatolicus, S. erythrocnemus, and re- lated species by a completely different shape of the aedeagus (shape and internal structures of median lobe, parameres extending distinctly beyond apex of median lobe, by the presence of a (sometimes very weak or in- distinct) median furrow on the pronotum, the much lighter basal halves of the femora, and by the absence of antero-lateral pronotal tubercles (usually absent also in S. cypriacus and S. prominens). Stenus sacrimontis Puthz and S. bosnicus Bernhauer have an aedeagus of highly distinctive shape and with completely different internal structures; for illustrations see PUTHZ (1970b). Distribution and intraspecific variation: As is illus- trated in Map 3, S. maculiger is a typical example of a Ponto-Mediterranean faunal element. Based on a study of the internal structures of the aedeagi of specimens from various regions, two morphs were recognized, the distribution of one of them ranging from the southeast of Central Europe to Ukraine, southern Greece (includ- ing many islands), and Northern Anatolia, and that pf the other confined to southern Anatolia (Map 3). The former morph has an aedeagus with an apically long and slender basal structure (Fig. 8, left); in addition, the elytra are often short and only slightly wider than the pronotum, and the hind wings are frequently of reduced length. In the latter morph, in contrast, the basal internal structure is apically stouter and shorter (Fig. 8, middle), the elytra are always distinctly wider and longer than the pronotum, and the hind wings are fully developed. One of the dissected males from Antalya (southern Ana- tolia), however, was found to have an internal structure (Fig. 8, right), which is apically almost as long and slender as in males e.g. from Greece, suggesting that the observed difference in the shape of the basal structure may not be constant. Both morphs may still represent species or subspecies, but in order to clarify the taxo- nomic status and rule out the possibility that the differ- ences are merely an expression of intraspecific varia- tion, more material especially from southern Anatolia must be examined. Fig. 8: Stenus maculiger Weise. Aedeagi of males from Greece (left) and southwestern Turkey (middle and right). 310 Bonner zoologische Beitráge 53 (2004) Map 3: Distribution of Stenus maculiger Weise, based on examined material, including four records from Austria examined and communicated to me by Volker Puthz. Filled circles: morph with long basal internal structure (Fig. 8, left); open circles: morph with short basal internal structure (Fig. 8, middle); grey circles: records represented only by females. Acknowledgements. My thanks are due to all the col- leagues indicated in the material section for the loan of ma- terial. In particular, | am grateful to Volker Puthz for sev- eral helpful discussions and for contributing unpublished data. 4. REFERENCES ASSING, V. & WUNDERLE, P. 2001. On the Staphylinidae of Cyprus (Coleoptera). Entomologische Zeitschrift 111(2): 34-41. BAUDI DI SELVE, F. 1870. Coleopterorum messis in insula Cypro et Asia minore ab Eugenio Truqui congregatae recensitio: de Europaeis notis quibusdam additis. Ber- liner Entomologische Zeitschrift 13 (1869): 369-418. PuTHZ, V. 1970a. Stenus (s. str.) anatolicus nov. spec aus der Türkei (Coleoptera, Staphylinidae). Mitteilungen der Deutschen Entomologischen Gesellschaft 29: 22-24. PUTHZ, V. 1970b. Ein neuer griechischer Stenus aus dem Deutschen Entomologischen Institut Eberswalde (Co- leoptera, Staphylinidae). Beitrage zur Entomologie, Berlin 20: 589-593. Author’s address: Dr. Volker ASSING, Gabelsberger- straße 2, D-30163 Hannover, Germany; E-mail: vassing. hann(@t-online.de Received: 28.06.2003 Accepted: 10.12.2004 Corresponding editor: M. Schmitt Bonner zoologische Beitrage Band 53 (2004) | Heft 3/4 | Seiten 311-321 Bonn, Dec. 2005 Reproductive Types and Mobility of Carabid Assemblages: Effects of Landuse Intensity of Extensively Managed Orchards Jürgen DEUSCHLE” & Erich GLUCK” .” Kóngen, Germany 2 Donzdorf, Germany Abstract. From April 1995 to November 1997 investigations were carried out in orchards (48.36N/9.23E): data were collected on the type and frequency of grassland use and on the carabid fauna. Seventeen sample plots were selected. Part of the plots had 25 years of unchanged management regimes: three-cutting meadows (3), two-cutting meadows (3), mulched meadows (4), abandoned meadows (3), a horse pasture, a sheep pasture with rotational grazing, a continuously grazed sheep pasture, and a sheep pasture abandoned in 1994. A total of 68 carabid species was registered. Two-cutting meadows (20-24 species), three-cutting meadows (17-30 spe- cies), and pastures (16-27 species) showed a significantly higher diversity than mulched meadows (13-19 species), and abandoned plots (7-18 species). The diversity and activity densities on the studied plots decrease with lower land use in- tensity. Reproduction: Spring breeders. were more frequently found on sample plots than autumn breeders (63-87% to 12-38%) or species reproducing all year (0-11%). Increased shading by fruit trees lead to a decrease of the percentage of spring breeders. Type of alae: The percentage of macropterous species was between 29 and 69%. Alae dimorphous species showed a percentage between 0 and 40%, and for brachypterous species the percentage was between 10 and 71%. The percentage of brachypterous species diminished along the land use gradient, while the percentage of alae dimorphous species in- creased. Key words. Ground beetle, land use systems, mobility, reproductive types, species community. 1. INTRODUCTION Vegetation and fauna of extensively managed orchards are mainly determined by the site and its maintenance, the type of grassland management and its land use in- tensity (DEUSCHLE et al. 2002). Until now animal as- semblages of extensively managed orchards have not been analysed based on the direct comparison of different forms of grassland management. Forms like continuous and rotational grazing (sheep, horses) have not been taken into account. There are few investigations dealing with the influence of management systems on arthropod assemblages in extensively managed grassland, whereas the effects of grazing have been documented more fre- quently (HANSSEN & HINGST 1995; MAELFAIT et al. 1988; RUSHTON et al. 1989; SCHNITTER 1994). Normally, orchards are regarded as ecologically high valid systems, but it is yet not known if and to what ex- tent the management influences carabid assemblages. Is there an influence on the number of species or densities or on the capability for large or small species? Further- more, it is asked whether resources are equal or ran- domly distributed in the orchard area or overwhelmed by the management regimes of the individual plots, thus the captured individuals show differences in their occur- rence due to different management forms on the basis of their ecological characteristics. Larval and adult hibernation are the two basic forms of reproductive strategies of carabids (autumn breeders, spring breeders). Differences in the reproductive strate- gies allow the carabids to live in spatial coexistence by exploiting the same resources at different times (MULLER 1987). Eurytopic woodland species are less dependent on specific reproductive forms (RODE 1993; OTTESEN 1996). The capability of carabids to disperse is regarded as an important prerequisite for the survival of the species un- der frequently changing external conditions (DE VRIES et al. 1996). This holds especially true for the cultural landscape (DEN BOER & VAN DIJK 1994; RODE & DULGE 1994). It is known that macropterous individuals inhabit dynamic, continuously changing habitats in open cultural landscape. The species are often very small and have a high potential of dispersion for the colonisation of new habitats. But their populations are therefore ex- posed to a significantly higher risk of extinction (DEN BOER 1987; DINGLE 1996). Brachypterous species are thought to have a smaller po- tential of dispersion. Especially eurotopic woodland species are brachypterous and in comparison they are quite mobile. From their original habitat they can mi- grate to explore and colonise suitable new habitats (GLUCK & KREISEL 1986). Woodland species are re- garded to be typical inhabitants of habitats with nearly constant living conditions. During the advancement of succession they are found more frequently in the spec- trum of species (BUREL & BAUDRY 1994; POLLARD 1968). So far the ecological characteristics of the species be- longing to the carabid assemblage should be different if the beetles react to resources (GLUCK & DEUSCHLE 2003). The capability of using the resources should then be reflected in their occurrence on different plots. The aim of this investigation is to demonstrate how dif- ferent management systems (meadows: two or three cut- tings a year, pastures: continuous and rotational grazing with horses and sheep, mulched meadows, abandoned meadows) have influence on the mobility of the carabid assemblage. Mobile species should be more common under rapidly changing conditions like intensive mowing. It will be examined, if the reproductive strategies (spring breeders/autumn breeders) of carabid assemblages are influenced by the management system in specific areas. It will also be shown, which type of management fa- vours brachypterous or macropterous individuals of cer- tain species. The advantages and disadvantages of re- Table 1. Morphology and land use of sampled plots Bonner zoologische Beitráge 53 (2004) productive strategies and increased mobility will be discussed considering the small-patterned mosaic struc- ture of extensively managed orchards with its inherit changing living conditions. 2. METHODS 2.1. Census and registration of land use in the orchards From the beginning of April until mid October 1997 in the study area in the nature reserve “Limburg” (48.38N/9.23E, Fig. 1, Table 1), southwest Germany, the following parameters were recorded weekly or every two weeks: type of use, time of mowing, “mowing de- vice”, “whereabouts” of mowed grass, and number of grazing animals. The amount of ground shaded by trees under perpendicular solar radiation was estimated and assigned to the following categories (0, 1-20, 21-40, 41- 60, 61-80, 81-100%). All areas, which are mowed com- pletely on a regular basis every year and where the cut grass 1s removed, will be called “typical meadows”. On “mulched meadows” the cut grass remains on the areas. “Continuous grazing” defines the management type of pastures, where the grazing animals can be found per- Plot Land use Type of Area Circumference Altitude Shading Sampling land use (Ar) (m) (mNN) (%) period 3CM2 3-cutting meadow meadow 26 209 405 20 97 3CM3 3-cutting meadow meadow 51 287 425 20 oF, 3CM1 3-cutting meadow meadow 16 204 392 20 99596397. 2CM3 2-cutting meadow meadow 40 446 415 80 7 2CMI 2-cutting meadow meadow 10 218 445 60 195,905 97 2CM2 2-cutting meadow meadow 15 250 410 60 "97 SPA abandoned sheep pasture 34 234 405 60 95596297 pasture (cont. grazing), see text SPC sheep pasture pasture 15 213 415 80 97 (continuous grazing) HP horse pasture pasture 124 580 400 20 9596,97 SPR sheep pasture pasture 11 239 445 20 ‘9596597 (rotational grazing) MMI mulched meadow mulched 12 162 455 20 055 96.97 (4 - 6 cuttings) meadow MMEI mulched meadow mulched 47 380 440 100 Oy (3 cuttings) meadow MME2 mulched meadow mulched 29 217 420 60 ‘95596, 97 (2 — 3 cuttings) meadow MME3 mulched meadow with I mulched 25 206 517 80 il cutting meadow AMR recently abandoned succession 13 153 445 0 95,965.97, meadow (4 years) AMO old abandoned meadow succession 13 202 420 100 9390397 (10 years) WDL woodland succession 13 222 520 100 "97 Jiirgen DEUSCHLE & Erich GLUCK: Reproductive Types and Mobility of Carabids manently on the same area, while “rotational grazing” is the type, where a particular area is grazed for a period of only a few days, but several times a year. Combinations of the three types of management forms can also be 313 found in the extensively managed orchards of the nature reserve area. Management oriented ranking of plots on the basis of land use intensity was calculated (Table 2). SPA CM2 | x | MM \ —É \ AS NN \ eZ, A X JA x SWC MME2 Landuse 1997 Mown crop — hey [I] mulching Pasture sheep, rotational grazing WDL sheep, cont. grazing horse Mixed landuse for grazing and mowing (sheep) mulching and pasturing == mulching and mown crop with clear up mulching, mown crop and pasturing Succession recently abandoned meadow old abandoned meadow ~~ | Woodland 3 miscellaneous [-_] sample plots 50 0 50 100 Meter es Fig. 1: Land use intensity of grassland in the study area during the vegetation period of 1997. 314 Bonner zoologische Beitráge 53 (2004) Table 2. Management oriented ranking of plots. The column rank I scales exclusively the nine in 1995 to 1996 studied plots, rank II scales all studied plots in 1997 (see Figs. 2, 3) Area RankI Rank II Explanation plot (95/96) (97) 3CM2 - 2 3CMI | 2 From all plots here the strongest interference in the ecosystem appeared. Essen- tial characteristic is a high biomass withdrawal in combination with compara- tively frequent mowing. 3CM3 - 2 2CM3 - 5 2CMI 2 5 Same, however on a lower intensity. SPA - 7 The change in management from intensively pasturing in 1995/96 allows no clear allocation in the ranking. The up to now management as mowing pasture allows the position between the categories pastures and mowing meadows. SPC - 8 A nearly throughout the year intensively pasturing entails a high withdrawal of material during the vegetation period. HP 3 9 In dry years the intensive pasturing leads to a high withdrawal of material during the entire vegetation period, whenever not only on etiolation spot a considerable part of the plant material is trampled down and therefore a further mulching is necessary. SPR 4 10 During the yearly grazing, only a small part of biomass is removed, the larger part is trampled down by the animals and remains at the plot. The grazing corre- | sponds to one mowing event on a mulching meadow. 11 MMI 5 11 On the whole area there is no withdrawal of biomass, but frequently mowing happens. The entire material is reduced to small pieces and remains until it is de- cayed as mulching layer on the soil surface. MMEI - 12 Same, but with a lower cutting frequency (3x) and partly with a beam mower. MME2 6 13 Same, but with a lower cutting frequency (2-3x) and partly with a beam mower. MME3 en 14 Same, but with a lower cutting frequency (1x) and partly with a beam mower. AMR 7 15 Since 5 years there is no withdrawal of material and no mowing. AMO 8 16 Since 11 years there is no withdrawal of material and no mowing. WDL - 17 The area is at least 50 years out of management. 2.2. Selection of sample plots and population census of carabidae In 1995 the selection of sample plots took into consid- eration areas which had been under the same manage- ment system for more than 25 years (Fig. 1, Tables 1, 2). Six pitfall traps were placed in a single line 10 m apart on a transect through the centre of each sampling plot. Ethylene glycol (50%) was used as preservative solu- tion, with detergent added to reduce surface tension. A cover made of Perspex (120 by 120 mm) was installed 30 to 50 mm above ground level. In 1995 (5 April to | November) traps were controlled and emptied regu- larly every week. In 1996 (5 May to 4 November) and 1997 (8 April to 4 November) the control interval was two weeks (DEUSCHLE & GLUCK 2001). 2.3. Analysis of data and statistical methods All registered data concerning management practices, vegetation, soil condition, and carabid population were integrated into a database. After examining the neces- sary conditions the following statistical tests were ap- plied: U-test by Mann & Whitney (MWU-test), and chi”-test. The rank correlation was calculated according to Spearman and the coefficients were tested on their significance. 3. RESULTS 3.1. Number of species and activity density of carabids A total of 5229 beetles representing 68 carabid species were caught in pitfall traps during the three years of in- vestigation. Eighteen species were caught only once. Regarding their mean number of species, meadows cut once or twice a year do not differ essentially from pas- tures. But all three management forms show a signifi- cantly higher number of species compared to mulched meadows or abandoned plots (Fig. 2). The number of Jiirgen DEUSCHLE & Erich GLUCK: Reproductive Types and Mobility of Carabids 315 species on the sample plots decreases with lower land use intensity. This trend is significant over all three years of investigation as well as for the cumulative number of species for the years 1995 to 1997. A de- creasing density in the number of individuals along a gradient of land use intensity does not show a consistent annual tendency. The activity densities of the sample plots for the years 1995 to 1997 show significant differences (Chi’-Test, p<0.001, n=9). The mean densities of the number of individuals according to the management type do not differ substantially. Only three-cutting meadows show a tendency towards higher activity densities (Fig. 3, Table 3). Table 3. Differences in mean number of carabid species and individuals on sampled typical meadows, mulched meadows, pas- tures, and abandoned meadows: Level of significance of paired MWU-tests (n.s. = not significant, * = p < 0.05, ** = p < 0.001, *** =n < 0.001) 3-cutt. 2-cutt. Pastures Mulched Areas under meadows meadows (n = 10) meadows Succesion (n = 5) (n= 5) (n = 8) (n= 7) 3-cutt. meadows - 1:8: iS: p < 0,05 ns. 2-cutt.meadows ns. - ns. ns. ns. Number of Pastures ns. ns. - ns. ns. Individuals Mulch.meadows p < 0,05 p < 0,01 p < 0,05 - ns. Areas and Suc. p < 0,05 p < 0,01 p < 0,01 n.s. - Number of species o leur) 2 16 + | 8. o 14. = | 287 10 | ee RO a ER EIE 8 1 _I_ Mean+SD 67 Mean-SD van L_|] Mean+SE ES VEREEEE 1 L E M E 3-cutting meadows pastures abandoned meadows pod O Mean 2-cutting meadows mulched meadows Fig. 2: Mean number of carabid species on sampled two-cutting and three-cutting meadows (each n = 5), pastures (n = 10), mulched meadows (n = 8) and abandoned meadows (n = 7), SE = standard error, SD = standard deviation. 316 Bonner zoologische Beitráge 53 (2004) 3.2. Reproductive types The highest percentages of adult hibernators were found on meadow 3CMI (1995: 87%, 1996: 86%) and on the mulched meadow MMI (1995: 84%). The lowest per- centages in 1997 responded to woodland (61%), as well as to the areas MMEI (62%) and 2CM3 (64%), and in 1996 to the mulched meadow MMI (64%). The areas which were sampled for three years showed differences in the yearly percentages between 2% and 14%. The smallest difference were registered on the horse pasture HP and on the mulched meadow MME2, the highest on the mulched meadow MMI (15%), the pasture SPR (14%), and on meadow 2CM1 (13%, Table 4). Species reproducing the whole year — for example Abax parallelepipedus — were not found on pastures SPA, HP, and SPR during the whole investigation period, while on meadows 2CMI and 3CMI, they were only detected in 1996. They were present on the mulched meadow MME2 and on the abandoned plots AMR and AMO in all three years. The only areas, which were sampled for the first time in 1997 and included reproducing species all year, were the plots 3CM3, 2CM3, SPC, MME3 and the woodland. This group had a low percentage of the total carabid assemblage, the highest appearing in 1996 on the abandoned plots AMR and AMO (14% and 15%, Table 4). The reproductive strategies hardly seem to correlate with parameters like area, perimeter, and altitude of the sample plots. But the percentage of all-year reproduc- tion types increased with decreasing land use intensity during the whole investigation. For the years 1995 and 1997 the correlation was significant (Table 5). The main reason was the high activity density of Abax parallele- pipedus, as well as the lower number of typical carabids of extensively used open lands on mulched meadows and abandoned land. In 1997 the percentage of spring breeders diminished significantly with decreasing land use intensity. A simi- lar trend could not be detected for 1995 and 1996. It is also quite possible that the shading by fruit trees has an effect on the presence of spring breeders. During the three years of investigation the percentage of spring breeders decreased with increasing shading. The corre- lation is significant for 1997 (Table 5). 600 + pasos 500 + 5 400) , = o = & 300! o < > > —— 200 ME | O E 100 A A rs A A O ann A ey vibes ds _T_ Mean+SD e Mean-SD [—] Mean+SE 0 FT | E | Mean-SE 3-cutting meadows pastures abandoned meadows O Mean 2-cutting meadows mulched meadows Fig. 3: Mean density of carabid individuals on sampled two-cutting and three-cutting meadows (each n = 5), pastures (n = 10), mulched meadows (n = 8) and abandoned meadows (n = 7), SE = standard error, SD = standard deviation. Jürgen DEUSCHLE & Erich GLUCK: Reproductive Types and Mobility of Carabids 317 3.3. Mobility and Dispersion Forty of the total of 68 species caught in pitfall traps were macropterous and 16 were brachypterous species. Twelve additional species do not show a constant de- velopment of the alae and are therefore considered to be (alae) dimorphous. Brachypterous species are especially relevant, because their distribution is exclusively related to ground movement. The percentage of species in this group varied between 10% on sheep pasture SPA (1995 — 1997), and 71% on the abandoned plot AMO (1996). All meadows showed a smaller percentage. On pastures and mulched meadows it was generally higher. Espe- cially on plot AMO, which was in a stage of succession, and in woodland, species of flightless carabids were quite frequent, while on the recently abandoned plot AMR few brachypterous species were found (Table 6). In all three years the percentage of brachypterous spe- cies increased significantly with decreasing land use in- tensity. The smaller the perimeter of a sample plot, the less individuals of these species were found. On the other hand the percentage of alae dimorphous and com- pletely macropterous species had the tendency to de- crease with lesser land use intensity. In 1995 and 1996 the corresponding correlations for all flying species and in 1997 also for alae dimorphous species were high and significant (Table 7). The alae morphology of all beetles caught was exam- ined. Only Bembidion properans presented the macrop- terous and brachypterous forms (Table 8). Females were significantly more frequent than males (sex ratio 1:2.1 (m: f) MWU - Test p < 0.05). The sex ratio of macrop- terous beetles was 1: 3.2 (m : f). In the population of Table 4. Dispersion strategies of carabids on sampled plots with different management regimes 3CM1, 2CM1, SPA, HP, SPR, MMI, MME2, AMR and AMO during the years 1995 to 1997 and 1997 only (species percentages) Macropterous Dimorphous Brachypterous 3CM1 1995 60.9 26.1 13.0 1996 58.8 23.3 17.6 1997 47.6 33.3 19.0 2CMI 1995 66.7 16.7 16.7 1996 66.0 25.0 15.0 1997 60.0 20.0 20.0 SPA 1995 33:6 EE 11:] 1996 60.0 30.0 10.0 1997 40.0 35.0 25.0 HP 1995 50.0 31.8 18.2 1996 45.0 40.0 15.0 1297 38.3 23:0 16.7 SPR 1995 33.0 22.2 27.8 1996 99.6 22.2 22:2 1997 50.0 31.3 18.8 MMI 1995 52:6 21:1 26.3 1996 43.8 25.0 3 1997 56.3 250 18.8 MME2 1995 47.4 15:8 36.8 1996 33.3 6.7 40.0 [997 31.3 18.8 43.8 AMR 1995 32.9 17.6 29.4 1996 58.8 11.8 29.4 1997 50.0 25.0 23.0 AMO 1995 41.7 8.3 50.0 1996 28.6 0 71.4 1997 45.5 18.2 36.4 3CM2 1997 47.6 38.1 14.3 3CM3 1997 50.0 36.7 13:3 2CM3 1997 50.0 31.8 18.2 2CM2 1997 52.6 26.3 vA | SPC 1997 99.0 23.9 18.5 MMEI 1997 69.2 Tel 23.1 MME3 1997 52.6 2161 26.3 WDL 1997 61.1 0.0 38.9 318 Bonner zoologische Beitráge 53 (2004) Table 5. Rank correlation coefficients regarding percentage of carabid species of different reproductive strategies with area pa- rameter (1995/96: n= 9; 1997 n= 17; level of significance: * =p < 0.05, ** = p<=0.01, ***=p<0.001) Portion of species Land use Area Perimeter Altitude Shading 1995 Spring breeders -0,38 -0,58 -0,25 0,26 -0,54 Autumn breeders -0,15 0,56 0,76% -0,38 0,56 All-year reproductive 0,84% 0,05 O57 - 0,09 inal le 1996 Spring breeders -0,09 0,04 0,10 -0,25 -0,56 Autumn breeders -0,20 0,10 0,27 0,19 0,20 All-year reproductive 0.45 0,02 -0,53 PLA 1997 Spring breeders -0,65** 0,12 -0,19 -0,48 OOPS Autumn breeders 0,31 -0,07 0,40 0,33 0,47 All-year reproductive 0,64** -0,10 -0,29 0,33 0,42 surveyed B. properans, females seem to be more alate than males. A total of 25% of all males and 38% of all females were macropterous. Bembidion properans was only detected on three-cutting meadows and pastures. But it was present on all plots of this type. Only a few macropterous males were caught on sheep pastures. The presence of B. properans on typical meadows and on the horse pasture was much higher. Females were found more frequently than males (Table 8). 4. DISCUSSION Sixty-eight species of carabids were found on the sam- ple plots under different conditions of land use intensity. The analysis of the reproductive types (adult hiberna- tors, larval hibernators, and all year reproductive spe- cies) show that the percentage of spring breeders per plot decreases only slightly with diminishing land use intensity and shading (land use 1995: r= -0.38, 1996: r= -0.09, 1997: r,=-0.65; shading 1995: r= -0.54, 1996: r= -0.65, 1997: r= -0.70). The corresponding correla- tion coefficients were significant in 1997 (Table 5). Adult hibernators show declining numbers with decreas- ing land use intensity. This could be the result of less shading of more intensively used plots, which holds true not only for the sampling plots but for the entire area of investigation (Table 7). Therefore, the presence of adult hibernators does not depend on the type of management regime. An altitudinal moisture gradient does not seem to influence the distribution of spring breeders very much. Only for the year 1997 a decrease in the percent- age of spring breeders can be detected with increasing altitude (Table 5). A significant increase in the percent- age of all year reproductive species in 1995 and 1997 can be explained by the higher presence of the two woodland species A. parallelus and A. parallelepipedus. There does not seem to exist an overall tendency for the distribution of these reproductive types to be “driven” by land use intensity (FADL et al. 1998). The reproduc- tive forms of both species are especially successful and present on areas without high interference (mulched meadows and areas under succession). Table 6. Species percentages of reproductive types of carabids on sampled plots 3CM1, 2CMI, SPA, HP, SPR, MMI, MME2, AMR, and AMO during the years 1995 to 1997and 1997 only (Sb = spring breeders, Ab = autumn breeders, y = all-year reproductive) Sb Ab y 3CMI 1995 87.0 13.0 0 1996 16:3 17.6 5.9 1997, 852, eee 2CM1 1995 83.3 16.7 0 1996 70.0 25.0 5.0 1997. 15.0. 250 0 SPA 1995 66.7 33.3 0 1996 75.0 25.0 0 1997700 300 0 80 HP 1995 71.3 22:7 0 1996 75.0 25.0 0 SPR 1995 83.3 16.7 0 1996 77.8 222 0 1997 6883130 MMI 1995 84.2 15.8 0 1996 62.5 31.9 6.7 1997 68.8 31.3 0 MME2 1995 1331 Die 5.3 1996 66.7 26.7 5.9 | 1997 68.8 25.0.2 2063 AMR 1995 82.4 11.8 5.9 1996 82.4 11.8 14.3 1997 75018863 AMO 1995 75.0 16.7 8.3 1996 71.4 14.3 14.3 1997 63.6 27.3° Wok 3CM2 1997 81.0 190 00 3CM3 _ 1997 800 167 1, 138 2CM3- 1997 636 38 Da 2CM 197 77 203 MO SPC 197 Al 222 mE MMEI 1997 61.5 385 500 MME3 1997 37.210 SS WDL 1997 61.1 27.8 an Jiirgen DEUSCHLE & Erich GLUCK: Reproductive Types and Mobility of Carabids 319 Table 7. Rank correlation coefficients of percentage of species regarding different dispersion strategies of carabid assemblage with area parameter (1995/96: n = 9; 1997: n= 17; level of significance: * =p < 0.05, ** =p < 0.01, *** p < 0.001) Portion of species Land use Area Perimeter Altitude Shading 1995 brachypterous U) Ei -0,17 -0,40 DIT 0,12 dimorphous -0,63 0,43 0,52 -0,43 -0,41 _ Macropterous ______-0,76* _____-0,26. 003 -0,02 0,25! 1996 brachypterous 0,82+ -0,20 -0,63 0,36 0,13 dimorphous -0,69* 0,25 0,54 -0,20 -0,21 1997 brachypterous Oates -0,28 -0,35 0,30 0,48 dimorphous -0,80*** 0,29 0,13 -0,55 -0,57 macropterous 0,17 -0,17 0,29 0,41 0,25 The removal of grass alters the ecological conditions on the surface and in the upper layers of the ground resulting in pronounced changes of temperature and moisture. On mulched meadows the cut grass remaining on the ground offers hiding places. It also serves as a buffer to weather changes during the following days and weeks. Therefore, the event of cutting grass has a much stronger effect on typical meadows than on mulched meadows. Flightless (brachypterous) species colonise mulched meadows and areas under succession more often than typical meadows or pastures. On the other hand more flying (macropterous and alae dimorphous) species can be found on typical meadows and pastures (Tables 6, 7). Mulched meadows do not seem to be barriers to brachypterous woodland species colonizing scattered areas under succession. During the three years of inves- tigation the percentage of wingless woodland species was higher on mulched meadows and abandoned land than on typical meadows and pastures. Mulched meadows offer better resources to woodland species in spite of equal frequencies of mowing (BUTTERFIELD et al. 1995). Large forms of brachypterous woodland carabids, which are dependent on shade vegetation, have the tendency to look for shaded areas, if the contours of these areas stand out against their surroundings. it seems that a population can comprise parts of actively exploring in- dividuals (direct movement vs random walk, see CHAR- RIER et al. 1997), which are capable of overcoming not optimal areas (BURKE & GOULET 1998; GARDNER et al. 1997; GLUCK & KREISEL 1986). If they reach suitable habitats — in this case plots like AMO or MME2 - they can found new populations. Woodland species are espe- cially dominant or sub-dominant on the abandoned plot AMO. Newly hatched individuals of Abax parallelepi- pedus and Molops elatus gave proof of successful re- production of these species in area AMO. Directed migration seems to be difficult under a closed canopy. The unshaded abandoned plot AMR, however, has a small percentage of woodland species. This is not typical for areas under succession and may be partly due to the influence of the neighbouring two-cutting meadow. Larger, eurotopic woodland species like 4. parallelus and A. parallelepipedus were only found as single specimens during the investigation period. A growing number of woodland species could not be de- tected with increasing age of the abandoned area. On the heavily shaded mulched meadows MME2 and MME3 both species were more common (Fig. 3). Area AMR is lacking trees and dark contours. Both woodland species seem to frequent this area less. Nearly all specimens of each of the nine alae dimor- phous species were either completely macropterous (Pterostichus vernalis, Synuchus vivalis and Stomis pumicatus) or brachypterous (Carabus granulatus, Ca- lathus fuscipes, Pterostichus melanarius, Clivina fossor, Pterostichus anthracinus, Notiophilus palustris, Dyscir- ius globosus, Bembidion lampros). They, therefore, showed a constant expression of the development of their alae. There were four wingless species found typi- cal for three-cutting meadows (C. granulatus, C. fusci- pes, Pt. melanarius, C. fossor). These ubiquitous forms were able to take advantage of the intensive use of grassland (TIETZE 1985). In a pessimal environment the allele frequency of the corresponding recessive gene can increase within the population and minimize the risk of extinction (AU- KEMA 1987, DEN BOER 1987). Dimorphous species also show numerous evidence of flight. The flight activity of both forms seems to be lower under high colonisation density (optimal conditions) than under low abundance (pessimal conditions, DESENDER 1986, MEIJER 1974). Numerous marked individuals of the small, eurytopic species Calathus fuscipes, which is alae dimorphous but mostly brachypterous, overcame larger distances on the ground than individuals of the larger, stenoecic and macropterous species Harpalus dimidiatus. 320 Bonner zoologische Beitráge 53 (2004) Table 8. Distribution and percentage of macropterous individuals of the alae dimorphous species Bembidion properans on sam- pled plots ro Sex distribution (%) Macropterous (%) males females males females 3CMI (1995-1997, n = 9) 44 56 22 22 SPA (1995-1997, n= 1) 100 0 100 0 HP (1995-1997, n= 15) 13 87 0 53 SPR (1995-1997, n= 1) 100 0 100 0 3CM1 (1997, n= 19) 26 74 0 26 3CM2 (1997, n= 16) 38 62 0 6 SPC (1997, n= 1) | 100 0 100 OA Y (n = 62) 32 68 25 38 The only species which included macropterous and brachypterous individuals was B. properans (Table 8). But the activity densities of this species were very low. Sheep pastures were colonised discontinuously by this species. A total of 63 specimens were caught during the investigation period. Male beetles were less alate than females, though they were present on more sample plots. Females were disproportionally more alate and less present, albeit the fact that their colonisation den- sity was higher. Considering the reproductive behaviour of the species, females are more important. The ability to fly gives them the chance to leave an area when liv- ing conditions become pessimal. They can colonise other more profitable areas in a direct way and therefore minimize risk concerning their reproductive success. In the studied case this applies to three-cutting meadows and especially to the horse pasture. The high activity on this pasture corresponds with the findings of DESENDER & POLLET (1986), who describe an increase in the colo- nisation density of B. properans during intensified graz- ing. The radii of action of carabids stretch beyond the area defined by management practices. This leads to edge and spillover effects. Their dimensions depend on the individual mobility of the beetles, weather condi- tions, and availability of food. For reproduction both strategies need nearly one year, es- pecially the larval overwintering species can live for longer than one year and reproduce for more times (GRUM 1976). The high loading variables of factor 1 show the meaning of the importance of mobility and therefore the possibility leaving areas with pessimal liv- ing conditions rapidly independent from reproductive success. Resulting adaptations to extreme disturbances as they occur in orchard by pasturing or mowing, are the most important influences for the distribution of carabid beetles. Different strategies of reproduction are less im- portant. The fact that species can migrate into new habi- tats is well demonstrated (VAN DIJK 1987). A high mobil- ity reduces extinction risk of pessimal living conditions and further more buffers the inter- and intraspecific com- petition by the larvae as well as in adult carabids. Mobil- ity ensures and may maximize the reproductive success independent from the overwintering strategy. Mowing and pasturing are the ancient practises of culti- vating grasslands during the past few hundred years, on which the species corresponding could adapt them- selves. No negative influence could be measured so long on carabid assemblages, because immediatelly af- ter the impact higher or lower activities on the areas of investigation could not be measured (DESENDER et al. 1994) Obviously the impact is very grave and sustained for a long period. Only highly mobile species assem- blages can profit from these living conditions. But altogether only a combination of several factors might explain the distribution and composition of the assemblage. Acknowledgments. We thank very much Dipl. Agr.-Biol. Ernst Rupp, Jutta Deuschle and Bradley Sinclair (ZFMK) for revising the English text. REFERENCES AUKEMA, B. 1987. Differences in egg production and egg- laying period between long- and short-winged Ca- lathus erythroderus (Coleoptera, Carabidae) in rela- tion to wing morph frequencies in natural populations. Acta Phytopathologica Academiae Scientiarum Hun- garicae 22: 45-56. BUREL, F. & BAUDRY, J. 1994. Reaction of ground beetles to vegetation changes following grassland dereliction. Acta ecologica 15: 401-405. BURKE, D. & GOULET, H. 1998. Landscape and area effects on beetle assemblages in Ontario. Ecogeography 21: 472-479. BUTTERFIELD, J., LUFF, M., BAINES, M. & EYRE, M. D. 1995. Carabid beetle communities as indicators of conservation potential in upland forest. Forest Econ- omy & Management 79: 63-77. CHARRIER, S., PETIT, S. & BUREL, F. 1997. Movements of Abax parallelepipedus (Coleoptera, Carabidae) in woody habitats of a hedgerow network landscape — a radio-tracing study. Agriculture Ecosystems & Envi- ronment 61: 133-144. Jiirgen DEUSCHLE & Erich GLUCK: Reproductive Types and Mobility of Carabids 321 DEN Boer, P. J. 1987. On the turnover of carabid popula- tions in changing environments. Acta Phytopathologica Academiae Scientiarum Hungaricae 22: 71-83, DEN Boer, P. J. & VAN DUK, TH. S. 1994. Carabid beetles in changing environment. Wageningen Agriculture University Papers 94: 30 S. DESENDER, K. & POLLET, M. 1986. Adult and larval abun- dance from carabid beetles (Col, Carabidae) in a pas- ture under changing management. Mededelingen van de Faculteit voor Landbouwwetenschapen Rijksuni- versiteit de Gent 51: 943-955. DESENDER, K. 1986. On the relation between abundance and flight activity in carabid beetles from a heavily grazed pasture. Journal of Applied Entomology 102: 225-231. DESENDER, K., DUFRENE, M., LOREAU, M., LUFF, M. L. & MAELFAIT, J.-P. 1994. Carabid beetles: Ecology and Evolution. Kluwer Academic Publishers, Netherlands. 474 pp. DEUSCHLE, J. & GLUCK, E. 2001. Laufkáfer-Zónosen in Streuobstwiesen Sudwestdeutschlands und ihre Diffe- renzierung entsprechend unterschiedlicher Bewirt- schaftungsweisen (Coleoptera: Carabidae). Entomolo- gia Generalis 25: 275-304. DEUSCHLE, J., GLUCK, E. & BOCKER, R. 2002. Die Vegeta- tion von Streuobstwiesen. Veröffentlichungen zu Natur- und Landschaftspflege Baden-Wiirttemberg 74: 5-56. DE VRIES, H. H., DEN BOER, P. J. & VAN DUK, T. S. 1996. Ground beetle species in heathland fragments in rela- tion to survival, dispersal, and habitat preference. Oecologia 107: 332-342. DINGLE, H. 1996. Migration — The biology of life on the move. New York Oxford: Oxford University Press. GARDNER, S. M., HARTLEY, S. E., DAVIES, A. & PALMER, S. C. F. 1997. Carabid communities on heather moor- lands in Northeast Scotland — the consequences of grazing pressure for community diversity. Biological Conservation 81: 275-286. GLUCK, E. & KREISEL, A. 1986. Die Hecke als Lebens- raum. Refugium und Vernetzungsstruktur und ihre Bedeutung fiir die Dispersion von Waldcarabiden. Laufener Seminarbeitráge 10/86: 64-83. GLUCK, E. & DEUSCHLE, J. 2003. Habitat- und Feuchteprá- ferenz von Laufkáfern (Coleoptera, Carabidae) in Streuobstwiesen. Bonner zoologische Beitráge 51: 51-69. GRÚM, L. 1976. An attempt to characterize matter transfer by carabid communities inhabiting forests. Ekologia Polska 24: 365-375. HANSSEN, U. & HINGST, R. 1995. Einfluß systementlasten- der Nutzungsformen auf die biozónotische Struktur im Feuchtgriinland. Mitteilugnen der Deutschen Gesell- schaft fiir allgemeine und angewandte Entomologie 9: 475-480. MAELFAIT, J.-P., DESENDER, K. & DE KEER, R. 1988. The arthropod community of the edge of an intensivly grazed pasture. Miinstersche Geographische Arbeiten DOES. MEIJER, J. 1974. A comparative study of the immigration of Carabids (Coleoptera, Carabidae) into a new pol- der. Oecologia 16: 185-20. MULLER, J. K. 1987. Period of adult emergence in carabid beetles: an adaption for reducing competition? Acta Phytopathologica Academiae Scientiarum Hungaricae 22: 409-41. / OTTESEN, P. S. 1996. Niche segregatin of terrestrial Alpine beetles (Coleoptera) in relation to environmental gra- dients and phenology. Journal of Biogegraphy 23: 353-369. POLLARD, E. 1968. Hedges IV. A comparison between the Carabidae of a hedge and field site and those of a woodland glade. Journal of Applied Ecology 5: 649- 657. RODE, M. 1993. Habitatpräferenzen häufiger Carabiden- Arten des nordwestdeutschen Feuchtgrünlandes. Mit- teilugnen der Deutschen Gesellschaft für allgemeine und angewandte Entomologie 8: 417-425. RODE, M. & DÜLGE, R. 1994. Flügelausbildung bei Pre- rostichus strenuus Panzer und Prerostichus diligens Sturm (Coleoptera, Carabidae) in Abhängigkeit vom Habitat. Zoologische Jahrbücher für Systematik 121: 159-170. RUSHTON, S. P., LUFF, M. L. € Eyre, M. D. 1989. Habitat characteristics of grassland Prerostichus species (Col- eoptera, Carabidae). Ecological Entomology 16: 91- 104. SCHNITTER, P. H. 1994. The development of carabid com- munities from uncultivated fields and meadows in the first five years of a succession. Pp. 361-366 in: K. DE- SENDER et al. (eds.) Carabid beetles: Ecology and Evo- lution.Kluwer Academic Publishers, Netherlands. TIETZE, F. 1985. Veránderungen in der Arten- und Domi- nanzstruktur der Laufkáferzóosen (Coleoptera — Cara- bidae) bewirtschafteter Graslandókosysteme durch In- tensivierungsfaktoren. Zoologische Jahrbücher für Systematik 112: 367-382. VAN DUK, T. S. 1987. The long-term effects on the carabid fauna of nutrient impoverishment of previously arable field. Acta Phytopathologica Academiae Scientiarum Hungaricae 22: 103-118. Authors’ addresses: Dr. Jürgen DEUSCHLE (corre- sponding author): Káthe-Kollwitz-Str. 14, 73257 Kóngen, Germany; E-mail: deuschle@tloe-deuschle.de; Prof. Dr. Erich GLUCK: Gingener Weg 61/1, 73072 Donzdorf, Germany, E-mail: glueck.donzorf(@t-online.de. Received: 06.04.2004 Accepted: 15.09.2004 Revised: 04.04.2005 Corresponding editor: M. Schmitt hg er wo Bonner zoologische Beitráge Band 53 (2004) Heft 3/4 Seiten 323-332 Bonn, Dec. 2005 Rogambara and Cabamofa, Two New Genera of Enigmatic Sciaroids from Costa Rica F (Insecta: Diptera: Sciaroidea) Mathias JASCHHOF Swedish Museum of Natural History, Stockholm, Sweden Abstract. In Costa Rica, two new species of peculiar sciaroids were found that are assigned to two new genera, Rogam- bara dentata gen. et spec. nov. and Cabamofa mira gen. et spec. nov. The new taxa are described, illustrated and their systematic relationships are discussed. Rogambara and Cabamofa appear to be sister groups and the two together are the sister group of Ohakunea+ Colonomyia. All four genera form what is here called the Ohakunea group that is distinct from any of the existing families in the Sciaroidea and presumably closest to the Sciaridae. Zusammenfassung. In Costa Rica wurden zwei bemerkenswerte, zu den Sciaroidea gehörige neue Arten gefunden, für die zwei neue Gattungen begriindet werden: Rogambara dentata gen. et spec. nov. und Cabamofa mira gen. et spec. nov. Die neuen Taxa werden beschrieben, illustriert und ihre verwandtschaftliche Stellung wird diskutiert. Rogambara ist Schwestergruppe von Cabamofa; Rogambara+Cabamofa ist Schwestergruppe von Ohakunea+Colonomyia. Diese vier Gattungen bilden die so genannte Ohakunea-Gruppe, die keiner der existierenden Familien der Sciaroidea ange- schlossen werden kann und vermutlich den Sciaridae am nachsten steht. Key words. Phylogeny, taxonomy, new species Stichwórter. Diptera, Sciaroidea, Phylogenie, Taxonomie, neue Gattungen, neue Arten, Costa Rica 1. INTRODUCTION Fungus gnats in the broadest sense (Diptera: Sciaroidea) are poorly studied in Central America including Costa Rica, a fact applying absolutely to the species level and restrictively to all supraspecific levels. The preparation of the respective chapters for the Manual of Central American Diptera clearly meant a boost for systematic research on the Sciaroidea of this region, and one may hope that, once the Manual is published, this period of increased research activity and output will continue. In the course of studies by the author on various groups of Costa Rican Sciaroidea, mainly Cecidomyiidae, My- cetophilidae and Sciaridae, two species of enigmatic sciaroids were found which could not be assigned to any of the existing family group taxa. Such ‘unplaceable’ sciaroids are usually rare, both in nature and collections. Moreover, they are much sought-after objects of phy- logenetic studies aiming to further illuminate interfamil- ial relationships. Detailed study of the two Costa Rican enigmatics showed that they belong to two different new genera, Rogambara and Cabamofa, that are closely related to one another and to two other ‘unplaceable’ genera, Ohakunea Tonnoir and Edwards, 1927 and Colonomyia Colless, 1963, with the last also present in Costa Rica (HIPPA & JASCHHOF 2004). These four genera together form what is here called the Ohakunea group. In the fol- lowing, the new taxa are described, illustrated and their phylogenetic relationships are discussed. 2. MATERIAL AND METHODS Specimens of the new species were picked from un- sorted Malaise trap samples in the care of the Instituto Nacional de Biodiversidad (INBio), Santo Domingo, Costa Rica. It is noteworthy that only eight specimens in total were found among some 110 Malaise samples from various regions within this small but highly di- verse country. Specimens were made transparent by treatment with KOH, dehydrated in ethanol and eventu- ally, after treatment with beechwood creosote, mounted on microscopic slides in Canada balsam. All material, including types, are deposited in the INBio collection. Usage of morphological terminology follows that of SOLI (1997) for Mycetophilidae. Drawings were made using an Olympus BX50 microscope in combination with the U-DA drawing unit. 3. TAXONOMY 3.1. Genus Rogambara gen. nov. Type species. Rogambara dentata spec. nov., described below. Description Habitus: Slender, humpbacked sciaroids some 1.5 mm in size; moderately long antennae, legs and wings; rather narrow waist; slightly downcurved abdomen; and distinctive wing venation. Ethanol-preserved specimens coloration brownish. 324 Bonner zoologische Beitráge 53 (2004) Head: Head capsule higher than long; vestiture consist- ing of setae of various lengths, but short in all. Postfrons slightly bilobed, setose; frontal tubercle slightly two- pointed. Face larger than clypeus, non-setose. Clypeus setose, separated from face along its upper margin. An- tenna shorter than body, little longer in females than in males, inserted slightly above midheight of head. Scape somewhat conical, subequal to subglobular pedicel, both setose. Antennal flagellum with 14 flagellomeres sub- equal in length, terminal flagellomere as long as or slightly longer than penultimate; each flagellomere cy- lindrical with very short conical neck and node less than twice as long as wide, neck and node weakly demar- cated from another. Flagellum without microtrichia, ex- cept some basally on flagellomere 1. Each flagellomere with even cover of short setae arising from membranous rings and occurring in two sizes: fine and short setae, and stronger and longer ones arising from large basal pores, with latter more numerous in males than in fe- males; setae interspersed with sensory spines; most proximal flagellomeres without setae arising from sock- ets (present in related genera); terminal flagellomere with 3-4 apical setae arising from sockets. Ocelli absent. Eyes reniform, with complete bridge at vertex, without interommatidal setulae. Mouthparts well developed, with short proboscis. Labrum sclerotized, triangular, non-setose. Maxilla with lacinia well developed, latter style-like and fringed terminally. Maxillary palpus with 3 segments, lacking “presegment”; first segment thick- ened, with densely set, long hyaline sensory hairs on in- ner side; second and third segments elongate; all seg- ments setose with some setae spine-like. Labium with prementum (including premental apodemes) poorly de- veloped, with strong setae. Labial palpus 2-segmented; labellum 1 smooth, with some setae; labellum 2 small, with some spine-like setae. Thorax: Cervical sclerite well sclerotized. Anteprono- tum well developed, setose with 1 seta very long and strong; antepronotal sclerites interconnected by post- pronotum forming very narrow ridge above neck. Epi- sternum | clearly separated from pronotum, non-setose. Epimeron 1 small, subtriangular, situated at postero- ventral margin of episternum 1. Scutum with dorsal sur- face slightly evenly arched; with anterior parapsidal su- ture distinct and median transverse suture weak; with lateral, dorsocentral and acrostichal setae of various lengths, with some setae very long and strong. Parater- gite distinct from scutum and well sclerotized. Scutel- lum setose, with 2 central setae very long and strong. Mediotergite in lateral profile high, slightly arched. Laterotergite large, pronounced, its ventro-anterior mar- gin in touch with posterior margin of preepisternum 2, non-setose. Postphragma well developed, produced into abdominal cavity for about half length of first abdomi- nal segment. Anepisternum 2 setose, large and elongate, subequal in length to preepisternum 2, both sclerites separated by distinct (anapleural) suture. Preepisternum 2 subtriangular. Pleural suture distinct. Pleural pit dis- tinct. Epimeron 2 poorly sclerotized and demarcated, sometimes with anepimeral portion more distinct. Inter- nal mid-coxal fork distinct. Metanotum poorly defined, forming narrow collar above mediotergite, with 1 long and strong seta on either side. Episternum and epimeron 3 both poorly sclerotized and demarcated. Openings of thoracic spiracles without striking features. Legs: Long, i.e. about 1.2 times length of body. Coxae moderately short, 1.e. 0.6 (fore coxa) to 0.5 (hind coxa) times height of thorax. Femora slightly shorter than tibiae; tibiae shorter than tarsus. Tarsomeres 1-4 gradually decreasing in length, tarsomeres 1 more than twice as long as tar- someres 2, tarsomeres 4 and 5 subequal in length. Coxae flattened, with long setae largely confined to an- terior portions (fore coxa) or antero-distal margins (mid and hind coxae). Femora barely flattened, with longest setae (1.e., those along hind margins) clearly shorter than width of femur, with large non- and sparsely setose portions. Tibiae with tight cover of comparatively short setae interspersed with large trichia, the latter half to two-thirds as long as setae. Tibial spurs 1:1:1, compara- tively long. Fore tibia with anteroapical depression de- lineated by indistinct subtriangular rim and bearing nu- merous setae in irregular (proximally) and comb-like (distally) arrangement. Hind tibia with subapical comb of stiff setae similar to those on fore-tibial depression. Vestiture on tarsi largely as on tibiae, additionally with 2 longitudinal rows of short, spine-like setae and 1 dense row of short, scale-like setae underneath. Pretar- sal claws very small, slightly curved, without teeth. Empodia and pulvilli each consisting of some hairs shorter than claws. Wing: Clearly shorter than body, about 2 times as long as wide, widest near midpoint. No distinct alular or calypterous areas, anal area moderately developed. Membrane transparent, with light-brownish tinge, densely and evenly covered with microtrichia, without setae. Venation: C extending to apex of wing, near apex of M;; Sc broad but very short, ending free; h broad but poorly demarcated; Rs pale, visible mainly through its tracheae, situated unusually close to wing base, with oblique inclination; R (1.e., vein portion be- tween arculus and Rs) very short; Ry (1.e., vein portion between Rs and junction point with anterior wing mar- gin) excessively long, joining C near midpoint of wing; Ry absent; Rs (i.e., vein portion between diverging point of M,., and junction point with C which is clearly be- fore wing apex) excessively long, almost straight; frm (i.e., vein portion between Rs and diverging point of M +2) present but extremely short; tb present but short and faint and visible mainly by its tracheae, with oblique inclination; mcu nearly absent; Mj,. furcate with stem and proximal portions of fork pale, with point where M;¡,> diverges from Rs very far basally and situ- Mathias JASHHOF: Enigmatic Sciaroidea from Costa Rica 325 ated beyond Rs, fork clearly shorter than stem; origin of CuA, and CuA; closely approximated, 1.e. forming fork without common stem; CuP broad, running close to and reaching half length of CuA>; A, apparently running very close to CuP for some distance and thus indistin- guishable; A, practically absent. Setae present along wing margin and ventrally and dorsally on R, R; and Rs. With number of sensory pores somewhat variable, i.e. on Ry, 1-2 distally, and Rs, 1-2 distally and 2-3 proxi- mally, apart from numerous pores on Sc and stem vein. Halter club-shaped, with very few setae on both stem and knob. Abdomen: Segments | and 8 shorter than other seg- ments. Long, strong setae on all sclerites, except ster- nites 1 and 2. Number of spiracles uncertain. Tergal plaques small and inconspicuous, situated anterolater- ally on each sclerite, with their pattern (as far as visible) 0/1-2/1/1/1/1/1/0. Terminalia: Male. Sternite 9 absent as distinct sclerite. Gonocoxites without lobes; gono- coxal apodemes broad. Gonostyli simple (1.e., not lobed), with apical tooth-like structure. Ejaculatory apo- deme long, broad and flat, for most part poorly scle- rotized. Parameres merged, i.e. forming a tegmen; dor- sal parameral apodemes very short and interconnected by sclerotized transverse bridge. Tergite 9 large, plate- like. Tergite 10 absent. Cerci with numerous setae of various lengths distally and dorsally. Hypoproct consist- ing of one weakly sclerotized lobe, with 2 strong setae. Female. Tergite 8 large, sclerotized, setose. Gono- coxites 8 extended to proximal cercus, truncate termi- nally, with less than 10 strong setae distally and finer, shorter setae elsewhere; gonapophyses 8 distinct, weakly sclerotized; tergite 9 short, non-setose; go- napophysis 9 weakly sclerotized (1.e., not visible); ter- gite 10 well-developed but short, setose with some setae longer than any other setae on abdomen, with short pos- terior extension in which proximal cercus is embedded; sternite 10 present as weakly sclerotized ribs possibly merged posteriorly, with 1 strong lateral seta on either side; cercus one-segmented with distal segment absent; proximal cercus segment bearing numerous strong, straight setae, longer than wide, rounded posteriorly. Two sclerotized, disc-shaped spermathecae . Immature stages. Unknown. Diagnosis and discussion. The genus Rogambara is characterized by the following apomorphous characters: (1) the absence of ocelli; (2) the complete eye bridge; (3) the number of palpus segments reduced to three; (4) enlarged anepisterna 2; (5) the set of tibial spurs re- duced to 1:1:1; (6) the basalization of wing veins; (7) simplified male terminalia; and (8) female terminalia with the distal cercus segments absent. Character 6 ac- tually stands for a number of derived features following from the basalization of Rs (JASCHHOF & HIPPA 2003). Character 7 is actually a complex of characters, pf which the most important are: the absence of a separate sternite 9; the absence of gonocoxal lobes; and the pres- ence of a tegmen resulting from the fusion of the pa- rameres. Characters 4 and 5 are truly autapomorphous characters of the genus Rogambara and found nowhere else in the sciaroids considered its closest relatives (see Phylogeny Section below). Etymology. The genus name is composed of the family names of three of the parataxonomists working with INBio, Elias Rojas, Billen Gamboa and Wilfredo Arana, in appreciation of their enthusiastic and productive work in the field. The gender is feminine. Rogambara dentata spec. nov. (Figs. 1-15) Types. Holotype: Male, Costa Rica, Prov. Limon, Re- serva Biológica Hitoy Cerere, Sendero Espavel, 550 m, in tall secondary rain-forest, 18 Sept.-7 Oct. 2003, by Malaise trap, E. Rojas, B. Gamboa, W. Arana, M. & C. Jaschhof. Paratypes: 1 male, 2 females, same data as holotype; 1 female, same locality as holotype, but 11 March-1 April 2003; 1 male, Prov. Puntarenas, Parque Nacional Corcovado, 800 m Sendero a Cerro Rincón, 745 m, 20 Dec. 2001-7 Feb. 2002, by Malaise trap, J. Azofeifa. Other material studied. Costa Rica: 1 male, Limón, Siquerres, Pacuarito, Los Brisas, Reserva Ecologica Rio Dantas, near Estacion El Palenque, 500 m, 22-26 Au- gust 1996, by Malaise trap, B. Gustafsson, T. Pape and B. Viklund (in Swedish Museum of Natural History, Stockholm). Panama: | male, Caribbean coast, San Lorenzo Protected Area near Colon (9°17°N, 79258 W), wet evergreen forest, 130 m, 18 October 2003, by ground flight interception trap, A. Tishechkin (in Zool- ogy Department, Canterbury University, Christchurch). Description (for characters not mentioned here, see ge- nus description above) Body length: Males — 1.4 mm, females — 1.5-1.6 mm. Male. Head: Postfrontal lobes each with 1 seta. An- tenna (Fig. 2) with fourth flagellomere twice as long as wide, vestiture clearly shorter than width of flagel- lomere. Eye bridge at vertex 3 facets long. Labrum comparatively wide. 326 Bonner zoologische Beitráge 53 (2004) AE, Figs. 1-4: Rogambara dentata spec. nov.; — 1: Head of female, frontal view (0.1 mm); — 2: Antenna of male, lateral view (0.1 mm); — 3: Antenna of female, lateral view (0.1 mm); — 4: Flagellomeres 3-5 of female, lateral view (0.05 mm). 1, 3 and 4: Para- types from Hitoy Cerere; 2: Paratype from Corcovado (In parentheses: Length of scale bar.). Mathias JASHHOF: Enigmatic Sciaroidea from Costa Rica 327 Figs. 5-7: Rogambara dentata spec. nov.; — 5: Head and thorax of female, lateral view (0.2 mm); — 6: Apical portion of fore tibia | of male (0.05 mm); — 7: Apical portion of hind tibia of male (0.05 mm). 5: Paratype from Hitoy Cerere; 6, 7: Paratype from Cor- | covado (In parentheses: Length of scale bar.). 328 Thorax: Anepisternum 2 with maximum 6 setae in up- per half. Legs: Fore tibia with anteroapical depression bearing comb of some 15 stiff setae (Fig. 6). Hind tibia with subapical comb of some 15 stiff setae (Fig. 7). Terminalia: Gonocoxites (Fig. 10) ventrally with wide, V-shaped emargination, below emargination largely membranous and non-setose, with setae of various lengths elsewhere; gonocoxal apodemes broad and strong. Gonostyli (Fig. 10) tapering to tip; with heavy, curved tooth terminally and setae of various lengths elsewhere. Ejaculatory apodeme (Fig. 11) present as broad, flattened, weakly sclerotized rod almost as long as tegmen. Tegmen (Fig. 11) somewhat longer than wide, rounded distally; ventral gonocoxal apodemes swept ventrally; dorsal gonocoxal apodemes very short, interconnected by strong, short, sclerotized transverse bridge. Tergite 9 (Fig. 12) plate-like with long anter- olateral bars and broadly rounded distal margin, with numerous setae of various lengths 2 of which are very long and strong. Cerci (Fig. 10) setose, large, broadly rounded terminally. Hypoproct (Fig. 10) consisting of smooth, rounded lobe bearing 2 strong setae. Female. Head: See Figures 1, 5. Antenna as in Figure 3, with fourth flagellomere as in Figure 4. Thorax: See Figure 5. Wing: See Figures 8, 9. Terminalia: See Figures 13-15. Etymology. The name is Latin meaning ‘toothed’, re- ferring to the gonostylus tooth in males of this species. TÁ 272 — FE ie eg = > IR EEE TEE I a TOR ELS a Figs. 8-9: Rogambara dentata spec. nov., paratype female; — 8: Wing, setae omitted (0.25 mm). — 9: Wing base (0.1 mm) (In parentheses: Length of scale bar.). Bonner zoologische Beiträge 53 (2004) 3.2. Genus Cabamofa gen. nov. Type species. Cabamofa mira spec. nov., described below. Description (based on females) Habitus: Slender, humpbacked sciaroids some 3 mm in size (males might be considerably smaller); with mod- erately long antennae, legs and wings; rather narrow waist; and distinctive wing venation. Ethanol-preserved specimens coloration light-brownish. Figs. 10-12: Rogambara dentata spec. nov., holotype male; — 10: Terminalia, ventral view; — 11: Tegmen and ejaculatory apodeme, ventral view; — 12: Tergite 9, dorsal view (Length of scale bar = 0.05 mm.). Head: Head capsule subglobular; vestiture consisting of setae of various lengths, but short in all. Postfrons bilobed, non-setose; frontal tubercle slightly two- pointed. Clypeus setose. Antenna shorter than body, in- serted slightly above midheight of head. Scape some- what conical, subequal to subglobular pedicel, both se- tose. Antennal flagellum with 14 flagellomeres subequal in length, first and terminal flagellomeres clearly longer; each flagellomere cylindrical with short conical neck and node barely longer than wide, neck and node clearly Mathias JASHHOF: Enigmatic Sciaroidea from Costa Rica 329 demarcated from another. Flagellum without microtri- chia, except some basally on flagellomere 1. Each flag- ellomere with irregular cover of long setae arising from sockets intermixed with short setae arising from mem- branous rings; setae interspersed with sensory spines. Ocelli absent. Eyes reniform, with complete bridge at vertex and long interommatidal setulae. Mouthparts well developed, with short proboscis. Labrum small, sclerotized, non-setose. Maxilla with lacinia well devel- oped, the latter style-like and fringed marginally and terminally. Maxillary palpus with 5 segments including distinct "presegment" bearing at least 1 strong seta; third segment swollen, with sensory pit on inner side; fourth and fifth segments elongate; all segments setose with some setae spine-like. Labium with premental lobe bearing strong setae. Labial palpus 2-segmented; label- lum | very small and non-setose; labellum 2 large, with spine-like setae in rows. IE Figs. 13-15: Rogambara dentata spec. nov., paratype females; — 13: Terminalia, ventrolateral view; — 14: Terminalia, dorso- lateral view; — 15: Spermatheca (Length of scale bar = 0.1 mm.). Thorax: Cervical sclerite well sclerotized. Anteprono- tum well developed, setose; antepronotal sclerites inter- connected by postpronotum forming very narrow ridge above neck and bearing setae laterally. Episternum | clearly separated from pronotum, setose. Epimeron | small, subtriangular, situated at postero-ventral margin of episternum 1. Scutum with dorsal surface evenly and slightly arched; anterior parapsidal suture distinct and median transverse suture weak; lateral, dorsocentral and acrostichal setae of various lengths. Paratergite poorly developed. Scutellum setose, with 2 central setae very long and strong. Mediotergite in lateral profile compara- tively short, slightly arched. Laterotergite large, with rather smooth transition into mediotergite, its ventro- anterior margin approximated with posterior margin of preepisternum 2, non-setose. Postphragma well devel- oped, produced into abdominal cavity for short distance. Anepisternum 2 short, non-setose, both sclerites sepa- rated by distinct (anapleural) suture. Preepisternum 2 large, subtriangular. Pleural suture distinct. Pleural pit distinct. Epimeron 2 poorly sclerotized and demarcated. Internal mid-coxal fork distinct. Metanotum poorly de- fined, forming narrow collar above mediotergite, with 1-2 setae on either side. Episternum 3 large, weakly sclerotized, more clearly demarcated posteriorly with sometimes two parts recognizable. Openings of thoracic spiracles without striking features. Legs: Fore leg little shorter and hind leg little longer than body. Coxae mod- erately short, 1.e. 0.6 (fore coxa) to 0.4 (hind coxa) times height of thorax. In fore leg, femur and tibia subequal in length, tibia shorter than tarsus. In mid leg, femur shorter than tibia, tibia shorter than tarsus. In hind leg, femur shorter than tibia, tibia and tarsus subequal in length. Tarsomeres 1-4 gradually decreasing in length, tarsomere 1 more than twice as long as tarsomeres 2, tarsomeres 4 and 5 subequal in length. Coxae flattened, with long setae largely confined to anterior portions. Femora barely flattened, with longest setae (1.e., those along hind margins) clearly shorter than width of femur. Tibiae with dense cover of comparatively short setae in- terspersed with large trichia, latter half to two-thirds as long as setae. Tibial spurs 1:2:2, comparatively short. In mid and hind tibiae, one spur clearly shorter than other. Fore tibia with anteroapical depression subtriangular, very weakly delineated, bearing numerous setae in ir- regular (proximally) and comb-like (distally) arrange- ment. Hind tibia with subapical comb of stiff setae simi- lar to those on fore-tibial depression. Vestiture on tarsi largely as on tibiae. Pretarsal claws crescent-shaped, without teeth. Empodia well developed, as long as claws. Pulvilli delicate, two-thirds as long as claws. Wing: Shorter than body, more than 2 times as long as wide, widest slightly beyond midpoint. No distinct alu- lar or calypterous areas, anal area moderately devel- oped. Membrane transparent, with light-brownish tinge, densely and evenly covered with microtrichia, without setae. Venation: C extending to apex of wing, between apices of Rs and Mı; Sc broad but very short, ending free: h broad but poorly demarcated; Rs pale, situated unusually close to wing base, with oblique inclination; R (1.e., vein portion between arculus and Rs) very short; 330 Bonner zoologische Beitráge 53 (2004) R, (1.e., vein portion between Rs and junction point with anterior wing margin) excessively long, joining C be- yond midpoint of wing; Ry absent; Rs (i.e., vein portion between diverging point of M;.5 and junction point with C which is practically at wing apex) excessively long, curved in distal third; frm (1.e., vein portion between Rs and diverging point of M,.,) present, longer than Rs; tb present, as long as one vein width; mcu present, pale, longer than Rs; M,.> furcate with its stem almost en- tirely absent, point where M;,. diverges from Rs very far basally and situated clearly beyond Rs, fork clearly shorter than stem; CuA, and CuA, arising separately; CuP distinct, running close to and reaching beyond half length of CuA;; A, extending to two-thirds length of CuP; A, nearly absent. Setae present along wing mar- gin, ventrally on R, R,, Rs, M-fork, CuA, and CuAg, and dorsally on distal portion of Rs. Number of sensory pores somewhat variable, i.e. on Rj, 3-4; frm, 2-3; and Rs, 2 distally, apart from numerous pores on Sc and stem vein. Halter club-shaped, with setae on knob. Abdomen: With segments 1 and 8 shorter than other segments. With setae on all sclerites except sternite 1. Number of spiracles uncertain. Tergal plaques appar- ently absent. Terminalia: Tergite 8 large, sclerotized, setose. Gonocoxites 8 extending up to proximal cercus, densely setose; gonapophyses 8 indistinct, weakly scle- rotized; tergite 9 not recognizable as distinct sclerite, possibly merged with tergite 10; gonapophysis 9 present as internal sclerotized fork; tergite 10 setose, with poste- rior extension in which proximal cercus 1s embedded; sternite 10 not clearly recognizable; cercus two-segmented, setose; proximal segment subtriangular in lateral view, longer than distal segment; distal segment rounded. Two sclerotized, disc-shaped spermathecae . Male and immature stages. Unknown. Diagnosis and discussion. In the genus Cabamofa, apomorphous characters include: the absence of ocelli; the complete eye bridge; and the basalization of wing veins. There is no autapomorphous character recogniz- able for Cabamofa; instead, all its derived features are found also in Rogambara. However, quite a number of derived features in Rogambara dentata, including two autapomorphies recognized, lack in Cabamofa mira, a fact making it problematic to argue that these two spe- cies are congeneric. In Cabamofa, the vestiture of the antennal flagellum includes an even cover with setae arising from sockets, a feature unique within the Oha- kunea group (see Phylogeny Section below), but at pre- sent it is hard to tell whether this is of more than spe- cies-specific importance. Etymology. The genus name is composed of the family names of three of the parataxonomists working with INBio, Khanaki Caballero, Marco Moraga and Alejan- dro Azofeifa, in appreciation of their successful field work to that we owe these very peculiar flies. The gen- der is feminine. Cabamofa mira spec. nov. (Figs. 16-26) Types. Holotype: Female, Costa Rica, Prov. Puntare- nas, Parque Nacional Corcovado, Quebrada Ceniza, 300 m, 14 March-5 April 2003, by Malaise trap, K. Cabal- lero, M. Moraga & A. Azofeifa. Paratypes: | female, same data as holotype. Description (for characters not mentioned here, see ge- nus description above) Figs. 16-19: Cabamofa mira spec. nov., female; — 16: Eye bridge, dorsofrontal view (0.1 mm); — 17: Head, lateral view (0.1 mm); — 18: Maxillary palpus, lateral view (0.05 mm); — 19: Antennal flagellum, lateral view (0.1 mm). 16: Holotype; 17-19: Paratype (In parentheses: Length of scale bar.). Body length: Females — 2.9 and 3.1 mm. Male. Unknown. Female. Head: See Figure 17. Antenna (Fig. 19) with node of fourth flagellomere (Fig. 23) 1.1 times as long as wide; longest setae shorter than width of flagel- lomere. Eye bridge (Fig. 16) at vertex 5 facets long. Maxillary palpus as in Figure 18. Thorax: See Figure 20. Postpronotum with 3 setae lat- erally. Antepronotum with | seta. Episternum 1 with 4-5 Mathias JASHHOF: Enigmatic Sciaroidea from Costa Rica 331 setae. Legs: Fore tibia with anteroapical depression bearing comb of some 12 stiff setae (Fig. 21). Hind tibia with subapical comb of some 18 stiff setae (Fig. 22). Wing: See Figures 24, 25. Terminalia: See Figure 26. Figs. 20-22: Cabamofa mira spec. nov., holotype female; — 20: Thorax, lateral view (0.25 mm); — 21: Apical portion of fore tibia (0.05 mm); — 22: Apical portion of hind tibia (0.05 mm) (In parentheses: Length of scale bar.). Etymology. The name is Latin meaning “strange”, refer- ring to the peculiar assemblage of morphological char- acters in this species. 4. PHYLOGENY The peculiar wing vein pattern and well developed postphragma in Rogambara and Cabamofa give reason to roughly assign them to a group of sciaroid genera which cannot be classified within any existing family of the Sciaroidea. Knowledge of these unplaceable sciar- oids has increased significantly in recent years (see CHANDLER (2002) and JASCHHOF (2004) for Heterotri- cha Loew and allies; JASCHHOF & DIDHAM (2002) for Rangomaramidae; JASCHHOF & HIPPA (2003) for Oha- kunea Tonnoir & Edwards; HIPPA & JASCHHOF 2004, for Colonomyia Colless); however, the phylogenetic re- lationships between these taxa and to the long- recognized family-level taxa within the Sciaroidea re- main debatable. The basalization of Rs and other wing veins (Fig. 27, character 1) serves as an argument to consider Rogam- bara, Cabamofa, Ohakunea and Colonomyia belonging to the same monophyletic group, the Ohakunea group. Rogambara appears to be most closely related to Ca- Figs. 23-26: Cabamofa mira spec. nov., female; — 23: Anten- nal flagellomeres 3-5, lateral view (0.05 mm); — 24: Wing, setae omitted (0.5 mm); — 25: Wing base (0.5 mm); — 26: Terminalia with one spermatheca, lateral view (0.1 mm). 23, 24 and 26: Paratype; 25: Holotype (In parentheses: Length of scale bar.). bamofa. Postulation of their sister-group relationship is based on three arguments: the presence of a complete eye bridge, absence (loss) of ocelli and shortening of the antennal flagellum (Fig. 27, characters 2-4). As regards the last character, one must explain that shortening here means shortening of the individual flagellomeres, not shortening through reduction in the number of flagel- lomeres. The clade Ohakunea+Colonomyia is based on two synapomorphies: regaining of the Ry and loss of sclerotized spermathecae (Fig. 27, characters 5 and 6). As argued by JASCHHOF & HIPPA (2003), Ry was pre- sent in the ground plan of the Sciaroidea and later lost in the hypothetical common ancestor of a clade comprising Diadocidiidae, the Heterotricha group sensu CHANDLER (2002), Ohakunea, Colonomyia, Sciaridae, Rango- maramidae and Cecidomyiidae. Among the genera dis- cussed here, Rogambara possessed the most apomor- phous character states (Fig. 27, characters 7-10), which find their plesiomorphous counterparts in both Oha- kunea (JASCHHOF & HIPPA 2003; and Fig. 27, characters 11-12) and Colonomyia (HIPPA & JASCHHOF 2004; and Fig. 27, characters 13-16). Among the derived features found in Rogambara, the three-segmented maxillary palpus and simplified male terminalia are most notewor- thy as these correspond with the conditions found in the Sciaridae, a family considered most closely related to Ohakunea and Colonomyia. The structure of the maxil- lary palpus in Rogambara and Sciaridae is practically identical. As regards male terminalia, Rogambara and Sciaridae differ only in small details. The most important differences lie in the structure of the aedeagus/tegmen complex: in Rogambara, the ejaculatory apodeme is broad, flattened and as long as the tegmen and aedeagal teeth are absent; in Sciaridae, the ejaculatory apodeme is usually narrow, tube-like and shorter than the tegmen and aedeagal teeth are usually present. I explain the pres- ence of a three-segmented palpus and simple male ter- minalia in both Rogambara and Sciaridae by homoplasy. Ohakunea Cabamofa Rogambara Colonomyia Fig. 27: Postulated relationships between the genera of the Ohakunea group (after JASCHHOF & HIPPA (2003); HIppA & JASCHHOF 2004; and this study). Black blocks = apomorphous characters; ? = no apomorphy recognized. Numbers 1-16 refer to apomorphous character states as follows: 1, Rs and other wing veins basalized; 2, eye bridge complete; 3, ocelli lost; 4, antennal flagellum shortened; 5, Ry regained; 6, sclerotized spermathecae lost; 7, number of maxillary palpus segments reduced to three; 8, anepisternum 2 enlarged; 9, one spur of mid and hind tibia lost; 10, male terminalia simplified; 11, male terminalia with parameres subdivided into ventral and dorsal pairs; 12, male terminalia with gonostyli strongly nar- rowed; 13, epimeron 2 shortened; 14, basitarsus with longitu- dinal row of specialized setae; 15, pretarsal claws deeply bi- furcated; 16, male terminalia with gonocoxites bearing interior sclerotized ribs. The absence of any autapomorphous characters in Ca- bamofa may be explained by the fact that its male is yet to be found, because experience shows that sciaroid male terminalia, due to their structural complexity, often can provide features of significance for phylogenetic considera- tions. No attempt was made in order to place the Ohakunea group among other Sciaroidea, as from a phylogenetic viewpoint the here introduced taxa cannot provide hints additional to those considered earlier (CHANDLER 2002; JASCHHOF & HIPPA 2003; HIPPA & JASCHHOF 2004). Bonner zoologische Beitráge 53 (2004) Acknowledgements. Study of the insect collections at IN- Bio was made possible through the generosity of INBio providing various support both in the laboratory and field during a three-month visit from September to November, 2003. In particular, I am much indebted to Manuel Zum- bado for his attentiveness and interest, and to the paratax- onomists Elias Rojas, Billen Gamboa and Wilfredo Arana who shared with us their knowledge on Costa Rican insects and wildlife. We thank the staff of the field station of the Reserva Biológica Hitoy Cerere facilitating greatly my four-weeks visit. My wife, Catrin Jaschhof, Greifswald, Germany, greatly assisted in the field and in sorting through numerous insect samples. Dr Raphael Didham, Zoology Department, Canterbury University, Christchurch, New Zealand, and Prof Dr Heikki Hippa, Swedish Mu- seum of Natural History, Stockholm, Sweden informed me of material of Rogambara dentata in their collections. Language of the manuscript was kindly improved by Dr Bradley Sinclair, Zoologisches Forschungsmuseum Alex- ander Koenig, Bonn, Germany. The helpful comments on the manuscript by two anonymous reviewers are gratefully acknowledged. REFERENCES CHANDLER, P. 2002. Heterotricha Loew and allied genera (Diptera: Sciaroidea): offshoots of the stem group of Mycetophilidae and/or Sciaridae? Annales de la So- ciété Entomologique de France (n.s.) 38(1-2): 101-144. JASCHHOF, M. 2004. The Heterotricha Loew group (Dip- tera: Sciaroidea) in New Zealand. Beitráge zur Ento- mologie 54(1): 3-30. JASCHHOF, M. & DIDHAM, R. K. 2002. Rangomaramidae fam. n. from New Zealand and implications for the phylogeny of the Sciaroidea (Diptera). Studia dip- terologica Supplement 11, 60 pp. JASCHHOF, M. & HIPPA, H. 2003. Sciaroid but not sciarid: a review of Ohakunea Tonnoir & Edwards, with the de- scription of two new species (Insecta: Diptera: Bibio- nomorpha). Entomologische Abhandlungen, Dresden 60: 23-44. HIPPA, H. & JASCHHOF, M. 2004. A re-evaluation of the genus Colonomyia Colless (Diptera, Sciaroidea), with the description of two new species. Insects Systemat- ics and Evolution 35(3): 335-352. SOLI, G. 1997. The adult morphology of Mycetophilidae (s.str.), with a tentative phylogeny of the family (Dip- tera, Sciaroidea). Entomologica scandinavica Supple- ment 50: 5-55. Authors’ address: Dr. M. JASCHHOF: Swedish Museum of Natural History, Box 50007S-10405, Stockholm, Sweden; E-mail: mjaschhof@yahoo.de. Received: 02.03.2004 Revised: 10.05.2004 Accepted: 18.05.2004 Corresponding editor: B. J. Sinclair Bonner zoologische Beiträge Band 53 (2004) | Heft 3/4 | Seiten 333-357 Bonn, Dec. 2005 | The Net-Winged Midges (Diptera: Blephariceridae) of the Russian Far East / Peter ZWICK" 8: Tatyana AREFINA” | "Limnologische Fluss-Station Schlitz, Schlitz, Germany Institute of Biology and Soil Science, Russian Academy of Sciences, Vladivostok, Russia Abstract. A revision and synopsis of the Diptera: Blephariceridae of the Russian Far East is presented; it includes ge- neric keys to all life stages. Constitutive characters supporting the monophyly of the genera Bibiocephala Osten-Sacken and Agathon von Réder are named. Genera Philorus Kellogg and Blepharicera Macquart are recorded from the Russian Far East for the first time. New species described are Philorus levanidovae sp. n., Blepharicera parva sp. n. and Agathon setosus sp. n. The number of species known from the Russian Far East rises to eight. Agathon decorilarva (Brodsky, 1954), A. eoasiaticus (Brodsky, 1954) and Neohapalothrix manschukuensis (Mannheims, 1938) are wide- spread, ranging west to Mongolia and Kazakhstan, respectively, and 4. decorilarva east to Kamchatka. All species were compared with actual specimens of close relatives from Korea and Japan and are distinct from them, except perhaps Bibiocephala maxima Brodsky, 1954. In the case of Bibiocephala, the world fauna is compared and keys to all stages of the four nominal species are provided. Agathon kawamurai (Kitakami, 1950) from Korea, A. ezoensis (Kitakami, 1950) from Sakhalin and Hokkaido, and the Korean Philorus chosenensis Kitakami, 1931 are redescribed from lectotypes, which are here designated. Key words. Taxonomy, faunistics, new species, lectotype designation, keys, Far East Asia 1. INTRODUCTION The Blephariceridae of the Russian Far East were first studied by BRODSKY (1954) who named several taxa. Unfortunately, Brodsky was apparently unaware of sev- eral studies by S. Kitakami who published on the Ble- phariceridae of Japan, Taiwan, Korea, Sakhalin, and Manchuria, from 1931-1950, and of the description of a peculiar Manchurian species (MANNHEIMS 1938). Two of Brodsky's taxa proved in the meantime to be syno- nyms of species named from nearby countries in the above papers. The validity of the remaining nominal species needed confirmation. BRODSKY'S collection is in the Zoological Museum, St. Petersburg, but not all of his specimens could be lo- cated. Kitakami's collection in Kyoto (KITAKAMI 1951: 70) is apparently lost. Very few specimens remain in the Otsu Hydrobiological Station of Kyoto University at Otsu (K.N. Satake, in lit. 1987); we were unable to study them. However, Kitakami gave specimens in al- cohol, including syntypes, to the United States National Museum (Washington, D.C.) and to B. Mannheims (Museum Koenig, Bonn, Germany). This material has uniform printed labels, occasional additions in ink are all in the same style, probably in Kitakami's own hand. A typed page by Kitakami sent to Mannheims lists every individual specimen with place and date of collec- tion. The list shows that specimens belonging to some particular species and kept in the same vial may never- theless have been collected at different places or times, respectively. Similar information is not available for the material in USNM. Therefore, even though the Wash- ington specimens are authentic Kitakami material, they are eligible as lectotypes only under certain conditions, mainly taxa named in Kitakami's last taxonomical paper (1950) where all available specimens were listed as equal syntypes. NARTSHUK (1999) presented keys to the then known larvae and pupae of the entire Russian fauna, to species or genus, respectively, apparently largely relying on published information. Adults were not considered. 2. MATERIAL, COLLECTION SITES AND DEPOSITORIES During three weeks in June 1998, net-winged midges were the specific target of fieldwork by the authors and several of their colleagues in Primorye and the Kha- barovsk district; collecting and treatment of collected insects conformed with Russian legal regulations. Whenever possible, large samples were collected to permit safe association of different life stages by dissec- tion of pharate specimens, and to give an idea of the population structure at the time of sampling, and per- haps of volitinism and phenology. This alcohol- preserved material provides also information on habitat preferences and species associations. To avoid frequent repetition, locality information is gathered in Table 1. Additional material collected by workers of the Institute of Biology and Soil Science, Academy of Sciences, Vladivostok (IBSSAN), consists of many smaller sam- ples, most from the same general area. Some data were already published (AREFINA 1995). We list this addi- 334 Bonner zoologische Beiträge 53 (2004) Table 1. List of localities successfully sampled for Blephariceridae by the authors during field work in Primorye and the Kha- barovsk District, in June 1998; material housed in coll. P. Zwick. Coordinates were read from Tactical Pilot Charts (prepared and published by the Defense Mapping Agency Aerospace Center, St. Louis, Missouri, 1986) and are approximate RFEOI RFEO3 RFEO4 RFEO4 RFEO6 RFEO6 RFEO7 RFEOS RFEO9 RFE10 RFE10a RFEI2 RFEI3 RFE14 RFEIS RFE16 RFE17 RFEIS RFE19 RFE20 RFE21 RFE22 RFE23 RFE24 RFE2S RFE26 RFE27 43°24'E, 132°48'N; Primorje, Steklyanukha River, tributary of Shkotovka River, Sevy Gorbatov Klyuch, waterfalls and cascades accessible from logging road beyond Novaya Moskva, 11.06.1998 and 26.06.1998 43°03'N, 133°17'E; Primorje, Vodopadnaja River nr Nikolayevka, E of Partizansk [43.08N, 133.09E], 12.06.1998 43°05'N, 133°18'E; Primorje, small rocky tributary to RFEO3, at forest edge, 12.06.1998 43°05'N, 133°18'E; Primorje, small rocky tributary to RFEO3, at forest edge, 12.06.1998 43°25'N, 133°28'E; Primorje, Povorotny River NE Partizansk [43.08N, 133.09E], on road to Lazo, 12.06.1998 43°25'N, 133°28'E; Primorje, Povorotny River NE Partizansk [43.08N, 133.09E], on road to Lazo, 12.06.1998 43°29'N, 133°42'E; Primorje, Lazovka River 30 km W of Lazo[43.25N, 133.55E], 12.06.1998 43°28'N, 133°44'E; Primorje, Lazovka River 25 km W of Lazo [43.25N, 133.55E], at bridge, 13.06.1998 43°12'N, 133°15'E; Primorje, Kiyevka River, 5 km S of Lazo [43.25N, 133.55E], 13.06.1998 43°35'N, 134°36'E; Primorje, upper course of Milogradovka River, 25 km NW Milogradovo, 13.06.1998 Primorje, small forest tributary to RFE10 in forest, 14.06.1998 43°38'N, 134°20'E; Primorje, Ussuri River, upper course on logging track from Margaritovka, N of pass, near Nishni, 15.06.1998 43°42'N, 133°58'E; Primorje, Krasnaya River, tributary of Ussuri near Beryozovi, 15.06.1998 43°43'N, 133°53'E; Primorje, Pravaya Poperechka River S of Arkhipovka, 15.06.1998 43°53'N, 133°55'E; Primorje, Izvilinka River at Izvilinka, near mouth into Ussuri, 15.06.1998 44°19'N, 133°54'E; Primorje, Ussuri River 6 km SW Kamenka, 15.06.1998 44°41'N, 134°04'E; Primorje, Zadovka River, tributary of Zhuralyovka River 10 km SW Samarka, 16.06.1998 44°43'N, 134°0O8'E; Primorje, Zhuralyovka River (tributary of Ussuri) at Samarka, 16.06.1998 44°50'N, 134°19'E; Primorje, Komisarovski River near Lesogorje, tributary of Otkosnaya River/ Us- suri, 16.06.1998 44°57'N, 134°24'E; Primorje, Malinka River S of Pozhiga, 16.06.1998 45°05'N, 134°32'E; Primorje, Malinovka River ESE of Pozhiga, 16.06.1998 45°06'N, 134°43'E; Primorje, cold forest tributary to RFE21, 3 km upstream on logging track, 17.06.1998 45°26'N, 134°35'E; Primorje, Orekhovka River near Marevka, tributary Malinovka, 17.06.1998 45°27'N, 134°42'E; Primorje, small tributary to RFE23, about 5km on, near Polyany, 17.06.1998 45°28'N, 135°00'E; Primorje, small upper course of Orekhovka River, near pass to Perevalnaja River, 17.06.1998 45°29'N, 135°04'E; Primorje, small forest stream just N of pass, Rogaty River, tributary of upper Pere- valnaja River, 17.06.1998 45°32'N, 135°04'E; Primorje, Rudny River, larger tributary of upper Perevalnaja River, 17.06.1998 RFE28 RFE30 RFE31 RFE33 RFE34 RFE35 RFE36 RFE37 RFE38 RFE39 RFE41 RFE42 RFE43 RFE44 RFE45 RFE46 RFE47 RFE48 RFE49 RFE49a RFESO RFES2 RFE53 RFES4 RFE55 RFE56 RFE100 Peter ZWICK & Tatyana AREFINA: Blephariceridae of the Russian Far East 335 45°38'N, 135°10'E; Primorje, Perevalnaja River, near mouth of Dorozhny River, 17.06.1998 46°32'N, 134°53'E; Primorje, branch of Bikin River near ferry, 25 km E of Verkhny Pereval, 18.06.1998 Primorje, unnamed small stream on road, much disturbed through building of new culvert, turbid, 19.06.1998 / 46°41'N, 135°27'E; Primorje, Bikin River and major tributary near Soboliny, 19.06.1998 46°43'N, 135°28'E; Primorje, Stolbovaya River above Soboliny, tributary of Bikin River, 19.06.1998 46°45'N, 135°33'E; Primorje, Tahalo River, large tributary to Bikin River at bridge, 19.06.1998 46°48'N, 135°38'E; Primorje, Maly Tahalo River, 20.06.1998 47°12'N, 135°34'E; Khabarovsky Kraj, Ivanov River at large bridge, between Yushnyi and Metsa, 20.06.1998 47°22'N, 135°37'E; Khabarovsky Kraj, Dolmi River at Dolmi, S tributary of Khor River, 20.06.1998 47°33'N, 135°33'E; Khabarovsky Kraj, Matai River, 20.06.1998 48°04'N, 136°23'E; Khabarovsky Kraj, upper course of Nemtu River on road from Sidima to Sukpai, 21.06.1998 Khabarovsky Kraj, small tributary to RFE41, about 3 km towards Sukpai, 21.06.1998 4805'N, 136°33'E; Khabarovsky Kraj, Petin River, tributary of Nemtu/Khor River, 21.06.1998 48°02'N, 136°44'E; Khabarovsky Kraj, tributary to Ku River, 21.06.1998 48°02'N, 136°44'E; Khabarovsky Kraj, cascades of small cold spring-fed stream 100 m uphill from RFE 44, 21.06.1998 4801'N, 136°46'E; Khabarovsky Kraj, Ku River, upper course at first bridge on road Sidima-Sukpai, 21.06.1998 47°56'N, 136°46'E; Khabarovsky Kraj, Khor River below Sukpai village, 21.06.1998 47°59'N, 136°47'E; Khabarovsky Kraj, Sukpai River at Sukpai village, 22.06.1998 48°00'N, 136°53'E; Khabarovsky Kraj, large tributary to Sukpai River, 22.06.1998 48°00'N, 136°53'E; Khabarovsky Kraj, tributary to large tributary of Sukpai River, 22.06.1998 47°47'N, 136°55'E; Khabarovsky Kraj, Sukpai River upper course at dead end of road, 22.06.1998 Khabarovsky Kraj, Avan River on main road Khabarovsk — Vladivostok, just N of Kotikovo [47°27'N, 134°37'E], 23.06.1998 Primorje, Bolshaya Ussurka, immediately above Dalnerechensk [45.55N, 133.43E], 23.06.1998 44°45'N, 131°43'E; Primorje, Poperechnaya River, small tributary to Molokanka River W of Lake Khanka, 24.06.1998 44°45'N, 131°52'E; Primorje, tiny trickle, tributary to Komissarovka River, at old bridge, 24.06.1998 (Kommissarovo [44.59N, 131,.46E]) 44°45'N, 131°48'E; Primorje, Komissrovka River, at mouth of Reshetinka River, SE of the Pogranichny Ridge, 24.06.1998 43°21'N, 131°38'E; Gryaznaya stream, waterfalls, 10.07.1998, T. Arefina & T. Tiunova Lo oS) o tional material under the respective species only if taxo- nomically important, or if originating from places dis- tinctly outside the range sampled in 1998. We were also given valuable comparative material from Mongolia (P. Surenkhorloo, Ulan Bataar), Kazakhstan (V. Devyatkov, Ust-Kamenogorsk) and Japan (S. Uchida, Toyota). All of these collections are presently kept at Schlitz in collection P. Zwick (PZ) and will eventually be deposited in a public museum. Comparative material was borrowed from the following collections: SAWL — Sächsische Akademie der Wissen- schaften zu Leipzig, Arbeitsgruppe Limnologie, Leng- feld, Germany; USNM — United States National Mu- seum, Smithsonian Institution, Washington, D.C., USA; ZFMK — Zoologisches Forschungsinstitut und Museum Alexander Koenig, Bonn, Germany; ZIAS — Zoological Institute, Academy of Sciences, St. Petersburg, Russia In lists of studied material, capital letters E, L, P stand for exuviae, larvae, and pupae, respectively. Larval in- stars 1-4 are identified by the respective numeral fol- lowing the letter L. 3. KEY TO GENERA OF BLEPHARICERIDAE IN THE RUSSIAN FAR EAST The Far East Russian fauna comprises five genera. Ge- nus Neohapalothrix Kitakami is endemic to the eastern part of Asia, its closest relative being the European ge- nus Hapalothrix Loew. The position of these genera in the family is doubtful — see under Neohapalothrix Kita- kami. The other four genera belong to tribe Ble- pharicerini and exhibit amphi-Pacific disjunctions, with close relatives in western North America. In the Palae- arctic Realm, Bibiocephala Osten-Sacken, Agathon von Róder and Philorus Kellogg are restricted to the eastern part of Asia, while Blepharicera Macquart is wide- spread in the northern hemisphere. Adults l Branches of R on long fork (Figs. la-b) ................. 2 l' Branches of R sessile, or from short fork (Figs. Ic-e) 2 An unconnected branch M3 originating freely in wing membrane and ending in wing margin, and a basal crossvein between branches of M and Cu pre- sent(Pres: Dl ación Philorus 2' No unconnected vein in wing, no basal crossvein be- tween M and Cu (Figs. la) ............. Neohapalothrix 3 R with three long branches, anterior branch ending in Ry, shortly before wing margin; basal crossvein between branches of M present (Fig. Ic) .................. PR RE EEE RER IE RL OPEN Bibiocephala 3' R with two long branches; if a rudimentary anterior branch is present it ends in R, near base of latter and Bonner zoologische Beitráge 53 (2004) appears like an oblique crossvein forming a small triangular cell. Basal crossvein between branches of M present or absent (Figs. 1d, €) ann 4 4 Rudimentary branch R, delimiting small triangle present or absent; crossvein between M and Cu pre- sent (FiE: A A eet Agathon 4' Neither rudimentary R, nor crossvein between M and Cu present (Fig. le)... ee Blepharicera Neohapalothrix Philorus Sc R2+3 7 R1-3 Sa Bibiocephala RS @ M1 M3 él M4 te d Agathon Blepharicera e Fig. 1: Wing venation of Blephariceridae from the Russian Far East. a, Neohapalothrix manschukuensis (Mannheims) (af- ter BRODSKY 1954); b, Philorus asiaticus Brodsky (after BRODSKY 1972); c, Bibiocephala maxima Brodsky (after Bropsky 1954); d, Agathon japonicus (Alexander) (after KI- TAKAMI 1931); e, Blepharicera asiatica Brodsky (after BRODSKY 1976). Nomenclature of wing veins after HOGUE (1987); tc = triangular cell delimited on outside by R>,3. Pupae | Respiratory organs composed of four erect lamellae (Fig. 11); three or four pairs of abdominal adhesive MM iia 2 l' Respiratory lamellae in the form of short crests or flanges (Figs. 12d, e); three pairs of abdominal ad- hesive Organs a eee meee Philorus Respiratory lamellae resemble slender pointed trian- gles. Body cross-section approximately triangular, several abdominal tergites with median spines; four pairs of abdominal adhesive organs ........................ Pd: Neohapalothrix 2' Respiratory lamellae tongue- or leaf-shaped, apices rounded; body cross-section rounded, regularly curved, no dorsal armature; three pairs of abdominal [55] adhesive OF CANS: „un. omen 3 3 Body highly domed, cross-section semicircular ........ US Bibiocephala 3' Body moderately arched, cross-section lower than a halt Carle mn... 4 Peter Zwick & Tatyana AREFINA: Blephariceridae of the Russian Far East 337 4 All respiratory lamellae structurally similar, flexible. Dorsal surface matt, granulation strong, mesothorax with rugose zig-zag pattern... Agathon 4' Anterior and posterior respiratory lamellae hard, stiff, two inner ones soft. Dorsal surface shining, granulation fine and sparse, no zig-zag pattern on MINES OUNOLAR Go! A actinic see setvssecvasvcvvaccanes dens Blepharicera Larvae (instars 2-4, with one to six filaments per gill) 1 Head capsule entire, posterior notches insignificant ISP ANC HS An en ne 2 l' Rear of head capsule laterally distinctly notched, usually deeply divided (to eye-spot; Fig. 12 f) ....... 3 2 Antenna 2-segmented. Labrum sclerotized, forming part of a continuous prestomal ridge between left and right posterior mandibular articulations (Fig. 2b) earnest A Bibiocephala 2' Antenna 3-segmented. Sclerotizations above mandi- bles extend between the two mandibular articula- tions on each side but are medially separated from each other by the soft labrum (Fig. 2c) ...... Agathon 3 Body compact, abdominal segments lack “neck pieces”. Highly convex, body cross-section triangu- lar, dorsum of last instar with median row of spines. Two lateral pseudopods, both appressed to ground (L4), or single pseudopod forked (L3; Fig. 19e), or with knob (L2; Fig. 19d) ................ Neohapalothrix 3' Body not compact, abdominal segments 2-6 with narrow anterior portions (neck pieces), not highly arched, cross-section not triangular. Dorsal armature variable; if forming a median row it consists of blunt knobs or warts; dorsal pseudopods variable but DEVEIRADPLESSEA LO-FTOUNG enncotonacononiónsononorinócarcncanesa 4 4 Dorsal sensory pseudopod missing altogether, only ventral ambulatory pseudopod present (Figs. 16a, d). Dorsal armature, if any, consists of single blunt WartstalonSmidline .......0462006000000000.0 Blepharicera 4' Dorsal sensory pseudopod present, with dorsal proc- ess or erect spine; abdominal armature consisting of six structures per segment (Fig. 12 f) ......... Philorus Larvae (first instar) 1 Dorsum with continuous transverse crests ............. 2 l' Dorsum with transverse rows of separate erect little Selene Bes wl OA) ters ne. en een 3 2 Cephalic division with five crests, one on each tho- racic and two on 1% abdominal segment ................... IA LE Bibiocephala and Agathon 2' Cephalic division with two crests (one on thorax, one on 1* abdominal segment) ............ Blepharicera 3 Dorsal sclerites are little triangular spikes arranged in transverse rows; cephalic division with four of these rows. Pseudopod with apical brush of slender Seta eH (TCS) ee eis nee Philorus 3' Dorsal sclerites long and slender, band- or hair-like, in transverse rows; five of these on cephalic divi- sion. Pseudopod with group of retractile tiny hooks BIER IE da Neohapalothrix 4. TAXONOMY 4.1. Genus Bibiocephala Osten-Sacken, 1874 i Bibiocephala Osten-Sacken, 1874. Type species: Bibio- cephala grandis Osten-Sacken, 1874, by monotypy (OSTEN-SACKEN 1874: 564). Amika Kitakami, 1950. Type species: Liponeura infus- cata Matsumura, 1916, by original designation (KITA- KAMI 1950: 37). The genera Bibiocephala and Agathon were not sepa- rated by KITAKAMI (1931) but later he recognized their distinctness (KITAKAMI 1950). Unfortunately the no- menclature was confused; Amika Kitakami, 1950 is a synonym of the North American genus Bibiocephala. Kitakami erroneously used the latter name for what is in fact Agathon. Our present use of the name Bibiocephala is based on the redescription of the type species, B. grandis Osten- Sacken (HOGUE 1982) and conforms to the restricted use by HOGUE (1973, 1987) and ZWICK (1990: 1992). Bibiocephala and Agathon differ from other northern hemisphere net-winged midges by characters in which they resemble the southern hemisphere Edwardsininae: undivided larval head capsule (Figs. 2a, c, 5); presence of a remnant of vein Ro.; (Figs. Ic, d), secondarily ab- sent in individual species of Agathon; 4 visible growth zones of larval head capsule behind the cephalic sclerites, like in Edwardsina Alexander, 1920 (TONNOIR 1924; Zwick 1977, fig. 22) and the Madagascan Paulianina (Eupaulianina) Stuckenberg, 1958 (STUCK- ENBERG 1958, fig. 28); Philorus also similar. The first two agreements are undoubtedly plesiomorphic character expressions, the third probably so. Additional resemblances between Bibiocephala, Agathon and Philorus are difficult to interpret: dorsal larval pseudo- pods with erect outgrowth or process (Fig. 12f); in some species of Agathon, only a knob or angle remains; unique in the family; larval abdomen with six spines, sclerites or appendages standing in a characteristic ar- rangement (Figs. 8a, 12f); abdominal armatures are not uncommon in larvae of net-winged midges but in other taxa their arrangement differs. In any case, the monophyly of the genera in question needs be established. As constituent apomorphies of ge- nus Bibiocephala we accept the following characters: Larva: Massive highly domed body (Fig. 2a). Mouth area forming a completely flat oral field surrounded in front by an essentially continuous triangular prestomal margin (Fig. 2b): Sides of prestomal margin formed by 338 Bonner zoologische Beitráge 53 (2004) sclerotized arcs supporting mandibular articulations, the short transverse middle piece by the sclerotized, trans- versely folded labrum. Two setiform labral sensilla stand above, two below the sclerotized edge. In the groundplan of the family (for example, Fig.2c) the oral field is not flat and in front not surrounded by a con- tinuous sclerotized prestomal margin because the la- brum is largely soft and bulges forward and ventrad be- tween the sclerotized arcs to its left and right which support the mandibular articulations. At the medial end, these sclerotized arcs turn dorsad towards the lower end of the median frontal sclerite. Larva: Short stout lanciform or spatulate setae present on head in front of eyes, around antennal foramen, and on anterior corners of cephalic division (Fig. 5a; last in- star only). Pupa: Highly domed. Respiratory lamellae very large, often leafy. Adult: Body unusually setose, especially males. Ab- dominal segment one fused to metathorax, tilted up, immobile; mobility of abdomen achieved via well de- veloped membrane between abdominal segments one and 2. ¢: antenna short, compact; front femur arched; last ter- gite posteromedially sclerotized and spinulose. ©: Pleuron and tergite 9 enlarged, both with fringe of dense, exceptionally long setae surrounding abdominal tip like a basket (see ZWICK 1990, fig. 1). From the description by KITAKAMI (1931), the first in- star larva of Bibiocephala infuscata has the full com- plement of transverse cuticular crests, that is, three tho- racic and two abdominal ones on the cephalic division, and two each on the other abdominal segments. The pseudopod bears eversible hooklets. Fig. 2: a, Bibiocephala grandis Osten-Sacken, frontal view of last instar larva; b, B. maxima Brodsky, ventral view of head; c, Agathon eoasiaticus Brodsky, oblique frontolateral view of cephalic division, left antenna and maxilla removed. dp, dorsal pseu- dopod: g, galea: hy, hypopharynx; 1, lacinia; la, labrum; md, mandible; mxp, maxillary palpus; ps, ventral pseudopod; t, tentorial invagination. Peter ZWICK & Tatyana AREFINA: Blephariceridae of the Russian Far East Nearctic Bibiocephala belong to a single species, B. grandis Osten-Sacken, 1874, which is widespread in western North America (HOGUE 1982). The number of Asian species is uncertain, see under B. komaensis Kita- kami, 1950. We studied abundant material of B. maxima, B. infuscata, B. grandis as well as some larvae of B. komaensis. We summarise our knowledge in keys, which replace formal descriptions of the previously un- known larva of B. maxima. Keys also contain supple- mentary details on adults and pupae. There are excellent figures of male genitalia of B. grandis (HOGUE 1982, reprinted 1987) and of B. infuscata (KITAKAMI 1937), but dististyle structure of B. maxima in the existing fig- ure (BRODSKY 1954) is confusing. We draw and com- pare the closely related Asian species (Figs. 3, 4), mainly to facilitate a decision on possible synonymy be- tween B. komaensis and B. maxima, should fresh Ko- rean material become available. Fig. 3: Bibiocephala infuscata (a, b) and B. maxima (c, d). a, c, apical views of left dististyle; b, d, left dististyle and apex of tegmen in dorsal view. All to same scale. 4.1.1. Key to all known Bibiocephala-species Adult males (B. komaensis (Kitakami) not known) 1 Hypopygium elongated, its anteroventral margin po- sitioned much in front of anterior margin of tergite 9; anterior lobe of dististyle strongly spinose; inner dististyle with erect basal lobe on outside and simple band-shaped apex which is gently curved back; inner hypopygial sclerites partly rotated, penis filaments polonia ern. B. grandis Osten-Sacken l' Hypopygium not elongated, anteroventral margin level with anterior margin of tergite 9; anterior lobe of dististyle finely pubescent; inner dististyle with- out basal lobe; inner hypopygial sclerites and penis filaments ‘of normal lemsthy in... nee 2 2 Dististyle with massive apex, ventral lobe wide. In apical view, ventral lobe flat, much narrower than plump dorsal lobe; notch between lobes shallow, tri- angular (Figs. 3a, b). Apex of inner dististyle nar- 339 row, ribbon-like. Tegmen with broadly rounded spoon-shaped apex .......... B. infuscata (Matsumura) 2' Dististyle slender, both lobes slender and narrow; ın apical view, notch between them deeply U-shaped. Apex of inner dististyle narrow and pointed, semi- circularly curved (Figs. 3c, d), surface before bend with curved rugosities. Apex of tegmen narrowed, tongue-shaped nie B. maxima Brodsky Adult females (B. komaensis (Kitakamı) not known) 1 Eyes almost meeting on vertex, ocelli on raised tur- ret. Antenna shorter than head, beaded, flagellar segments two and following as long as wide, distal ones may be wider than long. Oviscapt ventrally with several large setae on bulging base, in addition to fine trichomes; laterodistal corners obtuse; para- median tips with falcate sensilla barely visible in ventral view. Lobes covering dorsal side of oviscapt short, broadly divided, covered with stiff trichomes about as long as width of lobe (Fig. 4a). Seminal re- ceptacles larger than oviscapt, basally gradually nar- rowed, coiled, eventually blending into long narrow UCT ee B. grandis Osten-Sacken l' Eyes widely separate on vertex, no ocellar turret. Antenna variable. Oviscapt without setae, only with fine trichomes; paramedian tips of oviscapt long, of- ten partly visible in ventral view. Lobes covering dorsal side of oviscapt narrowly separated, stiff trichomes brush-like, much longer than width of lobe. Receptacles sharply set off against their short CIS Sat ORC) tates tisha rhvncheanaqgaieraceacuabeeeereesonges 2 Antenna longer than head plus rostrum, setiform, flagellar segments longer than wide. Laterodistal corners of oviscapt obtuse (Fig. 4c, d). Receptacles pyriform, about twice as long as oviscapt, surface of receptacles everywhere with fine pale punctures ...... Fagan Sa ee hacen meee OSS B. infuscata (Matsumura) 2' Antenna short, subuliform, basal flagellar segments wider than distal ones, only first flagellar segment longer than wide. Oviscapt with prominent laterodis- tal corners. Receptacles oval, about as long as ovis- capt, (Fig. 4b) pale punctures only in basal half, dis- tal half with fine dark trichomes inside ..................... [59] Pupae 1 Gills normal, the lamellae diverge only slightly; granula on pupal dorsum spinous .............c:cccceeeeeees kN rast eis le TE B. grandis Osten-Sacken I' Gill lamellae diverge strongly; pupal granula A Fe Ld tev eea eh 2 2 Gill lamellae spread out widely, slightly curled, sug- gesting a cabbage leaf. Wide margins of lamellae pale, covered with visible plastron ......................... en B. infuscata (Matsumura) 340 Bonner zoologische Beitráge 53 (2004) 2' Gill lamellae divergent but in regular alignment, one behind the other. Lamellae smooth, not curled, uni- formly dark and shining, no plastron .....................- .. B. maxima Brodsky and B. komaensis (Kitakami) Fig. 4: Bibiocephala spp., oviscapts (a-c) and seminal recepta- cle (d), all to same scale. a, B. grandis, dorsal view of ovis- capt; b, B. infuscata, combined ventral (left) and dorsal views (right) of oviscapt; c, d, B. maxima, ventral view of oviscapt and one of the two lateral receptacles. Larvae (only last instar specimens with five to six gill filaments per tuft) 1 Six gill filaments per tuft; peristomal margin medi- ally notched (labrum shorter than laterally adjacent areas; Fig. 2a); dorsal pseudopods slender and long, directed obliquely backward; ventral face laterally from suckers with heavy dark spine-like setae; dor- sal face with flat sclerotized glossy calluses ............. BE EE B. grandis Osten-Sacken 1” Five gill filaments per tuft; peristomal margin entire (Fig. 2b); dorsal pseudopods short, pointing side- ways; ventral face laterally from suckers with short pilosity; dorsal face variable ici sau 2 2 Distinct sclerotized warts or calluses on dorsum; dis- tal antennal segment little longer than basal one. La- brum with several long setae in addition to four normal setiform sensilla . B. infuscata (Matsumura) (B. infuscata minor Kitakami, 1931 is said to differ from B. infuscata by paler pigmentation and smaller size. Bibiocephala jezoensis (Matsumura), 1931, from Hokkaido was synonymized with B. infuscata (Zwick 1990). Largest species of genus). 2' No dorsal sclerotizations but sometimes low unscle- rotized swellings present; labrum only with the nor- mal four setiform sensilla; antenna variable ........... 3 3 Distal antennal segment littler longer than basal one ae B. maxima Brodsky 3' Distal antennal segment almost twice as long as basälonen. anne B. komaensis (Kitakami) 4.1.2. Bibiocephala maxima Brodsky, 1954 (Figs. lc, 3c, d, 4c, d, 5) Bibiocephala maxima Brodsky, 1954 (BRODSKY 1954: 229) Our specimens from the entire known range match the description of B. maxima Brodsky, 1954 and we did not study types. Unfortunately, BRODSKY (1954) compared B. maxima only with Nearctic Bibiocephala. Bibio- cephala maxima occupies a vast range, from Kazakhstan and the Altai mountains to the Ayano-Mayski Region on the coast of the Sea of Okhotsk. Its southernmost Russian localities are not far from North Korea and Manchuria from where B. komaensis was recorded (see below). Bibiocephala maxima has clear to faintly tinged wings; despite its name it is smaller than B. infuscata. Material (See Table 1; all in PZ). RFEO3, 16 E; RFE06, 12, 49P, 6E; RFEOS, Russian Far East, Primorje, La- zovka r. 25 km W of Lazo, at bridge, 13.06.1998: 1E; RFEO9, 4E; RFE10, 2E; RFE14, 36P, 8E; RFE15, 30, 28P, 72E, 2L; RFE16, 4P, 2E; RFE21, 566, 39, 16 P, 31E; RFE27, 29; RFE28, 23, 19, 6E; RFE33, 2E; RFE35, 19, 1E; RFE38, 2E; RFE39, 2E; RFE40, 14; RFE41, 563, 39, 170P, 22E; RFE42, 83, 19; RFE47, 73, 149, 2P, 20E; RFE48, 24, 19, 6E; RFE49, 2P; RFESO, 48P, 2E, 1L4. Russian Far East: Ayano-Mayski Region, Aldoma R., 11/12.8.1999, M&T Tiunov, 2E. Khabarovsk Territory: Ochotsky Region, River Kuchtui, 10 km above mouth, 30.06.1999, 10.3°C, M.&T. Tiu- nov, 164, 19, 38P, 1L4. Kazakhstan, surroundings of Ust Kamenogorsk [49°58'N, 82°36'E], V. Devyatkov (PZ): River Kur- chum, 25.9.2002, 1 L3, 1L4; mouth of River Khamir (tributary of River Bukhtarma), 11.6.2003, 44, 19). Mongolia: Selenge, Khonin Nuga _ [49°05'260N, 107°17'440E], Eroo River, 21.V1.2003, P. Surenkhor- loo, 1 pharate Y, 8 pupal exuviae (PZ); Turgen-Gol [50%05'18"N, 91°37'50"E], 20.8.1997 and 17.8.1999, 8 L2, 22 L3, 4 L4, W. Hom & M. Paul, 272871999 (SAWL); Tesiyn Gol [49°39'29"N, 95°43'47"E], W. Horn & M. Paul, 27.8.1999, 2 L3 (SAWL). Larvae of B. maxima undergo considerable change dur- ing development. Second and 3" instar specimens have six short dorsal spines on each abdominal segment, their arrangement is the same as the dorsal calluses in B. in- fuscata, or of spines in Agathon spp. Last instar larvae Peter ZWICK & Tatyana AREFINA: Blephariceridae of the Russian Far East 34] of B. maxima lack dorsal sclerotizations, but spatulate setae on the fore body become important. A few larvae collected in autumn looked very unfamiliar because they are freshly moulted, their gut still empty, and body not fully extended. In this condition, spatulate setae on head capsule and corners of prothorax stand close to- gether and appear large, in comparison with the trunk; in places, the true body contour is veiled by setae (Figs. 5a, b). By spring, larvae accumulate much biomass whereby the body becomes extended, the head is pushed forward and downward; general shape is strikingly dif- ferent from autumn specimens. Spatulate setae on fore body stand widely separate and appear tiny against ex- tended body, remaining easily unnoticed. Figure Sc shows the single extended last instar larva not yet in prepupal condition that we have seen (Mongolian specimen); shape of its fore body resembles B. infus- cata. In prepupae, corners of cephalic division are ef- faced, because of further body extension. Some Far Eastern last instar larvae look slightly variegated. They are light brown with anterior portions of several body divisions darker brown. Fig. 5: Bibiocephala maxima, cephalic division of Mongolian last instar larvae. a, freshly moulted specimen showing ar- rangement of spatulate setae; b, same specimen, at same scale as specimen with extended fore body (c). Biological notes. Larvae and pupae of B. maxima live on the underside of stones under which current rushes through and are never found on the upper side; the pre- ferred larval habitat of B. grandis is similar (HOGUE 1982). Bibiocephala maxima emerges earlier than other net-winged midges in the Russian Far East. In early June, at several places, only fragments of pupal exuviae were left. Elsewhere there were many adults and pupae, but al- together only two prepupae were still found in June. The few earlier instars we saw were collected in autumn. Ap- parently the life cycle resembles B. infuscata, which hatches from eggs in autumn and overwinters in the 4" larval instar (KITAKAMI 1950). Hibernation of B. maxima larvae takes place under harsh environmental conditions. For example, winter temperatures are below -30°C near the Mongolian Turgen Gol, and a gauging station 25 km upstream from the sample site records no measurable dis- charge between December and March. The river bottom consists of coarse gravel with large near-surface intersti- tial spaces that may have contained flowing water also in winter (M. PAUL, pers. comm. ). HOGUE (1982) suggested that B. grandis might overwin- ter as eggs or very small larvae. However, the species probably has the same life cycle as the Asian congeners because large numbers of inactive, closely aggregated last instar larvae but no other life stage were collected immediately after snowmelt, in May, in the lower St. Vrain River, Colorado, and in several streams near Provo, Utah (P. Zwick). Indeed, 4" instar larvae were collected through the winter in Oregon, and even be- neath winter ice in the Canadian Rockies (G.W. COURT- NEY, pers. comm. ). Adult males resting in vegetation did not hang down from it like most other Blephariceridae. They often sat on grass stems, head up, grasping the support with all six legs. Because of the long hind legs, the body strongly diverged from the grass stem. Large swarms of only males were observed at Malinovka River with bin- oculars, in the morning and in the afternoon. Swarms were over land, 3-6 m above ground, above or on the sunny side of riparian willow bushes (Salix sp.). Speci- mens kept distances of 40-90 cm between each other, flying quietly in a floating manner. The front legs were held obliquely up and back between the wings, over the thorax, with the tarsal tips curved forward; short middle and very long hind legs hung down. Males that came close or other insects flying into the swarm, for example a fly (Calliphoridae), were rapidly approached from be- low by some swarm member and chased away. At evening dusk, numerous males were noticed flying over swiftly and smoothly flowing sections of the upper Nemtu River. They flew upstream for short distances, then dropped back again. Various pale insects emerging from the river, mainly small Ephemeroptera and Plecop- tera: Chloroperlidae were well visible under the low light conditions. Many of them suddenly changed direc- tion of flight, or dropped abruptly. When caught with a net, the pale insect had each time been seized by a fully pigmented male Bibiocephala. In two cases, however, the pale specimen was a freshly emerged Bibiocephala female. It appears that males emerge before females and catch these for copula, upon evening emergence. Where and for how long females live is unknown. We saw only few females, either at emergence, or spent and dying be- tween stones on the shore of River Khor, below Sukpai. Distribution. Bibiocephala maxima was first described from the Altai. It occurs even further west, near Ust- Kamenogorsk. It is also widespread and common in the Russian Far East. The new Mongolian records fit in well. ZASYPKINA & RYABUKHIN (2001) report unidenti- fied larvae of Bibiocephala from further northeast than the area sampled by us. 342 4.1.3. Bibiocephala komaensis (Kitakami, 1950) Amika komaensis Kitakami, 1950 (KITAKAMI 1950: 41). Material. 5. komaensis: 1 last instar larva, Korea, Ryeongha, Hamgyeongbuk-do (= Reika, N. Korea), 12.6.1936, leg. Kawamura. In the same vial are also IL2, 1L3, Hakugan, N. Korea, 6.9.1940, leg. Kitakami - (ZFMK); all three specimens are syntypes. | specimen of each larval instar, syntypes (no locality data; USNM). We were unable to borrow a larval syntype (Reika, N. Korea, leg. Kawamura 13.V1.1936) kept in Otsu Hydro- biological Station of Kyoto University (K. N. SATAKE 1988, pers. comm.). These few larvae seem to be all of the still remaining syntypes; we designate no lectotype. In the very old material we cannot assess slight differ- ences in body surface colour and gills between B. infus- cata and B. komaensis which Kitakami mentioned. In the last instar, the long distal antennal segment distin- guishes B. komaensis from B. infuscata (KITAKAMI 1950), and also from B. maxima. Details of the mouth region agree with B. maxima, and differ from B. infus- cata. Pupal exuviae of B. komaensis agree with B. maxima, by Kitakami's description; material is not available. Larval instars 2 and 3 lack dorsal spines, which are present in B. maxima; however, we saw few specimens. Whether B. komaensis is really distinct from B. maxima cannot be decided in the absence of pupae and espe- cially of adults of B. komaensis. The case of the two nominal species might resemble the species pairs Agathon eoasiaticus — A. kawamurai, Philorus levani- dovae — P. chosenensis, and Blepharicera parva — B. vamasakii, see below. 4.2. Genus Agathon von Róder, 1890 Agathon von Róder, 1890. Type species: Agathon ele- gantulus von Röder, 1890: l.c.: 232 (VON RÓDER 1890: 230). Apomorphies supporting the monophyly of 4gathon are found in the male genitalia: long widely separate male cerci connected only by a transverse basal bar (e.g., Fig. 6a). Mostly, cerci are slender straight rods although there is some variation in detail. For example, in 4. ele- gantulus cerci are curved mediad and apically narrowed (see figures in HOGUE 1973, 1987). In A. setosus Sp. n. and A. aylmeri (Garrett, 1923) (Nearctic) their slightly enlarged apices are gently turned ventrad, etc. Vesica with transverse apodeme, not a vertical one, as normal. Agathon was in the past confused with Bibiocephala be- cause a remnant of wing vein R) is retained in some species; it lacks in others. However, aberrant specimens occur in either group; they have a remnant of R; in one wing only. Bonner zoologische Beitráge 53 (2004) All Asian species are closely related and similar to the Nearctic type species. Outer dististyle bifid, shorter dor- sal branch arises basally and may end before, or extend over, subapical modifications on dorsal face of ventral branch. In 4. elegantulus and almost all other species a subterminal process or fold extends variably far across ventral branch so that a pocket or depression is formed between fold and apex proper (Figs. 7c, d, 9c, d, g). In- ner dististyle very wide, a largely membraneous foli- aceous structure (Figs 6a, 9h). Fig. 6: Agathon setosus sp. n. a, male genitalia in dorsal view; b, cerci and anal cone in oblique posterior view from left side, to show connection of cerci to sclerites supporting anal pouch. ac, anal cone; c, cercus; TIX, tergite 9; p, paramere tips; t, tegmen; id, inner dististyle; od, outer dististyle. The above diagnosis does not apply to 4. comstocki (Kellogg, 1903) and A. doanei (Kellogg, 1900) whose relationship remains doubtful. The genus Agathon re- quires further study; its relations with Asioreas Brodsky in the Altai and Himalaya will be discussed separately (ZWICK, in preparation). Each lobe of the female oviscapt bears distally a broad spinuliferous dorsal callus (not narrow and finger- shaped like in Bibiocephala). Three spheroid to pyri- form receptacles, basal half with tiny puncture-like pale spots, are sharply set off against short straight ducts. As usual, the median receptacle is more slender than the lateral ones. We name a new species, raising the number of Russian species to four, all of which are found also in the Far East. The Russian species have closely similar relatives in Korea and Japan, respectively (KITAKAMI 1931, 1950) with which they were previously never compared. We have carefully compared these species and consid- ered the Russian taxa are valid. Several large samples contained also first instar larvae, which by their association with later instars, can pre- liminarily be assigned to A. decorilarva and A. eoasiati- cus, respectively. We noticed no specific differences and none to distinguish them from first instar larvae of Bibiocephala. Peter Zwick & Tatyana AREFINA: Blephariceridae of the Russian Far East 343 4.2.1. Keys to Agathon species known or expected from the Russian Far East Adult males (Keys to females not attempted) 1 Dististyle almost entire, dorsal lobe appears only as a subapical tooth (Fig. 6a) .............. A. setosus Sp. n. 1' Dististyle deeply divided into dorsal and ventral [obesiventralone LONGED 22.2000 2 2 Dorsal lobe extends to or over subterminal tooth or crest on ventral lobe of slender dististyle ............... 3 2' Dorsal lobe of dististyle short, strong transverse crest across ventral lobe exposed. Ventral lobe api- cally very wide, transversely truncate (Fig. 9g) ........ AA cian dci A. ezoensis (Kıtakamı) 3 Ventral lobe of dististyle dorsally with subapical ON US AN A. decorilarva Brodsky 3' A subapical crest extends across dorsal face of ven- trallobejordististyle(ElS..D0) 4. desc. ceases acti asses .. A. eoasiaticus Brodsky, A. kawamurai (Kitakami) Pupae 1 Outer respiratory lamellae rounded with narrow base (racket-shaped), respiratory organs prostrate (Figs. NEO) ete eo A A. setosus Sp. n. l' Respiratory organs erect, lamellae slender, tongue- Sn fic Cnn re A 2 [59] Anterior end of pupa in dorsal view regularly trun- cate to rounded, cephalic sclerite not recessed be- tweenllateralllobes.of pronotum ............2.2.0se.002240.: BE SEO CELE EERE A. decorilarva Brodsky 2' Anterior end of pupa in dorsal view with cephalic sclerite recessed between anteriorly projecting lat- salas AAA sank A. eoasiaticus Brodsky, A. ezoensis (Kitakami), A. kawamurai (Kitakami) Larvae (final instar) 1 No spines or pointed granules on dorsum, only long soft setae on sides of body present (Fig. 8d) ............. NN cena rh cerro tado ida dera rias A. setosus Sp. N. l' Larval dorsum with pointed granules or long spines 2 Six long erect spines on each abdominal segment (RR EN EN A. decorilarva Brodsky 2' Dorsal armature consists of pointed granules not or hardly higher than width at base, and/or transverse LOWSTORETLECE SPITIULES. (cose. 222000002 sect ceseesee ¢ nspeenerene 3 3 Six well developed granules or short spines per ab- dominal segment (Fig. 8b); no rows of spinules. Thorax with single short spinule row immediately behind cephalic sclerites. Dorsal pseudopods with blunt.dorsalikneer.:......2....2 A. eoasiaticus Brodsky 3' Abdominal segments with two spinule rows, among which dorsal warts may or may not be distinct (Fig. 8c). Thorax with more spinule rows .................... 4 4 Three thoracic spinule rows (N. Korea) ...................- Seta Pe dead ore ree Pei A. kawamurai (Kitakam1) 4' Two thoracic spinule rows (Hokkaido, Sakhalin, Itu- TUB) ieee mene ee, A cis A. ezoensis (Kitakami) 4.2.2. Agathon decorilarva (Brodsky, 1954) (Figs. 7, 84) Bibiocephala decorilarva Brodsky, 1954 (BRODSKY 1954: 236). Material. 4gathon decorilarva was described from syn- types from: “Altaj: r. Yan-Tsili; r. Korbu; r. Kokschi; r. Bolschoj Korbu; r. Abakan; S. Primorje: r. Sitsa; Pigare- jeva gorge in St. Tigrovaja; Sutsan, r. Kamen; r. Kuatru, Kuretski Mts”. We studied some larval syntypes (ZIAS) labelled: Bibiocephala decorilarva sp. n. det. K. Brodskii // [illegible abbreviation] 64 [illegible abbre- viation] 78. The species is common and widespread in rhithral streams in the Russian Far East, see also ZASYPKINA & RYABUKHIN (2001). Material from localities in Table 1 (all in PZ): RFEO4, 1P; RFE06, 7P, 7L4; RFEO7, 57P, 11L4; RFEO8, 21P, 2L4; RFEO9, 1P; RFE12, 6E, 23L4, 28L3; RFE13, 55L4; RFE14, 17P, 22L4, 1L3; RFE15, 17P, 2314, 113: RFEIG,. UL4; REEI7, 24, 19, 131P, 59E, 8L4 ; RFE18, 8P; RFE19, 50P; RFE20, 41P, 10E, 214; RFE21, 60P, 1L4; RFE22, 14, 10P, 10L4; RFE25, 13, 1P, 1414; RFE26, 6P,-5E, 1L4; RFE27, 1¢,. 50P, 6E, 23L4; RFE28, 39, 19; RFE28, 52P, 18E, 114; RFE33, 18P, 4E, 1L4; RFE34, 1P; RFE36, 16P, 10L4, 1L3; RFE37, 21P, 3E, 2L4; RFE38, 1P; RFE41, 44, 80P, 65L4; RFE42, ca. 20P, 4L4; RFE43, 1P, 5L4; RFE44, 64L4; RFE45, 1L4, 11L3, 118L2, 1L1; RFE46, 41L4, 24L3; RFE49, 18P, 52L4, 10L3; RFE49a, 10L4, 5L3; RFESO, 2P, 8L4, 2L3. Additional material from the Russian Far East (PZ): Amur River Basin, IM. Levanidova: Ali River, 23.07.1959, 10L, 8P; Iski River, 02.07.1961, 4L. Maga- dan Territory, Makarchenko: Dukcha River, 7, 9.7.1979, 99L; 23.07.1979, 8L; Motyklejka River [Motyklejka 59°34'N, 148°38'E], 13,14.7.1977, 17L; Ulakan River, 17.07.1977, 6P, 2L. Ayano-Mayski Region [Ayan 56°27'N, 138°10'E], M&T Tiunov: Uika River 1 km up- stream of mouth, 21.-26.7.1999, 11L4, 14-15.8.1999, 9P; Ulujkan River, Lantar River basin, 29.07.1999, 19P, 8L4, 1L3; Aldoma River, 11/12.8.1999, 125, 89, 22P; Lantar River, 2.5 km upstream from mouth, 28.07.1999, 28,19, 28P; 214. Kamchatka, I.M. Levanidova (PZ): Kirpichnaya River, 18.7.1968, LIP, 4E; River Ewpitwajam (Lake Palan- skoje), 18.8.1963, 6P, 9E; nameless stream near village Zhupanova, 20.07.1964, 1L; Talalayevka River, 04.08.1972, Nikolayeva, 1L. Descriptive notes. BRODSKY (1954) described all sta- ges and both sexes. He compared A. decorilarva with 344 Bonner zoologische Beitráge 53 (2004) three Japanese species, 4. japonicus (Alexander, 1922), A. montana (Kitakamı, 1931), and A. bispina (Kitakami, 1931), none of which is really similar. We establish dis- tinctness of A. decorilarva from several much more similar Japanese species that were not considered by BRODSKY (1954). - Adult. R, present. Spur formula 0.0.2 in both sexes. Spurs on hind tibia very unequal in size. Male (Figs. 7a, c, d): Eyes widely separate, upper sec- tion little developed. Rostrum shorter than head height. Antenna short, with 14 segments, terminal one not lon- ger than penultimate and only slightly constricted. Geni- talia slender, outer dististyle with narrow apex and in- curved tip; subterminal fold short, almost tooth-like. Dorsal branch of dististyle curved, extends to close to subterminal fold. Female (Figs. 7b, e): Eyes large, narrowly separate, up- per section fully developed. Antenna even shorter than in male, barely reaching tip of rostrum. Rostrum only as long as head height but fully developed. Oviscapt with slightly concave sides and tongue-shaped distal lobes; spermathecae subspherical to ovoid. Fig. 7: Agathon decorilarva. a, b, head of pharate male and female , respectively; c, male hypopygium (tergite 9 and cercı removed); d, oblique dorsal view of left outer dististyle; e, fe- male terminalia. Not to scale. Pupa. Typical of genus, zig-zag pattern across thorax. Very similar to 4. eoasiaticus (see BRODSKY 1954: figs. 4/6+7), respiratory lamellae less than twice as long as wide at base, lamellae set in an oblique row so that op- posite first lamellae are wide apart while lamellae 4 are closer together,.their slightly recurved tips meet, or al- most so. Bases of lamellae on the outside terminating in integument as simple straight lines. The endpoints of the four lamellae lie on a slightly convex line because la- mellae 2 and 3 project slightly further sideways than lamellae 1 and 2. Cephalic sclerite not distinctly re- cessed between fore leg sheaths. Larva. Long erect spines, anterior ones longer and closer together than posterior ones (Fig. 8a). Lateral ones on segment seven smaller, about half the length of dorsal ones. Integument very variably contrastingly pat- terned, from mainly yellow to mainly dark. Also spines (all or only some) dark or yellow. Head capsule variably patterned in yellow and dark, often an extended pale center near anterior end of median frontal sclerite, lat- eral frontal sclerites with distinct yellow eye-brow-like mark above oblique suture. However, head capsule may also be almost completely black. Ventral pseudopods slender, with narrow recurved apex, posterior margin distinctly concave. Dorsal pseudopods slender, with erect narrow pointed subterminal outgrowth on narrow tip of pseudopod. Fig. 8: Agathon ssp., cross-sections through larvae, 3 ab- dominal segment from behind, diagrammatic. a, A. decori- larva; b, A. eoasiaticus, c, A. ezoensis; d, A. setosus. Not to scale. Comparisons. Agathon decorilarva is a close relative of (but certainly distinct from) two Japanese species whose larvae also have long erect spines and sharply angled dorsal pseudopods. Agathon bilobatoides (Kita- kami, 1931) has only the four central spines large. It al- so possesses additional transverse rows of abdominal thornlets, also three rows on the thorax. The subterminal fold on the male dististyle is long, standing in an obli- que position. The pupal gill lamellae are closely packed, all semicircularly curved towards middle, with tips of opposite organs meeting. Agathon longispinus (Kita- kami, 1931) has the median spines on the larval ab- dominal segments 2 and 3 connected by a transverse bar-like socket, and lateral spines on segment 7 are tiny, much shorter than on the other segments. Pupal lamellae Peter Zwick & Tatyana AREFINA: Blephariceridae of the Russian Far East 345 more slender than in 4. decorilarva (>3 times longer than wide at base), divergent, from 1“ to 4" succes- sively more curved and directed medially, tips of 4" la- mellae meeting. Male with long antenna (reminding one of A. eoasiaticus), dorsal branch of male dististyle short, with strongly widened apex. In the Russian Far East, A. decorilarva and A. eoasiati- cus often coexist; pupae can be distinguished by the re- lative length of the cephalic sclerite versus leg sheaths. This character can no longer be assessed in pupal exuviae which can, however, be identified by the shape and po- sition of the outer endpoints of the four respiratory la- mellae. 4.2.3. Agathon eoasiaticus (Brodsky, 1954) (Figs. 2c, 8b, 9) Bibiocephala eoasiatica Brodsky, 1954: 234). 1954 (BRODSKY Material. We identified the species from the description and saw no types. We did not find the species at the type locality ("St. Tigrovaja, tributary of r. Sitsa; Pigare- jeva gorge") but Stations RFEO1 and RFEO3 are close to it: RFEO1, 11.06.1998, 330L4, 25L3; 26.06.1998, 3 3, SP. 79EA, 1213; REEO9, 1L4, 1L3; RFE10a, 7P, SESREE12, M4, 2L3; RFEÍS, 18, 39, 1P, 4E; RFE19, 3P.3E; REE20, AP, 1E; RFE21, 1L2; RFE22, 3d, 29, 22SP29E., 2014; REE24, 19, IP, 2E; RFE25, 29, 2E; RFE27, 39, 1P, 2E; RFE28, 1P; RFE31, 10d, 39, 129P, 14E, 8L4; RFE33, 19, 1P, 3E; RFE34, 43, Q; [BaP e20E SLA: RFE36, 28P, 23E; RFE37,-6P, 12E, 112, 1L1; RFE41, 53, 39, 2P, 94P, 7E, 22L4; RFE42, 22P, 5L4; RFE44, 38L4; RFE49, 92P, 27L4; RFE49a, IRSA REES4, 12P, SE; RFESS, 99,1E, 114 (all in BZ): Mongolia, Selenge, Mandal, leg. P. Surenkhorloo (PZ): Khonin Nuga [49°05'260N, 107°17'440E], Tsagaan Chubut River, 30.V1.2003, numerous L3, L4, 8 P. Bar- Chuluut River [48°58'638N, 106°57'013E], 13.V11.2003, 6P (incl. 34). Bar-Chuluut River [48°58'368N, 106°59'564E], 4.VIII.2003, 23, 22. Eroo River near confluence with small tributary [49°05'260N, 107°17'440E], O1.VII.2003, 2 L4. Ataa River, 18. VII.2003, 56P (incl. pharate 44), 3Ex, 3L4, 1L3 (all in PZ). Adult. Previously unknown. R> present. Spur formula 0.2.2 (females) and 0.0.2 (males). Spurs on hind tibia unequal in size, those on female middle leg minute, not easily seen. Male (Figs. 9a, c, d): Antenna more than twice as long as head with rostrum, individual flagellar segments long, last one strongly bottle shaped. Genitalia compact, hypopygium rounded. Distal edge of dorsal dististyle wider than in 4. decorilarva, truncate, prominent sub- terminal fold across Y of dististyle width. Fold hidden by longer, slightly club-shaped dorsal branch of dististyle. Female (Figs. 9b, e): Very similar to 4. decorilarva, rostrum and antenna slightly longer. Oviscapt appegrs slightly more elongate but is not distinctive. Receptacles subspherical to ovoid. Pupa. Resembles 4. decorilarva, but cephalic sclerite distinctly recessed between leg sheaths. Bases of lamel- lae terminating in integument as simple straight lines. On outside, endpoints of lamellae 2-4 lie in straight line while lamella 1 projects distinctly further sideways and ends in small backwardly turned hook. Larva. Short brown or yellow spines on pale sockets (Fig. 8b), or only distinct cones on abdominal segments, sometimes two or three small cones together instead of spine. Anterior spines or cones higher and more widely apart than posterior ones. Integument greyish, not con- trastingly patterned. Head capsule with pale center near lower end of median frontal sclerite, remainder amber to dark brown; oblique suture across lateral frontal sclerites sometimes apparent as pale narrow line. Thornlets in transverse rows, tiny, but of variable ex- pression; all abdominal and three thoracic rows may be present. Dorsal pseudopod plump, with obtuse rectangu- lar knee instead of a dorsal process. Comparisons. Agathon eoasiaticus, the Korean A. ka- wamurai (Kitakami, 1950), and the Japanese A. ezoensis (Kitakami, 1950) and A. bispinus (Kitakami, 1931) are similar. The last is from Honshu and not available to us; only larvae and pupae were described. Male genitalia of A. ezoensis are distinct (see below) but those of A. kawamurai are not. Females (all with large upper portions of eyes, with large reddish ommatidia) and pupae are so similar that in view of the limited ma- terial available of the Japanese and Korean species, we attempt no distinction. Larvae can be distinguished by the dorsal armature of thornlets and spiny warts. KITAKAMI (1950) used pres- ence/absence and number of transverse rows of thornlets on thoracic and abdominal segments and the relative size of six dorsal warts to distinguish the taxa. In A. eoasiaticus, the six dorsal warts are all larger than the thornlets, the anteromedian pair being largest. In the other species (Fig. 8c; A. ezoensis), at best the lateral warts are distinct while the dorsal ones are effaced and barely larger than spinules in the transverse rows. How- ever, A. eoasiaticus varies, both within and between samples. There may be 0-3 rows of thornlets on the tho- rax, and none, indistinct, or two distinct rows on the ab- domen. The main warts may be prominent and stand by themselves, or be less distinct from adjacent thornlets. Warts may have several points each, or essentially con- 346 Bonner zoologische Beitráge 53 (2004) sist of a closely set group of thornlets. The distinction of larvae remains therefore provisional until more material of the relatives of A. eoasiaticus becomes available. 300 um 7 Kurt Fig. 9: Agathon eoasiaticus. a, b, head of pharate male and female, respectively; c, male hypopygium (tergite 9 and cerci removed); d, dorsal view of left outer dististyle; e, female ter- minalia. A. ezoensis. f, g, left outer dististyle in ventral and dorsal views; h, same as g, inner dististyle added. Not to scale. 4.2.4. Agathon kawamurai (Kitakami, 1950) Bibiocephala kawamurai kawamurai Kitakami, 1950 (KITAKAMI 1950: 44). 7 Material. Bibiocephala kawamurai kawamurai Kit., 16 (lectotype, here designated), 12, 2P (genitalia of pharate © on slide), 2L4 (paralectotypes, here designated), Korea, Baekam, Hamgyeongbuk-do (= "N. Korea, Hakugan"), 23.V11.1939 (ZFMK); additional paralectotypes; 14, 19, 2P, 2L4 (one lacking cephalic division) (USNM). Taxonomy. KITAKAMI (1950) suggested “perhaps we can expect this species in Manchuria”. Male genitalia do not differ from A. eoasiaticus. The nominal taxa may nevertheless be distinct because the rostrum of 4. kawamurai attains not even head height. Agathon eoasi- aticus (Fig. 9a) has also longer antennae than A. kawa- murai. Larvae of A. kawamurai have the full set of transverse rows of prominent thornlets among which dorsal warts are indistinct. However, larval armature of A. eoasiaticus is variable. The variation of the wide- spread A. eoasiaticus may eventually be found to en- compass the structural variants presently distinguished as A. kawamurai, which would become the valid name. Comments. KITAKAMI (1950) made no distinction be- tween specimens known to him, which therefore are all equal syntypes. Consequently, the specimens in USNM must have type status, even though their precise collec- tion data are unknown. We have not seen syntypes from Ryeongha, Hamgyeongbuk-do (= Reika) in North Korea in Otsu Hydrobiological Station: 2L3, 2L2, 12.V1.1936 (Kawamura) and 14, 19, 2P, 2L4, 22.VII.1939 (Kita- kami). 4.2.5. Agathon ezoensis (Kitakami, 1950) (Figs. 9f-h) Bibiocephala kawamurai ezoensis Kitakami, 1950 (KI- TAKAMI 1950: 46). Material. Bibiocephala kawamurai ezoensis Kit., 14 (lectotype, here designated) 2P (paralectotypes) [Takinozawa, S. Saghalien, 20.V11.1936]: 2L4 (para- lectotypes) [Sóunkyo, Hokkaido, 25.VII.1936] (ZFMK). Additional paralectotypes without loca- lity data: 14, 1P, 2L4 (USNM). Additional material (PZ): Kuriles, Southeastern Iturup Isl.: Usach waterfall, 29.07.1998, T. Arefina & V. Teslenko, 17, 10P, 6L4, 1L3: unnamed stream with wa- terfall nr Usach waterfall, 29.7.1998, T. Arefina, 44, 12, 3P. Very similar to A. eoasiaticus but Rz normally absent. However one of the males in USNM and also one from Iturup Island have a remnant of R> in one wing. Male genitalia (Figs. 9f-h) distinguished by outer dististyles with very wide apex and a short dorsal branch not closely appressed to larger ventral branch. Subterminal transverse fold across ventral branch of dististyle long, prominent, and exposed. Its caudal edge curves mediad and connects to apical edge, the curved connection projecting distinctly beyond general contour of dististyle. Inner dististyle wide, medial part distinctly elongated and projecting. Presently females not separable from A. eoasiaticus by external characters because lack of R> in A. ezoensis is no reliable difference. Receptacles not examined. For pupae and larvae (Fig. 8c), see under A. eoasiaticus. Remarks. Like A. kawamurai, this taxon was named from many syntypes from several localities in Hokkaido and on Sakhalin, between which no distinction was ma- de in the description; therefore, the specimens in USNM are syntypes even though locality data are missing. The locality information for the specimens in ZFMK (in square brackets) is from the list that accompanies KITA- KAMI'S material in coll. MANNHEIMS. Peter Zwick & Tatyana AREFINA: Blephariceridae of the Russian Far East 347 4.2.6. Agathon setosus sp. n. (Figs. 6, 10, 11) Material. Russian Far East: Holotype (pharate <), para- types (6 pharate Í, 2 pharate Y, 9P), [2773] Primorje, Sikhote-Alin, Edinka River, 10.08.1979, leg. Timoshkin. (Presently in PZ, will eventually be deposited in a mu- seum). Additional material: Primorje, Sikhote-Alin, Ada River, Bikin River basin, 22.07.1979, 4L, leg. Timoshkin. — Magadanskaya obl., Talok Stream at Susuman Town, Kolyma River basin, leg. Samokhvalov, 10.4.1982, 17 L; Khabarovskii krai, Basilii Stream, Tumnin River ba- sin, 16.7.1997, leg. Tiunov, 1L. Mongolia, Selenge, [49°05'260N, 107°17'440E], Eroo River near conflu- ence with small tributary, 01.V11.2003, P. Surenkhorloo, 5P, 8L4 (including prepupae) (all in PZ). Adult. Adults apparently similar to previous species in size, venation, and colour; details unknown because on- ly pharate specimens are available. Spur formula 0.0.1 in both sexes, both with small, widely separate eyes (Figs. 10a, b). Upper eye portion distinct but only small part of total eye area and not larger-facetted than lower part. Female with, male without functional mandibles and serrate hypopharynx. Antenna and palpus slender, much like A. eoasiaticus. Male genitalia (Fig. 10c): Widely separate, long and slender cercus (Fig. 6) distally slightly enlarged and gently curved ventrad. Outer dististyle slender and nar- row, with faint mediobasal swelling. Dorsal branch fused to ventral one over most of length, free tip resem- bles subapical tooth on ventral branch. Apex of ventral branch appearing simple because transverse fold practi- cally apical, vertical fold separating it from ventral face visible only in apical view. Inner dististyle slender, more sclerotized and narrower than in relatives. Female genitalia (Fig. 10d) similar to other species, oviscapt appears shorter and stouter. Seminal recepta- cles distinctly pyriform. Pupa (Fig. 11). General shape normal except respira- tory organs prostrate, less raised than in majority of spe- cies, extending distinctly further forward than long sheaths of fore legs which include wide straight front margin of cephalic sclerite between them. Outer lamel- lae racket-shaped, base narrower than distal part, stiff and shiny, distinctly wider than soft medial lamellae. Mesonotum with strong zig-zag folds. Dorsal granula- tion of metathorax and abdomen strong in middle, in- creasingly finer towards sides, lateral granules only 1/4 size of middorsal ones. ei Na Fig. 10: Agathon setosus sp. n. a, b, head of pharate male and female respectively; c, male hypopygium (tergite 9 and cerci removed); d, female terminalia. Not to scale. Larva. General shape resembles relatives. Cephalic sclerites amber, triangle above mandibles black, brown seams along anterior tentorial invaginations and along outer edges of medial and lateral frontal sclerites. Me- dial frontal sclerites additionally with faint traces of two usual brown longitudinal lines. Antenna 3-segmented. Body naked except long flexible setae arising from dis- tinct insertion points, mainly on posterior segment halves. Ventral pseudopods simple, dorsal ones almost simple, only faint angle in dorsal contour (Fig. 8d). Comparisons. The male genitalia are exceptional be- cause the dorsal branch of the dististyle is not a freely projecting finger. Apex of outer dististyle and relatively narrow inner dististyle also unique. Pupa distinctive through its prostrate respiratory organs with stiff rounded outer lamellae much wider than median ones. Larva unusual in having long slender erect fine setae, instead of the usual cones, spines or stout thornlets. At first glance, larva appears naked and insofar resembles A. montanus (Kitakami, 1931) from Japan which, how- ever, has prominently angled pseudopods. In particular, its exposed pseudopod 7 is so positioned that it resem- bles a backwardly bent knee. In contrast, pseudopods of A. setosus are almost simple. 348 Bonner zoologische Beitráge 53 (2004) 1mm 500 um Fig. 11: Agathon setosus sp. n. a, pupa in dorsal view; b, front end seen from left side; c, left respiratory organ, lamella 4 partly cut to expose lamellae two and 3. Figures a and b to same scale. 4.3. Genus Philorus Kellogg, 1903 Philorus Kellogg, 1903 (KELLOGG 1903: 199). Type species: Blepharocera yosemite (Osten-Sacken, 1877), by subsequent designation of Coquillett, 1910 (Co- QUILLETT 1910: 588). The phylogenetic relations of Philorus among Ble- pharicerini are doubtful because the genus still shares presumably primitive characters with Bibiocephala and Agathon; see under Bibiocephala. At the same time, the radial vein is more reduced (two short branches on a long stalk, instead of long independent branches) than in other Blepharicerini, supporting monophyly of genus Philorus. The pupal respiratory organ of four low, widely divergent lamellae is another obvious apomor- phy. The genus comprises almost 25 species from mainland Asia (south to Assam and west to Tian Shan), Japan, and western North America. Grouping of included spe- cies is not yet generally clear. However, the new Rus- sian species clearly belongs to a monophyletic group distinguished by: lateral digitiform appendages on male tergite 9; male with extremely short ventral bridge from which the bag-like, uncrested vesica extends freely for- ward; pupal respiratory organs medially touching, la- mellae very low, crest-like, divergent. The name Euliponeura Tonnoir, 1930 would be avail- able for this group which comprises P. asiaticus Brodsky, 1972, P. assamensis (Tonnoir, 1930), P. cho- senensis Kitakami, 1931, P. horai (Tonnoir, 1930), P. longirostris Kitakami, 1931, P. minor Kitakami, 1931; not all of the three characters listed above are known for each included species. Recognising Fuliponeura would turn the genus Philorus paraphyletic because the last two apomorphic characters in the above list are shared by P. vividis Kitakami, 1931 (ZWICK, 1997a). A com- plete phylogenetic analysis of the genus must be per- formed before any formal change of taxonomic status of included groups. 4.3.1. Philorus levanidovae sp. n. (Figs. 12, 13) Material: Holotype pharate Í, paratypes 1 pharate Í, 2 pharate Y, 4¢, 89 plus 4L1, 18L2, 35L3, 16P, 3E (some on slides) without type status: Russian Far East, Khasanski Region, Gryaznaya stream, Kravtsovski Cas- cades, 12. and 17.07.1998, T. Tiunova and T. Arefina (presently in PZ). Additional material: Russian Far East, Primorje, small tributary to Vodopadnaja River at forest edge nr Niko- layevka, E of Partizansk, 12.06.1998, 5L1, 13L2, 8L3, 20L4 (PZ). Adult. General structure, especially wing venation, ty- pical of genus. Small, wing length: 4 5.5 mm, Y 7.5 mm. Pale, dorsal aspect of head, thorax and abdomen light brown, antennal flagellum and male genitalia brown, remainder yellowish. Eyes widely separate, a tiny elongate-oval portion of eye towards ocelli more reddish, more finely facetted than the rest from which it is not set off by line of separation. Ocelli on flat wide frons, not raised on turret. Antenna with 13 segments, the 11 flagellar segments elongate and asymmet- rical: dorsal contour straight with short seta at base, ventral side with distinct basoventral swelling. Several structures sexually dimorphic. Antenna: each segment about 2.8-3.2 times as long as maximum width in males, 2.6-2.8 times in females (pharate specimens). Last segment 3.7x longer than wide in males, but simi- lar to other segments in females. Mouthparts: rostrum of male shorter than height of head, no mandibles, no ser- rate hypopharynx. Palpus of © short, segment four with prominent subterminal swelling, segment five long, whip-like, bare except some setae on convex face. Fe- male rostrum slightly longer than height of head, with serrate hypopharynx and mandibles, mandibles longer than labrum. Palpus of 7 generally similar to male but much longer. Scattered lateral setae along base of la- brum in female, male without. Legs long and slender, unmodified. Tibial spurs hairy, spur formula 0.0.1 in males, 0.0.2 in females, spurs on hind tibia very un- equal. Male genitalia (Figs. 12a, b): Epandrium transverse, broad, setose, particularly strong setae bordering con- cave rear edge. Cercus simple, conical, setose, proctiger Peter ZWICK & Tatyana AREFINA: Blephariceridae of the Russian Far East 349 with two setae projecting below. Hypopygium trans- verse and short, distal margin in middle slightly triangu- lar. On either side of dorsal face of gonocoxite arises sclerotized erect finger with strong apico-medial seta- tion. Outer dististyle profoundly bilobed. Dorsal branch with blunt setose medial projection at base, apex of branch slender, bent anteromediad, with strong apical spines. Ventral branch largely soft, surface covered with microtrichia, rough. From an apical oval lobe with few large ventral setae hangs dorsally flat but ventrally keeled appendage. Band-shaped medial edge of append- age with microtrichia, semicircular lateral portion with hyaline outwardly directed spicules. Inner dististyle slender in form of dorsally hooked rod. Tegmen not ap- parent. Inner sclerites: ventral bridge shallow, vesica in form of oval sac without apodeme; three subequal penis filaments. Parameres longer and wider than penis fila- ments, apically widened, canal distinct, with funnel- shaped apical opening. Fig. 12: Philorus levanidovae sp. n. a, male genitalia, dorsal view, left finger-shaped process cut; b, right dististyle, ventral view; c, female oviscapt; ventral view to the left, dorsal view to the right of the interrupted vertical line; the single curved spine on the side of one oviscapt lobe is apparently abnormal; d, pupa, dorsal view; e, pupal respiratory organs, dorsal view; slide preparation, position of lamellae caused by pressure of cover slip; f, last instar larvae, anterior and posterior body por- tions, dorsal views. Female genitalia (Fig. 12c): Sternite 9 forms two semi- circular lateral lobes with long setation including ton- gue-shaped base of oviscapt between them. Oviscapt plump, lobes rounded, separated by U-shaped notch. Microtrichia on ventral oviscapt face arranged in regular curved lines, middle strip bare. Dorsal face of each o- viscapt lobe with about 15 grouped stout short curved setae just beyond apex of tongue-shaped process origi- nating at oviscapt base; apex of process with slender microtrichia. Three pyriform spermathecae with short simple ducts. Pupa. 4.7-6.5 mm long, flat. Pale brown. Granulation exceedingly fine and pale. Contour ovoid, front blunt, little wider than rear (Fig. 12d). Respiratory organs meeting medially, lamellae reduced to shallow crests. Outer lamellae form a low transparent rim around rough and partly wrinkled respiratory field. Slightly higher soft central lamellae 2 and 3 differ in length and dis- tance from margin (Fig. 12e). Straight, open spiracular slit separating central lamellae; outer end of slit termi- nating in visible spiracular pore. Larva. Last instar up to 7.5 mm long (Fig. 12f). Ce- phalic sclerites distinctly divided. Antenna about as long as cephalic sclerites, with long pale median zone. An- tenna 2-segmented, separated by long ill-defined articu- lar membrane. Sometimes, dark basal segment with nar- row pale zone in middle, antenna then seemingly 3-seg- mented. Thoracic segmentation indistinct because segmental transverse rows of little erect spines on ce- phalic body division are irregular and somewhat ob- scured by numerous additional spinules. Abdominal ter- gites 1-6 with six pointed cones, each with several small spines at top. Two paramedian cones anteriorly, the other four widely spaced in posterior transverse line. Addi- tionally, a pointed projection on elongate dorsal feeler situated above simple pseudopod. Segment 7 lacking pseudopod as well as cone on dorsal feeler. Remnant of tergite 8 with two small raised warts. Posterior margin of anal body division shallowly bilobed. Ventral face typical, six suckers, six pairs of gill tufts, five filaments in each; four rectal papillae surrounding last sucker from behind. Gills are so positioned that they are not normally visible in dorsal view. Lobes of rear body edge with shiny medially divided sclerite band. Instars 3 and 2 similar, with three and one gill filament per tuft, respec- tively. Cephalic sclerites in instars 1-3 with distinct poste- rior growth zones. First instar larva (Fig. 13). No gills; rectal papillae present. Elongate egg burster on median frontal sclerite. Body surface finely wrinkled and with tiny flat cuticular spikes arranged in transverse slightly raised lines, crest- like. Long crest on only two of three thoracic segments and on abdominal segments 1-7. This long crest also in- cludes pair of tiny cone-shaped and minute hairlike sen- 350 Bonner zoologische Beitráge 53 (2004) silla (tergal and subtergal sensilla, HOGUE 1978). Later- ally, crest ends on small hump with 2-3 stronger spines above pseudopod (probably tergopleural sensilla of HOGUE 1978). Abdominal segments 1-7 additionally with short anterior crest lacking sensilla. Rudimentary segments 8 and following without crests and sensilla. Base of pseudopod with two, rarely three spines and tiny hair sensillum. Distally, each pseudopod with sin- gle very long seta and about 12 transparent rearward di- rected slender spines. Length of spines in each group in- creases from front to rear. sabana ba 44 Ff > o KARA Loe Ady rro ryan bhradadn y Veen Fig. 13: Philorus levanidovae sp. n., first instar larva, dorsal view. Habitus with enlarged insets of dorsolateral armature and of apex of pseudopod (large seta clipped). Comparisons. Because larvae of P. /evanidovae and P. chosenensis cannot presently be distinguished, speci- mens from a tributary to Vodopadnaja River are tenta- tively assigned to P. levanidovae which is the sister- species of P. chosenensis Kitakami from Middle Korea; see there for distinction of males and pupae. Members of the so-called Euliponeura-group and P. vividis Kita- kami have very similar pupae with medially meeting respiratory organs with low, crest-like lamellae and a spiracular slit with visible spiracular pore. Larvae of this group have similar dorsal armature; Japanese species are distinguished by forwardly shifted gills visible in dorsal view. First instar larvae of P. levanidovae agree with P. vividis (KITAKAMI 1931, fig. 2) and P. taiwanensis Ki- takami (KITAKAMI 1941b, fig. 2), but figures of the lat- ter two species are not sufficiently detailed to compare integumental ornamentation. Habitat. Kravtsovka stream flows in a narrow canyon shaded by forest; it forms many rapids and cascades. Philorus levanidovae was the only species of Ble- phariceridae found in the stream, larvae and pupae were collected from the hygropetric zone on big boulders, only a few larvae were found on pebbles in the main flow. Adults were swept from riverside vegetation. The small tributary to Vodopadnaja River flows in the shade of the forest edge; the collecting site was a few meters before the stream enters the Vodopadnaya River. Speci- mens were fully submerged in the main flow of the streamlet in which there were no other Blephariceridae. Conversely, no specimen of P. levanidovae was taken at a few meters distance in the fast-flowing unshaded Vodopadnaja river where Agathon, Bibiocephala and Blepharicera were common. The simultaneous presence of all life stages may suggest several overlapping gen- erations, like in P. chosenensis. Etymology. We are pleased to name this new Russian species in the honour of the late Dr. Iya M. LEVANI- DOVA, Vladivostok, much admired pioneer scientist and grand old lady of Far East Russian freshwater research. 4.3.2. Philorus chosenensis Kitakami, 1931 Philorus chosenensis Kitakami, 1931 (KITAKAMI 1931: 93); 1950 (KITAKAMI 1950: 64, figs. 40-43). Material. Lectotype (present designation) adult male, paralectotypes | pupa, | last instar larva (ZFMK); syntypes from Mt. Geumgang, Gangwon-do (= “Mt. Kongo”), Middle Korea, 25.V11.1939. — Addi- tional material: 1 adult male, 1 pupa, 1 last instar larva (USNM), authentic material identified by Kitakami, possible syntypes. P. chosenensis 18 thermophilic, hygropetric and plurivoltine (KITAKAMI 1950). The species is very simi- lar to P. levanidovae; we saw no females. Males can be distinguished by the genitalia; Kitakami's figure 1s cor- rect but too small to recognize details and consequently we provide a new illustration (Fig. 14). The above specimens appear to be all that still exist from Kita- kami's collection and those that are definite syntypes are here designated as lectotype and paralectotypes, respec- tively. Only characters distinguishing P. chosenensis from P. levanidovae sp. n. are redescribed. Adult. Male genitalia (Fig. 14): Dorsal branch of outer dististyle with broad but very blunt basal projection; apex short, rounded, not overhanging. Apex of ventral branch flat, resembling curved ribbon whose microsculpture changes from basal microtrichia to apical spinules but me- dial and lateral faces are similar. Inner dististyle sinuous, apex curved mediad, not dorsad. Penis of male in USNM is erect; parameres with small ovoid terminal section, enlargement being in parasagittal plane. Peter ZWICK & Tatyana AREFINA: Blephariceridae of the Russian Far East Pupa. Similar to P. levanidovae except fine but dark and distinct granulation. Larva. Similar to P. levanidovae, no distinctive charac- ters known. Fig. 14: Philorus chosenensis Kitakami, male lectotype. a, ge- nitalia, dorsal view, left finger-shaped process cut; b, same, right half, ventral view. 4.4. Genus Blepharicera Macquart, 1843 Blepharicera Macquart, 1843 (MACQUART 1843: 61). Type species: Blepharicera limbipennis Macquart, 1843, by monotypy, a junior synonym of Asthenia fasciata Westwood, 1842. Blepharicera is the most widespread genus of the fam- ily. It falls into several species groups (ZWICK 1990, ZWICK & ZWICK 1998) of which only the fasciata-group occurs in East Asia. The single species from the Russian Far East is new to science. 4.4.1. Blepharicera parva sp. n. (Figs. 15, 16) Material. Holotype, | pharate Í, paratypes 5 pharate <, 9 Y, River Manoma, basin River Anjui [a tributary of Amur River], leg. Arefina & Tiunova, 28.07.1996; the same sample also includes 31 P, 30 L3 & L4 which ha- ve no status as syntypes (leg. IBSSAN; presently in PZ). Additional material: RFEO3, 115 L1-L4; RFEO9, 25L1, 67L2, 85L3, 137L4, 34P; RFES6, 42 L2-L4 (incl. pre- pupae); RFE10, 4 L1-L3; RFE21, 9 L1-L2; RFE28, 1 2113; REE37, 1: U3; RFE48,°39 L1-L3; RFE54, 29 L2-L4 (incl. prepupae) (all in PZ). Primorje: Lazovskii Reservation, Lazovka River near vill. Lazo, leg. Vshivkova, 05.07.1978, 4L (IBP>PZ); 08:07:1979, 2P, IL (IBP>PZ). Lazovsky Reserve, Kievka River, 04.08.1985, leg. Gostyukhina, 5L, 2P; 13.06.1998, leg. Arefina, 72L, 2P (IBSSAN). Ussuri River Basin, Zubkin Stream, 27.07.1992, leg. Tiunova, IL (IBSSAN). Ussuri River near mouth of Izvilinka River, 08.08.1979, leg. Levanidova, 10L (IBSSAN). Ussuri-Bolshaya Ussurka River Basin, Perevalnaya Stream, 06.08.1997, leg. Tiunova, 4L, 2P (IBSSAN). 351 Yedinka River (middle part), 14.08.1986, leg. Korionov, 3L, 7P (IBSSAN). Narva River (lower part), 02.08.1987, leg. Teslenko, 4L, 3P; 14.08.1994, leg. Arefina, 2L, 9P; Sikhote-Alin Reserve, Bystraya Stream, 02.08.1997, leg. Tiunova, 7L, 10P (IBSSAN). - Khabarovsk Territory: Iski River, 12.08.1959, leg. Le- vanidova, 5P, 3L (IBSSAN). Bikin River Basin near mouth of Omorochka Stream, 30.07.1995, leg. Tiunova, Y (IBSSAN). Bikin River Basin, Klyuchevaya Stream, 28.07.1995, leg. Tiunova, IL (IBSSAN). Bikin River Basin, Zeva River (upper part), 17.07.1995, leg. Ti- unova, IL (IBSSAN). Bikin River Basin, Zeva River, near Beriozovaya Stream, 24.07.1995, leg. Tiunova, 10L, 8P (IBSSAN). Bolshekhekhtsirsky Reserve, Ussuri River Basin, Bychikha Stream (15.5° C), 22.06.1958, leg. Levanidova, 4L, 1P (2) (IBSSAN) (all in PZ). A typical Blepharicera. Adults available only as pharate specimens; dimensions, pigmentation and details of wing therefore unknown. Fig. 15: Blepharicera parva sp. n., adults. a, b, male genitalia, dorsal view, tergite and cerci shown separately; c, left dististyle, dorsal; d, head of female; e, head of male; f, Ble- pharicera japonica, part of male genitalia, dorsal (from ZWICK 1990). Adult. Male. Widely separate almost simple eyes, re- duced upper portion forms narrow section at mesal rear ¡99 Nn 159) margin of eye, behind ocelli. Rostrum shorter than head height, labella projecting slightly beyond labrum. No mandibles, hypopharynx, lacinia; palpal segments short except whip-like 5 segment. No facial pilosity. An- tenna slender, most segments more than twice as long as wide. Coxae normal, simple. No tibial spurs. Tarsal _ claw falcate, with some hairs on sides near base. Hypopygium slender, inner sclerites inverted as typical of B. fasciata-group. Tegmen lightly sclerotized, apex of common spade-shape. Inner dististyle divided into blade-like median and short finger-shaped lateral por- tions. Inner edge of outer dististyle with some basal se- tae, distally angularly bent mediad. Dorsal and ventral distal lobes deeply divided, dorsal lobe a bit angled and much shorter than the more regularly curved soft ventral lobe with medially projecting apex. Cercus narrowly rounded, no angles in contour. d Fig. 16: Blepharicera parva sp. n. (a-c) and B. japonica (d, e). a, d, terminal body division of last instar larva, dorsal; b, e, de- tails of integument on some middle abdominal segment; c, granula on abdominal segment of pupa. Figures a and d and c- e are to the same scales, respectively. Female. A very narrow crest between eyes runs to ocel- lar triangle which is not raised. Bare strip separates un- equal portions of widely separate eyes. Upper portion in frontal view slightly smaller than lower, convex, very large bulging ommatidia in centre, lateral ones much smaller. Antenna short, of 15 segments, acutely pointed, setiform, each flagellar segment with single longer seta on outside, resulting in sparse setal fringe. Distinct fa- cial hair patch between base of antenna and base of ros- trum. Rostrum about as high as head. Mandibles serrate, long, apex curved mediad. Hypopharynx serrate, hidden behind labrum which is little longer than labella. Lacinia distinct. Palpus segments short, especially 5". Mesocoxa with usual hairy appendage. Hind tibia with two small unequal spurs, base of hind tarsus with some procumbent long black setae. Tarsal claws as in male. Genitalia not distinctive, three spherical receptacles with sclerotized duct whose diameter widens abruptly short distance from receptacle. Bonner zoologische Beitráge 53 (2004) Pupa. Shape and respiratory organs typical of genus, 3.8-4.8 mm long, 2.0-2.5 mm wide. Integument glossy, dorsal granules on metathorax and abdomen tiny, each with central dome of closely packed hair-like pegs (Fig. 16c). Anterodorsally on abdominal tergites granules ag- gregate and form slightly raised warts. Anterior and posterior lamellae of gills of same width and almost twice as wide as lamellae 2 and 3 between them. Brown, also all four gill lamellae of same colour. Larva. Last instar larvae up to 6.5 mm long, typical of subgroup. Cephalic sclerites with usual mottling but each dark spot on raised area of integument, which is hemi- spherical on medial frontal sclerite, but flat and obliquely rising from low front to high posterior edge on lateral frontal sclerites. Antenna two-segmented, segments sepa- rated by pale section of same length as basal segment, lat- ter distinctly shorter than apical segment. Each abdominal tergite may have anteromedian wart, surrounding in- tegument and intercalary segments darker than rest of surface; although some specimens lacking warts alto- gether. Surface finely microreticulate, with numerous tiny insertion points; setae reduced to tiny acutely conical pegs and not easily visible. Soft pale hairs on sides of body. Ventral face of body with distinct but pale asper- ities between sucker and base of pseudopods. Seven gill filaments in each tuft. Body divisions medially brownish, anterior part of each division darker than posterior, pale mark at paramedian muscle attachments; neck—pieces and lateral portions of divisions pale. Last body division with truncate sclerotized blackish distal edge. Lateral lobes of segment bluntly triangular, their posterior contour meeting body contour at right angles. Last pseudopod reduced to sclerotized area around setal insertion, medial edge of sclerite often downcurved to form small grapple. Larval instars 3, 2 and |, respectively, with five, three and no gill filaments, less distinctly patterned, dull grey, lobes of last body division less distinct. Antenna one— segmented in instars | and 2, with very short dark ring representing basal segment only in instar 3. Cuticular ornaments of first instar larva as described for B. fas- ciata (ZWICK 1980), that is, only single prothoracic transverse cuticular crest present on thorax. Variation. The only variation concerns the expression of tergal warts and dorsal pilosity in larvae. In some populations, especially at the type locality, warts are shallow and inconspicuous; in most others, particularly in the Kievka river, warts are steeply rising and high. Similar variation is known in other species, for example B. fasciata. While some specimens appear almost bare, others are fairly pilose. However, this may depend less on absence or presence of hairs than on extent of bacte- rial and detrital coating which greatly affects visibility of hairs. Peter Zwick & Tatyana AREFINA: Blephariceridae of the Russian Far East Distinction from related species. On the Asian mainland occur only species of the fasciata-subgroup, which were all redescribed by ZWICK (1990). However, B. japonica Kitakami, 1931 is the only species of which all stages and both sexes are known (KITAKAMI 1931, 1950). B. yamasakii (Kitakami, 1950), from Manchuria: Adults are very similar to B. parva, but both sexes bear a hairy appendage on mesocoxa (in all other species, present only in females). The upper portion of the female eyes are “nearly flat”, and the sides of the lower division are “nearly vertical” whereas in B. parva both divisions are distinctly convex. The pupa is not recognizably differ- ent. The larva differs by the dense pale pilosity and lack of integumental warts although the shape of the caudal body division is similar to B. parva. B. japonica: Adults resemble B. parva, but the male cerci are slightly pointed, the upper lobe of the outer dististyle is smaller than the lower (opposite in B. parva), and the lobes are shallowly separate. Females lack a hairy mesocoxal appendage but the trochanter is covered in dense black hairs. Eyes are simple, with no facial hair patches, and the rostrum lacks mandibles and hypopharynx. The width of the sclerotized receptacular duct changes gradually, not abruptly. Blepharicera ja- ponica females have very slender, thin, skinny tarsi with strongly oblique articulations and heavy strongly curved claws; these details were previously overlooked. A simi- lar character syndrome was recently noticed to be com- mon in another tribe, Apistomyiini (ZWICK 1998). Pu- pae of B. parva and B. japonica can presently not be distinguished. Long lateral lobes of segment seven of B. japonica larva are directed obliquely backward (Fig. 16d). The surface bears pale, club-shaped flattened hairs among microreticulation (Fig. 16e) and no integumental warts. Three dark bands run over entire length of body. Cephalic sclerites are smooth, not sculptured as in B. parva. Instar 3 of B. japonica has already two distinct antennal segments, instar 2 possesses a small dark basal ring; otherwise not distinct from B. parva. B. yankovskyi (Alexander, 1953): The single male from North Korea has very large eyes: the upper section 1s small but much less reduced than in B. japonica, and hence also different from B. parva. B. dimorphops (Alexander, 1953) (from China, Fukien): Outer male gonostyle is very deeply divided, with both lobes slender, and this form is distinctly different from B. parva (figs. 8a-c of ZWICK 1990). 4.4.2. Blepharicera sp. Material. Mongolia, Selenge, Mandal, Khonin Nuga [49%05'260N, 107°17'440E], Eroo River, 21.V1.2003, 1L3, 2L2, leg. P. Surenkhorloo (PZ). 353 The larvae resemble.B. parva, but we are not certain of their conspecificity. 4.5. Genus Neohapalothrix Kitakami, 1938 Neohapalothrix Kitakami, 1938. Type species: Néoha- palothrix kanii Kitakami, 1938, by original designation (KITAKAMI 1938: 341). Asiobia Brodsky, 1954. Type species: Asiobia acan- thonympha_ Brodsky, 1954, by original designation (BRODSKY 1954: 245). Synonymy of ZWICK, 1990: 253. The phylogenetic status of the closely related genera Neohapalothrix and Hapalothrix (Europe; ZWICK, 1997b) is not firmly established because of conflicting characters. A dorsal larval feeler with geminate setae occurs only in Blepharicerini. ZWICK (1992) therefore included both genera in this tribe, assuming conver- gence for reduced wing veins and habitus characters. Most authors, however, regarded both genera as Pal- tostomatini because of reduced wing venation, curved front femora, short front and middle and very long hind legs. The labial brush here described for Neohapalothrix has so far only been observed in Paltostomatini (ZWICK 1998). In view of this last character we now assume that the genus actually belongs in that tribe. If so, the dorsal feeler must have evolved independently from Ble- pharicerini. Indeed, the feeler originates through split- ting of the pseudopod, and is positioned in front of the pseudopod, not above it as is typical of Blepharicerini. Both traits are unique. Neohapalothrix is endemic to Asia. The genus includes the type-species from Honshu and the widespread N. manschukuensis (Mannheims). 4.5.1. Neohapalothrix manschukuensis (Mannheims), 1938 (Figs. 17-20) Curupira manschukuensis Mannheims, 1938. — Type- locality: Weischache, Manschukuo [China, Manchuria] (MANNHEIMS 1938: 329). Asiobia acanthonympha Brodsky, 1954. — Type- localities: Russia: r. Angara, Padun; S-Primorje: r. Bi- kin; Kazakhstan: Altai, r. Bija (BRODSKY 1954: 250). — Synonymy of Zwick, 1990: 253, Neohapalothrix shirozui Saigusa, 1973: 226. — Type lo- cality: Japan, Hokkaido, Teshio, Piuka (SAIGUSA 1973: 226). — Synonymy of ZWICK, 1997b. Curupira manschukuensis was described from a single female and tentatively placed in the neotropical genus Curupira. In the same year, KITAKAMI erected the ge- nus Neohapalothrix for his new species, kanii, from Ja- pan. Subsequently, KITAKAMI anticipated and estab- lished, respectively, the correct generic combination of the mainland species (KITAKAMI 194la, 1951). How- ever, it was later described and named two more times. BRODSKY (1954) presented all life stages. The amazing male tarsal structures were again illustrated by SAIGUSA (1973) and ZWICK (1997b) who also described changes in pseudopod structure between larval instars, compar- ing them with Hapalothrix. Neohapalothrix manschukuensis was known from the Altai (BRODSKY 1954) and the Russian Far East; the new Mongolian record is therefore not surprising. The new record from Sakhalin closes a gap in the distribu- tion, because a single male was taken on Hokkaido (SAIGUSA 1973). Neohapalothrix manschukuensis 1s common in the Russian Far East, conspecificity of all stages is definitely established through the metamorpho- type-approach. We describe previously unknown details and present some ecological observations. 500 pm 300 um Fig. 17: Neohapalothrix manschukuensis. Female maxillo- labium, dorsal view, and enlarged detail of left half of figure. Muscles diagrammatic. Material (all in PZ). RFEO9, 21P, 12L4; RFE14, 33L3, 64L2, 6L1; RFEI5, 100L4, 36L3, 5L2, 3L1; RFEI6, 43, 189, 20P, SE, 18L4; RFE18, 43P, 8L4, 113; RFE19, 12L3; RFE21, 28L4, 84L3, 412; RFE23, 61P, 33L4, 3L3; RFE28, 24L4, 14L3, 1L2; RFE30, 88P, 13L4; RFE33, 341P, 19L4, 2L3; RFE35, 7P, 29L4, 1L2; RFE38, 23L4, 3L3; RFE39, 37P, 2114; RFE47, 39L4, 2L3; RFE48, 3P, 50L4, 7L3, 1L1; RFES2, IP; RFES3, 24; RFES6, 9P, 5L4, 1L3. Russian Far East, Sakhalin: Tym River at Ado-Tymovo Village, 10.07.1985, E. Ma- karchenko, 1 L4 (TA). Additional material: Mongolia, Selenge: Mandal, Kho- nin Nuga [49%05'260N, 107°17'440E], Eroo River, 21.V1.2003, 28P, 43L4, 1L3; 25.V1.2003, 359; Eroo River [49°05'N, 107°17'E], 26.V1.2003, 29; 13.V11.2003, 2%; Eroo hot spring, Irchleh River, [49°05'N, 107°17'E], 28.V1.2003, 16L4, 8P; Mandal, Ataa River, 18.V11.2003, 1L3; Mandal, Bar-Chuluut River Bonner zoologische Beiträge 53 (2004) [48°58'638N, 106°57'013E], 13.V11.2003, 22; Mandal, Bar-Chuluut River [49%02'784N, 106%58'669E], 09.V11.2003, 1L4 (all leg. P. Surenkhorloo; in PZ). Adults. Both sexes of similar habitus. Female front fe- mur slightly less sinuous than male. Female holoptic, eye little smaller than in male but also sharply divided into upper and lower portions. Rostra differ little, except longer maxillo-labium with base bent back distinctly only in females. Labella alike in both sexes. Soft scaly inner face forms anterior finger-shaped fold and larger posterior lobe with flat smooth medial face. Delicate brush at base of posterior lobe formed by slender cone with hair-like outgrowths of cuticle. Female genitalia remarkable for short palpus-like apex of cercus not extending backward beyond general con- tour line. Oviscapt with truncate deeply divided but nar- rowly adjacent lobes. Dorsal face of each lobe without cover, side forming lateral wall ending in small dorsal angle. Sclerotized base of oviscapt truncate with promi- nent corners, almost bifid. Median of three pear-shaped black receptacles slightly smaller, all with short straight stalks. Fig. 18: Neohapalothrix manschukuensis, female genitalia. a, sternite 8 and oviscapt; muscles diagrammatic, as seen by transparency; left half of figure shows oviscapt in ventral, right half in dorsal view. b, sclerites from female genital cav- ity, with attached receptacles. Larval instars. First instar larva (Figs. 19a-c) of normal habitus, dorsal side convex, rounded. Anal body divi- sion semicircular, no lobes; pseudopod seven repre- sented only by tiny wart with long seta. Long, flattened cuticular structures resembling blunt hairs stand in transverse rows across body. One long row per thoracic segment. Each abdominal segment with short anterior plus long posterior row. Slender hair-like tergal and subtergal sensilla standing in posterior rows recognized by insertion points only at magnification over 200. Pseudopods in dorsal view largely hidden under body. Peter ZWICK & Tatyana AREFINA: Blephariceridae of the Russian Far East Base of each pseudopod with group of three spine-like cuticular outgrowths and a large seta (tergopleural and/or pre-dorsopseudopodal sensilla, HOGUE 1978), geminate setae (dorsopseudopodal setae of HOGUE 1978) typical of first instar pseudopods close to apex. Colourless delicate cuticular tube everted from tip of pseudopod with about 10 tiny sharp hooks at end. An- tenna l-segmented. No gills. Lateral rectal tubules very long, median ones minute. Second instar larva with hard central cone on each ab- dominal segment. Integument with long curved hairs, some blunt, others apically thinned and finely pointed. On sides of body also short transparent spatulate and curved setae. Pseudopods forked (Fig. 19d), long poste- rior branch with rough ventral sole and three long dorsal setae. Short anterior branch with two or three long hairs in addition to geminate setae; ventrally smooth. Antenna 2-segmented. Single gill filaments, rectal tubules un- changed. ioe ER (ay WA 11722 Wal Sl Wak. Shee \ NOE E EOS L1 INN SAS \ LD pr (> IN wm >> c INS RANG 2 © Sr HEIN hel PLS pero UNA Fig. 19: Neohapalothrix manschukuensis. a, first instar larva (cephalic division plus abdominal segment 3; abdominal seg- ments 5 and following); b, detail of dorsal armature, not to scale. Left side of abdominal segment three and pseudopod: c, first instar, posterior view (from a slide mount); d, e, 3" and 4" instar, respectively, dorsal views. Same scale for c and d. Branches of pseudopod deeply separate in 3" (Fig. 19e) instar larva, completely divided in 4" instar. Hair cover denser, four and seven gill filaments, respectively. Ab- dominal segments 1-6 with strong dark dorsal spine on 355 raised middle portion; body cross-section resembies equilateral triangle (lateral view in BRODSKY 1954 is far too flat!). Dorsal side sometimes (3" instar) or normally (4th instar) patterned in dark brown and yellowish. Ex- tent of pale pigment varies from only abdominal seg- ments one and two and sides of five and six pale to mainly light specimens with dark cephalic sclerites, spines and pseudopods, and brown segments three and 4. Ecological notes. Scientists of IBSSAN repeatedly picked numbers of adults from flowers of Angelica saxatile (Turez.) (Daucaceae) and Filipendula palmata (Pall.) (Rosaceae). During our field work, adults were observed flying over the river banks, just before com- plete darkness. Dying and dead spent females were found between pebbles in stagnant water along the Khor River below Sukpai. Females seemed to have oviposited in areas where the river was receding. Individual last in- star larvae and pupae observed on pebbles in the same area were alive but thickly covered with mud. Normally, larvae and pupae were found completely submerged, in strong current. There is an obvious preference for large watercourses and potamal habitats. Population structure varied much between samples taken almost simultaneously. At sev- eral localities, water temperature was recorded at 20 min intervals (Gemini Tinytag datalogger ®) over night, between 19:00 and 8:00 hrs. The relationship between nocturnal minimum temperature and predominant de- velopmental instar is shown in Figure 20; accelerated development in warm water is obvious. The sample from Komissarovka River is an exception but may be too small for safe judgement. Ex a Ss Bikin (101 n P 7 Komissarovka (15) ] e ssuri E La 4 Sukpah (61) = Tahalo (37) E Perevalnaja (139) L3 | 8 Malinovka (116) L2 + 1 ; r 10 12 14 16 18 20 Nocturnal Minimum Temperature [°C] Fig. 20: Neohapalothrix manschukuensis, dominant instars (larval instars L2-L4, P = pupae, E = exuviae) in samples from rivers with different nocturnal minimum temperature, 11-24 June, 1998. Figures in parentheses are numbers of specimens in the sample. Acknowledgements. We acknowledge the loan of impor- tant historical specimens and thank E.P. Nartshuk (Zoo- logical Institute, Academy of Sciences, St. Petersburg), H.Ulrich and B. Sinclair (Museum Alexander Koenig, Bonn), W. Mathis (United States National Museum, Wash- (98) N nN ington) and G.W. Courtney (Ames, Iowa) for their cour- tesy. G.W. Courtney and B. R. Stuckenberg (Pietermaritz- burg) reviewed the final draft of the present study. K. Ta- nida (Osaka) supplied and partly translated a rare study by S. Kitakami. K.N. Satake kindly provided information on parts of the Kitakami collection in the Otsu Hydrobiologi- cal Station, Kyoto University. Prof. Y.J. Bae (Seoul) is sin- cerely thanked for providing transcriptions of Korean lo- calities originally published in Japanese. Interest in the Far East Blephariceridae arose during study of many occasional samples of Russian Blephariceridae, collected by I.M. Levanidova and other workers at the In- stitute of Biology and Soil Science, Russian Academy of Sciences, Vladivostok, which I.M. Levanidova made avail- able to P. Zwick. We cordially thank our friends and col- leagues Tatyana and Michail Tiunov, Valentina Teslenko and Valera Lyubaretz for help and comradeship during successful field work in the Russian Far East, in June 1998, and for the gift of samples collected at other occasions. Our thanks are also extended to Oleg Timoshkin (Institute of Limnology, Russian Academy of Sciences, Irkutsk) and Vladimir Samokhvalov (Institute of Biological Problems of the North, Russian Academy of Sciences, Magadan) for collecting material. Travel grant Nr. 436 RUS 18/28/97 by Deutsche Forschungsgemeinschaft to P. Zwick is gratefully acknowledged. We also thank the Max-Planck-Society for a grant allowing T. Arefina to participate in the work at Schlitz. The gift of a rich collection of Japanese net-winged midges by S. Uchida (Toyota) to P. Zwick is very much appreci- ated. Important comparative material from Mongolia was lent or given to us, respectively, by W. Horn and M. Paul, Sachsische Akademie der Wissenschaften zu Leipzig, Ar- beitsgruppe Limnologie, Lengfeld, Germany, and P. Surenkhorloo, Ulan Bataar; V. Devyatkov (Ust-Kameno- gorsk) kindly gave us a collection from Kazakhstan. We sincerely thank them all. REFERENCES AREFINA, T. I. 1995. A small collection of net-winged midges (Diptera, Blephariceridae) from the Russian Far East. Report of the Studies of the Structure and Function of River Ecosystems of the Far East 3: 3-4. BRODSKY, K. 1954. [Blepharoceridae (Diptera) from Altai and southern Primorye]. — Trudy Zoologitsheskogo In- stituta Akademii Nauk SSSR 15: 229-256. [In Rus- sian]. BRODSKY, K. 1972. New species and changes in the status of previously described species of Middle Asiatic Ble- pharoceridae (Diptera). Entomologitsheskoe Oboz- renie 51(3): 637-645. [In Russian]. [English transla- tion: Entomological Review, Washington, 51: 384- 389]. BRODSKY, K. A. 1976. Mountain Torrent of Tian-Shan Ecological-faunistical review. Akademia Nauk SSSR, Zoological Institute, Leningrad, 244 pp. [In Russian]. [English translation (1980): Mountain torrent of the Tien Shan: A faunistic-ecology essay. W. Junk: The Hague. 311 pp.]. Bonner zoologische Beitráge 53 (2004) COQUILLETT, D. W. 1910. The type-species of the North American genera of Diptera. Proceedings of the United States National Museum 37: 499-647. HOGUE, C. L. 1973. The net-winged midges or Ble- phariceridae of California. Bulletin of the California Insect Survey 15: 1-83, HOGUE, C. L. 1978. The net-winged midges of eastern North America, with notes on new taxonomic charac- ters in the family Blephariceridae (Diptera). Contribu- tions in Science of the Los Angeles County Museum of Natural History 291: 1-41. HOGUE, C. L. 1982. Revised status of net-winged midges of the genus Bibiocephala in North America based on a study of quantitative variation in the males (Diptera: Blephariceridae). Contributions in Science of the Los Angeles County Museum of Natural History 338: 1-16. HOGUE, C. L. 1987. Blephariceridae. In: GRIFFITHS, G. C. D. (ed.). Flies of the Nearctic Region. E. Schweizer- bart'sche Verlagsbuchhandlung, Stuttgart 2(4): 1-172. KELLOGG, V. L. 1903. The net-winged midges (Blepharo- ceridae) of North America. Proceedings of the Cali- fornia Academy of Sciences, Series 3, 3(6): 187-233, pls. XVHI-XXIL. KITAKAMI, S. 1931. The Blepharoceridae of Japan. Mem- oirs of the College of Science, Kyoto Imperial Univer- sity, Series B 6(2): 53-108, pls. VII-X VII. KITAKAMI, S. 1937. Supplementary notes on the Blepharo- ceridae of Japan. Memoirs of the College of Science, Kyoto Imperial University, Series B 12(2): 115-136, pls. FIL KITAKAMI, S. 1938. A new genus and species of Blepharo- ceridae from Japan. Memoirs of the College of Sci- ence, Kyoto Imperial University, Series B 14(2): 341- 352, pls. XXI-XXIII. KITAKAMI, S. 1941a. [The Blepharoceridae from Manchu- ria]. Dobutsugaku Zasshi [Zoological Magazine] 53: 89-90. [In Japanese]. KITAKAMI, S. 1941b. On the Blepharoceridae of Formosa, with a note on Apistomyia uenoi (Kitakami). Memoirs of the College of Science, Kyoto Imperial University, Series B 16(1): 59-74, pls. I-IL. KITAKAMI, S. 1950. The revision of the Blepharoceridae of Japan and adjacent territories. The Journal of Kuma- moto Women's University 2: 15-80, pls. 1-5. MANNHEIMS, B. J. 1938. Uber das Vorkommen der Gat- tung Curupira in Manschukuo nebst Beschreibung der Entwicklungsstadien zweier neuer Blepharoceriden aus Anatolien und Súd-Chile (Diptera: Blepharoceri- dae). Arbeiten über morphologische und taxonomische Entomologie aus Berlin-Dahlem 5(4): 328-332. MACQUART, J. M. 1843. Description d'un nouveau genre d'insectes dipteres. Annales de la Société Entomologi- que de France, 2éme ser. 1: 59-63, fig. I on pl. 3. MATSUMURA, S. 1916. Shin-Nippon Senchu-Zukai [New thousand insects of Japan, Additamenta]. Tokyo. Vol. 2 [Blephariceridae on pp. 413-414, pl. XXIV, fig. 7]. MATSUMURA, S. 1931. Six thousand illustrated insects of Japan-Empire. Tokyo. 1-111, 23, 1497 + 191, 2, 6, pls. 1-10. [Blephariceridae on p. 407. In Japanese]. NARTSHUK, E. P. 1999. Blephariceridae Setshatokrylie komari. Pp. 70-80, pls. 39-42 on pp. 448-455 in: TSA- LOLIKHIN, S.J. (ed.) Key to freshwater invertebrates of Russia and adjacent lands, Vol. 4, 1000 pp., St. Peter- burg. [In Russian]. Peter ZWICK & Tatyana AREFINA: Blephariceridae of the Russian Far East 357 OSTEN-SACKEN, C. R. 1874. Report on the Diptera col- lected by Lieut. W. L. Carpenter in Colorado during the summer of 1873. Annual Report of the United States Geological and Geographical Survey of the Ter- ritories for 1873 7: 561-566. SAIGUSA, T. 1973. A new species of the genus Neohapa- lothrix from Japan (Diptera: Blephariceridae). Sie- boldia 4(3): 225-235, pls. 16-17. STUCKENBERG, B. R. 1958. Taxonomic and morphological studies on the genus Paulianina Alexander (Diptera: Blepharoceridae). Mémoires de l'Institut Scientifique de Madagascar, Série E 10: 97-198, pls. I-III. TONNOIR, A. L. 1924. Les Blepharoceridae de la Tasmanie. Annales de Biologie Lacustre 13(1-2): 5-67. VON ROEDER, V. 1890. Zwei neue nordamerikanische Dip- teren. Wiener Entomologische Zeitung 9(8): 230-232. ZASYPKINA I. A. & RYABUKHIN, A. S. 2001. Amphibiotic Insects of the Northeast of Asia. Pensoft and Backhuys Publishers, Sofia, Moscow, Leiden; VHI + 184 pp.; Blephariceridae on p. 65. Zwick, P. 1977. Australian Blephariceridae (Diptera). Australian Journal of Zoology, Supplementary Series 46: 1-121. ZWICK, P. 1980. The net-winged midges of Italy and Cor- sica (Diptera: Blephariceridae). Aquatic Insects 2: 33-61. ZWICK, P. 1990. Systematic Notes on Holarctic Ble- phariceridae (Diptera). Bonner Zoologische Beiträge At 231-157. ZWICK, P. 1992. Family Blephariceridae. Pp. 39-54 in: Soos, A., PAPP, L. &. OOSTERBROEK, P. (eds) Cata- logue of the Palaearctic Diptera, 1 (Trichoceridae — Nymphomyiidae). ZWICK, P. 1997a. A redescription of Philorus novem Kaul, 1971, and a new synonymy in net-winged midges (Diptera, Blephariceridae). Mitteilungen der Schwei- zerischen Entomologischen Gesellschaft 70: 295-298. ZWICK, P. 1997b. Synonymies in the Genus Neohapa- lothrix (Diptera: Blephariceridae). Aquatic Insects 19: 9-13. ‘ ZWICK, P. 1998. The Australian Net-Winged Midges of the Tribe Apistomyiini (Diptera: Blephariceridae). Austra- lian Journal of Entomology 37: 289-311. ZWICK, P. & Zwick, A. 1998. Net Winged Midges (Dip- tera: Blephariceridae) from the Philippines, mainly from Palawan. Aquatic Insects 20: 1-27. Authors’ addresses: Peter ZWICK (corresponding au- thor): Limnologische Fluss-Station Schlitz, Postfach 260, D-36105 Schlitz, Germany, e-mail: pzwick@mpil- schlitz.mpg.de; Tatyana AREFINA: Institute of Biology and Soil Science, Russian Academy of Sciences, Far Eastern Branch, Vladivostok 690022, Russia, e-mail: arefina(wibss.dvo.ru. 22.06.2004 07.09.2004 B. J. Sinclair Received: Accepted: Corresponding editor: re 4 Ms ' 4 7 f= 7 zer a . E i ‘ , B 0 0 5 i; ‘ j . ot j j 4 ; o — i a . o : + I o Ks 7 vy ' A . A ‘ o o Ú o , if . y ze: dr o . . = wa Vion E 22) v y o 1 j \ A Ú 7 a o + a ? pl ot hi bd . y o a " 7 My hay Y . i tot Oo F Y . ae u u Bi = ot Ve 64 OF ” Y 0 UA o A \ ö ‘ bu u | y , . 14: . “ot A \ i \ i 1 ‘ J y 0 & i A 1 Hl oF . B a 0 1 7 ‘ ' i ' y ñ R “ eS | ren 0 y o 5 E ve x Y 7 T v . o = * . ] A , 5 B gr Ü 7 ' ry 7 A 7 ’ FIR b + 1 o MS dl i . of 5 y a \ hoo , “ Ñ . En . . i i . Y TAO oS s ' 1 \ fl ‘ Va 7 " y , i " 7 y \ . , a # a u ' ' - Y 7 ‘ A ' A R TR ñ h 7 e i ü 4 e . A ” ' ' Ñ 5 ' den ' Py. a \ er B in : hea ee o i . ns x . . 7 e ES iS g 7, : AF: B . om ? Da) ‘ 1 \ AS : . 7 ' ‘ A 4 , o 7 u \ ' o 7 fi . . . ‘ ñ . ; so. A ' . 7 va MO PS e Ne ANI A A y un, u Pr . O vty 1) . . e ON 4 o a i ’ hs \ y i 7 F wi i m . . abs . | i ' B B ' 7 as) ‘ ‘ o ' * . . 7 . Zu > 1 ' cl y \ : 2 . . , \= ' ñ . 4 B ñ ' ‘ : 7 ' ' A 7 » i 4 "| . f “ae ' . 1 i ; + 7 ' i \ ' 7 4 ' ñ ' ' ve . = ; i = \ 7 =i ' A = =!) = 5 | ON " i ] . ñ : : . a : a 7 ‘ 7 u 9 Er i . ‘ e ö , y toh ‘ u 6 . \ . = y I j rn, 7 i q 1 . 14 ur A “ u o " [vs i ' a . en \ . 7 ‘ i al y 7 U) 5 o A o . 6 ' ti 7 u 7 uy Editorial Dear Reader, In August 2005, the Federal Secretary of Culture — from which we receive 25% of our funds - imposed on us a fi- nancial cut of € 109,000. As a consequence, we could not afford to continue publishing issues of Bonner zoolo- gische Beitráge after the release of the double issue 53(1/2) in June. We regret that we had to ask authors and read- ers for even more patience than usual. Our aim is to catch up to the actual yearly schedule as soon as possible. Last year, we had sufficient manuscripts, but suffered from a crucial lack of financial means. There are good chances, that we can reach our goal in 2006. Due to an unfortunate coincidence of technical problems and personal misunderstandings, some of the colour illus- trations in the Clas-M.-Naumann-memorial issue have been printed in discontenting quality. We, editors and print- ers, apologise for any trouble and disappointments caused through this failure. In attempting to make amends, all colour illustrations are herein reprinted and placed at the end of this double issue, after p. 358. The authors will be provided with colour-corrected high resolution PDF files of their contributions. Sincerely Michael Schmitt 14 Bonner zoologische Beitráge 53 (2004) durch ihm Schloss Greiffenstein und die umliegenden Landereien zugefallen waren; damals nahm er auch den Namen RUSWORM an. Sein Vermögen ermöglichte es ihm, sich nun ganz seinen Interessen zu widmen. Ein Schliisselerlebnis war die Begegnung mit den ,,Mikro- skopischen Augen- und Gemütsergötzungen“ von Mar- tin Frobenius LEDERMULLER und schlieBlich mit dem Autor dieses bis heute berühmten Werkes selbst. W. F. VON GLEICHEN besuchte M. F. LEDERMULLER 1760 in Erlangen und bestellte bei ihm ein Mikroskop, und Mik- roskope sollten fortan auch zu seinen wichtigsten Ar- beitsbehelfen gehóren. Im Jahre 1762 hielt sich LEDER- MULLER etwa vier Wochen bei VON GLEICHEN auf Schloß Greiffenstein auf, die beiden Mikroskopiker verbrachten offenbar eine glückliche Zeit, in der sie sich in gemeinsam erlebter Freude und Begeisterung ganz dem Mikroskopieren hingaben. Schon vor seiner Be- gegnung mit M. F. LEDERMÜLLER, unmittelbar nachdem er sich auf sein Schloss zurückgezogen hatte, hatte W. F. VON GLEICHEN eine rege Publikationstätigkeit be- gonnen, wobei er sich mit ganz unterschiedlichen na- turwissenschaftlichen, landwirtschaftlichen und techni- schen Fragen befasste. Dass besonders diese frühen Publikationen manchmal eine genügend kritische Be- trachtungsweise vermissen lassen, darf nicht unerwähnt bleiben. Ab 1760 wurde W. F. VON GLEICHEN ein Mikroskopiker ersten Ranges, ausgestattet mit einer ausgeprägten Beobachtungsgabe und einer ebenso aus- geprägten Lust, das Gesehene zu interpretieren, Schlüs- se zu ziehen und zu veröffentlichen. So entstanden zahl- reiche weitere Publikationen über sehr unterschiedliche naturwissenschaftliche Themen (Bibliographie: FIKEN- SCHER 1801). W. F. VON GLEICHEN starb infolge eines Schlaganfalls in seinem 67. Lebensjahr am 16. (oder 18.) Juni 1783 auf Schloss Greiffenstein. Die Bedeutung W. F. VON GLEICHENS als Naturforscher wurde schon zu seinen Lebzeiten erkannt, und schon unmittelbar nach seinem Tod erschienen Darstellungen seines Lebenswegs: WEIKARD 1783; ANONYMUS 1784; FIKENSCHER 1801. Von den späteren biographischen Publikationen über W. F. VON GLEICHEN sind vor allem die Arbeiten von WILLNAU (1926), WEIDNER (1980), GEUS (1989, 1990) und DETTNER (1997) zu erwähnen. W. F. VON GLEICHEN war nicht nur ein begabter Expe- rimentator, sondern ein ebenso talentierter Zeichner, der seine Publikationen mit vorzüglichen Abbildungen aus- stattete. Seine berühmteste und durch hervorragend ko- lorierte Kupfertafeln ausgestattete entomologische Ar- beit ist ohne Zweifel jene über die „Geschichte der gemeinen Stubenfliege“ (VON GLEICHEN 1764), in der er überdies Beobachtungsfehler von M. F. LEDER- MÜLLER aufzeigte, was bedauerlicherweise zu einer Ent- zweiung der beiden führte. Entomologische Themen werden auch in mehreren an- deren Publikationen VON GLEICHENs behandelt. In zwei Werken setzt sich VON GLEICHEN mit Neuropteren aus- einander, nämlich in dem 1764 erschienene Band über „Das Neueste aus dem Reiche der Pflanzen ...“ (Abb. 1) und in seinem 1770 publizierten „Versuch einer Ge- schichte der Blatläuse und Blatlausfreßer des Ulmen- baums ...“ (Abb. 9). Das 1764 veröffentlichte Werk ist — wie man schon dem langen Titel (,,... und einem Anhang vermischter Beobachtungen ...*) entnehmen kann — so etwas wie ein Sammelband über verschiedene Themen. Er enthält auch zwei neuropterologische Artikel, von denen der eine („Der Blatlausfreßer und seine Eier an einem Hanfblat“) den präimaginalen Stadien einer Chrysopiden-Spezies, der andere („Die Hofdame“) einer im Winter gefundenen Imago einer Florfliege gewidmet ist. Das Neuefe aus bem Keigeden Viene, oder Mitroffopifóe Snserfuhunge und © Beodadkungen geheimen Zeugungötheile der "Manzen in ihren Sit der in denfelber befindlichen Sifters nebf i einigen Nerfuden von dem Keim, und einem Anhang i vermifiter Srobadtungen, aed mit Garben nad der Natur vorgefkellet von Wilhelm Fricderid) Freiheren von Gleiden, genannt Mupworm, Herrn auf Breiffenftein und Bonnland, Groß. Ereuz des Brandenburgifchen rohten Adler» Ordens und Hodfürftt, Brandenburgifc, Eulmbadhifhen Geheimen Naht, Herousgese bens verlegt mit den ndftigen In Kupfer dime und [Cuminirten Abblldungen verfegen von Sobann Chriffoph Keller, Maler in Nürnberg Geordt, bd Ebriflan de Launoy feel. Erbm, 1764, a] Abb. 1: W. F. VON GLEICHEN (1764), Titelblatt des Werks, das die Arbeiten über den „Blatlausfreßer ..“ (1764a) und „Die Hofdame“ (1764b) enthält (Bibl. H. & U. ASPOCK) In der 1770 veröffentlichten umfangreichen Arbeit wird die Biologie einer Hemerobiiden-Spezies ausführlich dargestellt. VON GLEICHEN hat sich wenig um die kor- Horst AsPöck & Ulrike Aspöck: Neuropterologische Beiträge in den Werken von Wilhelm Friedrich VON GLEICHEN — 15 rekte Bestimmung und Benennung der von ihm unter- suchten Insekten gekiimmert, vielmehr hat er einfache deutsche Namen verwendet. Sein Augenmerk konzent- rierte sich ganz und gar auf die funktionellen Aspekte, auf Lebensweise und Entwicklung, während ihn taxo- nomische und systematisch Aspekte offenbar gar nicht interessierten. GEUS (1990) zitiert eine geradezu abfalli- ge Aussage VON GLEICHENs über jene Naturforscher, „die nur Systeme ausheckten, von nichts als Nomenkla- tur, Papillons, Konchylien u.d.g. mit einem Worte, nur immer von der Schale der Dinge schwatzten. Solche Menschen setzen meistens die Naturkunde nur zu einem blossen Gedaechtniswerke oder blos zu einer leeren Liebhaberei herab.“ 2. 1764A: „DER BLATLAUSFREBER UND SEINE EIER AN EINEM HANFBLAT“ (ABB. 2-6) In diesem Aufsatz schildert VON GLEICHEN, wie er Ende Juli 1761 an einer Hanfstaude ein Blatt fand, „an dessen beiden Seiten weise subtile Fäden, deren Endungen gleichfärbige kleine Knöpfchen hatten, sehr regelmäßig angeklebet waren“. VON GLEICHEN brach das Blatt ab, nahm es mit nach Hause und fand seine Vermutung, dass es sich bei den „Knöpfchen“ um Insekteneier han- delte, bestätigt, als er bei Betrachtung der Knöpfchen unter dem Mikroskop die durchscheinende Larve sah und plötzlich Zeuge des Schlüpfens der Eilarve wurde. VON GLEICHEN beschreibt Eier und Larven außerordent- lich genau und bildet sie auch in hervorragender Weise ab; seine Zeichnungen dienten Johann Christoph KEL- LER (1737-1795) als Vorlagen für die Kupferstiche (Abb. 2). Zu seinem Bedauern konnte er keine weiteren Zeichnungen anfertigen, weil die Larven nach drei Ta- gen tot und eingeschrumpft waren. VON GLEICHEN wusste zu diesem Zeitpunkt nicht, welche Insekten er vor sich hatte, was ihre Nahrung ist, ja er schreibt sogar ganz offenherzig: „Allein meine Ungewissheit ihrer Be- stimmung verleitete mich, zu glauben, dass sie wenigs- tens nicht viel größer werden würden, als sie waren, da ich eines derselben Fig. 4. vergrößert abbildete.* Erst nach dieser Untersuchung kam VON GLEICHEN das be- rühmte Werk von REAUMUR (1734-1742) in die Hände, in dessen drittem Band (1737) die Biologie von Chryso- piden beschrieben wird. VON GLEICHEN gibt auch un- umwunden zu, dass er, hätte er von REAUMURs Werk vorher Kenntnis gehabt, mit seinen „Blatlausfreßern viel sorgfältiger umgegangen seyn würde“. Erst durch REAUMUR erfuhr VON GLEICHEN , dass die „Fresszan- gen“ nicht zum Fressen, sondern zum Saugen dienen und dass sie „hohl und an den Spitzen offen sind“. Dass diese Saugzangen von größter systematischer Bedeu- tung sind und das die Ordnung Neuroptera begründende Merkmal schlechthin (also in heutiger Terminologie: die Autapomorphie) der Neuroptera sind, wusste auch REAUMUR nicht. Erst mehr als 100 Jahre später erkannte der junge Friedrich BRAUER den außerordentlichen sys- tematischen Wert des Merkmals, indem er alle Familien der Neuroptera (im heutigen Sinn) in einer Ordnung (er nannte sie damals Megaloptera) vereinte (BRAUER & Löw 1857). USE OY en Tabula 2, Der Blatlausfreßer und feine Eier an einem HDanfblat, u Ende des Sulii bes 1761ten Sabre erblickte id) unvermuhe des Sul 1 ng et an einer Hanfilaude ein Blat, an ar ee Sa ne ee a deren Endungen gleichfärbige Fleine Knöpfen hatten, ae rad) diejes Blat ab, und nahm es mit nad) Haufe, wo id) e Sch br gar bald ee, a en ba id) die Sinöpfchen, womit fich bie herabhangenden 7 tee hielt, Der braune Strich, welder, als ich diefe Eier unter dem Vergröi ferungaglafe betradhtete, Durch die feine Haut defelben durchfchien Fig. 2. beftdette mid) fogíeid) in o Meinung, und Die folgende DBegebenheit beftdttigte folche vollkommen. * Denn als id) mit der Abseihnung ines Diejer Eier befehäftiget tar, Öfnete fic) folches an feinem obern Theile, und cines der befonderften Thierchen Y fo mir nod) iemats vor die Augen getommen ift, fam aus demfelbigen hervor, Die langía: ‚me und mühfame Beroegung defelben geftattete Mir, eg absuseichnen, ia fo gar aud) ausjumahlen, . Die cou des Cies fant hiebei mad) dem Maafe, als fie leer wurde, fichtbarlich sufammen und rourde Dieienigen , toelche bag entylickende Vergnügen fennen, das bei berglcidjen Unt i foríd)enden und wißensbegierigen Beobachter rühret, werden mir glauben, Se u fage an se fem Augenbticte das prächtigfte Schaufpiel der Welt von der Belhauung diefes Wurms nicht abgerufen haben würde. Die 3te Figur ftellet Diefe CErfcheinung auf das richtigfe vor. Mann ficht das nod) jarte Infete y Wie es auf der Seite liegt, und mit allen Theilen feines Leibes jur Welt fome. An dem Kopfe sigen fic) 1) sroei gegeneinander gefrimte Gpisen, mit denen es die Schaale ded Cies durchftofien har, 2) finf im halben Zirfel ftehende Fleine Augen, tweldhe das mittlere größere umgeben, und 3) wie id) nach ber Deutlich gefehenen Bervegung des untern braunen Theiles des Kopfes fliege, Das Maul a. Die Begierde, mit diefen fremden Gäften genauer befant ju werden, veranlaßete mid) fogleich, ein anderes völlig ausgefchloffenes Thierchen von bem Hanfblate zu nehmen, und unter das Vergrdferungss glas ju bringen, tveldes dem erftbetrachteten vollfommen dhnlic) war. Des andern Tages waren alle Eier eer, und mein Hanfblat voll Heiner Thierchen , die aber viertel, ía halbe Stunden ndhtig hatten, um den Weg von ein paar Linien zu machen. Sie zeigten fid) aber auch viel größer und dunfler von Farbe Fig. 4. alg bieienigen waren, Die id) Den Tag vorher im ihrer Kindheit gefehen hatte. Was für ein fürchtertis dhes Thier würde ung diefer ABurm feyn, wenn er die Größe eines Pferdes, oder Ochfens hätte? Und twit grof ift nicht dic Weifiheit und Liebe des Schäpfers, Da er dem Auge des Menfdyen diefe Elcinen Ofieder feiner Schöpfungskette verborgen hilt? Go wie aber in diefer Kette eines gu des andern Erhaltung und Nahrung beflime it, fo zeiger auch das fürchterlihe Zangengebiß , twomit der Kopf Diefes Snfefts gerwafs net ift, Daf es im der Reihe des Flcinen Gervürms ein eben fo (chrectbares Raub thier finn mag , als der Bar und Der Wolf in der Reihe größerer Thiere, Und vielleicht erfezet die Lift, feinen Raub zu fangen, wasihm die Natur an der Gefdromdigkeit verfage hat, tie toir diefes an dem Ameifenräuber und WWurmldiwen fehen, Um meine Abbildung diefes Snfekts aber mehrers zu erläutern, will ich es kürzlich befhreiben. Der in Dergleihung mit der Größe des Körpers fehr große und breite Kopf ruhet auf einem furs gen Hals. Oben ift er mit einem hornidtem Helm bedectt, defen vorderes Theil meisgrau, das hintere aber braun ift, An den beiden vordern Seiten des Kopfes ftehen gwei umgegliederte, Durchfidhtige und glatte Fühlhörner neben den Augen, Unter diefen raget das Fürchterliche Zangengebifi mit gelben Spies hervor. Die Augen ftehen’gleih am Anfange des Zangengebißes, und könmen hier nicht alle fechfe gefehen werden, wie bei der Dritten Figur. Der Körper ift Fegelfórmig , aber, gumal die Bruft , mebr plat de b rund, Abb. 2: W. F. VON GLEICHEN (1764a), Erste Seite der Arbeit über den „Blatlausfreßer ...* Wahrscheinlich verdanken wir indes dem Umstand, dass VON GLEICHEN das Werk von REAUMUR nicht frü- her zur Hand hatte, die schón kolorierte Kupfertafel der Eier und der Erstlarve. (Die auf Fig. 5 und Fig. 6 dar- gestellten leeren Eier stammen von einem spáteren Fund des Gártners VON GLEICHENS auf einer Zwetschke.) Un- seres Wissens handelt es ich dabei um die erste farbige Darstellung einer Chrysopiden-Larve mit einer Wieder- gabe des Pigmentierungsmusters in der gesamten ento- mologischen Literatur. Die Darstellung ist so genau, dass der Versuch, die Larve zu bestimmen, lohnend erschien. Die Larven fast aller in Mitteleuropa vorkom- menden Chrysopiden sind gut bekannt, zu tiberwiegen- dem Teil existieren auch (allerdings nicht durchwegs 16 Bonner zoologische Beitráge 53 (2004) WF GRobf. et pint IC. Keller exo Abb. 3: W. F. VON GLEICHEN (1764a), Tafel zu der Arbeit über den „Blatlausfreßer ...“ Abb. 4: W. F. VON GLEICHEN (1764a), Ausschnitt der Tafel zu der Arbeit über den „Blatlausfreßer ...“: Die in (für C. pallens charakteristischen) Reihen abgelegten Eier. Abb. 5: W. F. VON GLEICHEN (1764a), Ausschnitt der Tafel zu der Arbeit „Blatlausfreßer ...*: Die aus dem Ei schlüpfende Lar- ve (von C. pallens). auch vom ersten Larvenstadium) gute Abbildungen (vgl. KILLINGTON 1936-1937; PRINCIPI 1940; GEPP 1983; DIAZ-ARANDA 1992; DIAZ-ARANDA & MONSER- RAT 1995). Vergleicht man die Larve VON GLEICHENS mit den verfiigbaren Abbildungen, so findet sich die größte Übereinstimmung mit Chrysopa pallens (Ram- bur, 1838). Wir haben den derzeit vermutlich besten Kenner der präimaginalen Stadien von Chrysopiden, Herrn Prof. Dr. Peter DUELLI (Birmensdorf) um eine Beurteilung der Abbildung in VON GLEICHENS Arbeit gebeten. Nach seiner Meinung lassen die in Reihen ab- gelegten Eier sofort auf C. pallens schließen. Ebenso spricht die Pigmentierung der Kopfkapsel am ehesten für C. pallens. Weiters schreibt er: „Die langen, am En- de gebogenen Borsten lassen auf einen trash-carrier schließen. Das ist C. pallens ja nicht. Allerdings sind bei allen Chrysopiden-Arten die Borsten des ersten Larven- stadiums, relativ zur Körperlänge gesehen, viel länger als im 3. Larvenstadium. Auch tragen viele Arten dann doch noch etwas trash (debris) mit sich herum, später nicht mehr. Verglichen etwa mit Chrysoperla-Larven Horst Aspöck & Ulrike Aspöck: Neuropterologische Beiträge in den Werken von Wilhelm Friedrich VON GLEICHEN — 17 sind bei C. pallens auch bei der L3 die Borsten sehr lang.“ Zusammenfassend hält Prof. DUELLI die Larven in VON GLEICHENs Arbeit mit größter Wahrscheinlich- keit für Chrysopa pallens. Abb. 6: W. F. VON GLEICHEN (1764a), Ausschnitt der Tafel zu der Arbeit über den „Blatlausfreßer ....“: Erstlarve (von C. pal- lens). Diese Art (früher als Chrysopa septempunctata Wes- mael, 1841, bekannt) ist eine extrem expansive, poly- zentrische Art mit Ausbreitungszentren in der West- und in der Ostpaláarktis und einer Verbreitung, die na- hezu ganz Europa (mit Ausnahme der nórdlichen Teile, jedenfalls ganz Mitteleuropa von der Ebene bis in die kolline Stufe), NW-Afrika sowie weite (auch tropische) Teile Asiens östlich bis Japan und Taiwan umfasst (ASPOCK et al. 1980; ASPOCK et al. 2001). Mit Sicher- heit war und ist die Art im Bereich des RUBWORM’ schen Schlosses, wie úberhaupt in Franken, háufig. (Zur Verbreitung von Ch. pallens in Deutschland siehe SAU- RE et al 2003.) Diese durch die erste kenntliche Farbabbildung einer Chrysopiden-Larve und die Beschreibung des Schliipf- vorgangs bemerkenswerte Arbeit VON GLEICHENS ist of- fenbar weitestgehend in Vergessenheit geraten. Selbst in der an Vollständigkeit kaum überbietbaren Bibliogra- phie von OSWALD (2004) fehlt sie. Ältere und z. T. so- gar kolorierte Darstellungen von Chrysopiden-Larven finden sich u. a. bei GOEDART (1669), ALBIN (1720), FRISCH (1736), REAUMUR (1734-1742), BONNET (1945, Übersetzung: GOEZE 1773), sie sind allerdings durch- wegs grob ausgeführt und lassen jedenfalls keine Art- Bestimmung zu. 3.1764B: „DIE HOFDAME“ (ABB. 7-8) Auch diese Arbeit mit dem ungewöhnlichen Titel fehlt in der Bibliographie von OSWALD (I. c.), obwohl sie die vermutlich größte je publizierte Abbildung einer Chry- sopide enthält. VON GLEICHEN schildert in dem der Zeit entsprechen- den narrativen Stil, wie sich „zur Zeit des Carnevals im Jenner bei später Nacht“ ein Insekt auf eine Spielkarte setzte, das durch seine Schönheit auffiel (weshalb ihm VON GLEICHEN den Namen „Hofdame“ gab). Tags darauf untersuchte VON GLEICHEN das Insekt mit frei- em Auge ebenso wie unter dem Mikroskop und führte eine kolorierte Zeichnung aus, von der J. C. KELLER einen Kupferstich anfertigte, den VON GLEICHEN kolo- rierte. Das Insekt ist von jedem Entomologen sogleich als Chrysopide zu erkennen, und jeder Neuropterologe weiß, dass es sich dabei nur um eine Chrysoperla-Art handeln kann. Die in Mitteleuropa vorkommenden Chrysoperla-Arten überwintern als Imagines, und unter diesen nehmen manche (in Mitteleuropa als einzige Chrysopiden) im Herbst eine gelblich-bräunlich-rötliche Färbung an. Vor 40 Jahren wäre die Bestimmung völlig problemlos erschienen. Damals kannte man nur eine einzige Chrysopiden-Spezies unter den Chrysopiden Mitteleuropas, die als Imago überwintert und im Herbst ihre Färbung von grün in braun verändert: Chrysoperla carnea (Stephens, 1836). Im Verlauf der vergangenen Jahrzehnte hat sich gezeigt, dass „die alte Chrysoperla carnea“ nur als Kollektivnahme für mehrere morpholo- gisch schwierig zu differenzierende Spezies zu betrach- ten ist, von denen mindestens 5 in Europa und z. T. auch in Mitteleuropa vorkommen (Literatur hierzu: HENRY & DUELLI 1999; HENRY et al 1996, 2002; HENRY et al 2003; JOHNSON et al 2003; ASPOCK et al. 2001): Chrysoperla carnea (Stephens, 1836) Chrysoperla lucasina (Lacroix, 1912) Chrysoperla mediterranea (Hölzel, 1972) Chrysoperla pallida (Henry, Duelli & Johnson, 2002) Chrysoperla agilis (Henry, Brooks, Duelli & Johnson, 2003). 18 Bonner zoologische Beitráge 53 (2004) = Die Hofdame. 5 / des Heren von Reaumurs [ese , welche er PL i en von . 32, Fig. ir il i dl Albee bie A da ich fie beobachtete, nicht fo AA al e se E re = (hon ee Urfachen nicht gefehen, Der setae ; % e Faumt 15. bid 16, X in ei i a tocifer Seide einfpinnen, aus wilder le ly alates ene = E o eal fia) aber erft im Derbjte einfpinnen, das folgende Sribiabe eine fehr fd)öne lege ee ee ae, oe ihre Eier an die Aeffe, oder Blätter der ihr anfländigen. Pflanze, mehrers hievom finder der g. & am ae a en Erélárung der Figuren, Fig. 1. Ein Hanfblat mit Snfeften Giern, Fig. 2. Ein foldjes Gi vergrößert. Fig. 3. Der Blatlausfreßer, wie er aus dem Gi zur Welt Fomt, Fig. 4. Derfelbe einen Tag fpdter beobachtet, Fig. 5. Dergleichen Eier auf einem Stüdcyen Zmerfehenhaut vergrößert, Fig. 6. Diefes in natúrtider Größe. Fig. 7, Das von der Zmetichenhaut losgernachte untere Theil des Cierfabens , defen trichterförmige Geftate seiget, toie der Stil aus dens Flebrid)ten Anfaze, gleid) dem Fade aus dem Spiny socten, herausgefponnen ift, Tabula 3. Die Hofdame. (s, olte id} twol nicht yu entíchulbigen fon, Daß id) diefem fiegenden Spnfekte ben Namen bes Schmus des der Höfe beilege, da es cin fo prád)tiger Schmuck der Natur ¡ft 2 Und maffen fid) nicht die Blumiften und Gärtner gleiches Recht an, tenn fie ihren Blumen und Früchten fogar die prächtigen Titel der Kaifer und Könige geben ? Sie felbften, meine gndbige Damens, werden dieerften fepn , mir diefe Vergleidung zu vergeben, fobald fie einen Blick auf die Worftellung diefer (hdnen ¿liege thum mollen. hr eigen Geroiffen wird ihnen fagen, daß nid)t nur diefe Benennung ungemein rol getroffen fey, fondern dag au) diefes Snfett an Pradt und Schönheit fogar in ber Vergrößerung mance Hofdame in ihrer Robe übertreffe, zumal wenn fie bedenfen wollen, daß unter allen ihren Schönheiten nicht eine einige dem Kaufmanne und Schneider zugehöre. Nod) mehr, es gehöret diefes Infekt gerigermaßen zu Dem Gef'plechte der Sungfrauen. > Es liebet endlid) diefe Hofdame aud bas Spiel. Dann eben beiden Spiele war es, als fie fid) zur Zeit des Carnevals im Spenmer bei fpáter Nacht einfand , und auf bie Karte feste. Kaum hatte fie fich ein wenig hier umgefehen, als id) ihr, um des andern Tages nähere Befantichaft mit ihe ju machen, fogleich ein Quartier bei meinem Mitcoffop amvies. Die fhönen las furgeümen Augen, die id) (chon ohne Vergrößerung bei Licht fahe, liefen mid) auf die Entdecungen nod) "mehrerer Schönheiten hoffen. Die vergráferte prueite Sigur der erflen, welche die Hofdame im matüclicher Größe vorfielt, wird zeigen, dag id) mich’micht betrogen Habe, Am Kopfe fieht man eine erhabene Stim, und unter folder ¡woo von vorne länglicht nate Lippen a, An Des untern Lippe gehen 2: 2 Abb. 7: W. F. VON GLEICHEN (1764b), erste Seite der Arbeit über „Die Hofdame““. Diese fünf Taxa unterscheiden sich vor allem durch spezifische Vibrationsmuster, die sich durch verschie- den langes, durch unterschiedliche Intervalle und Fol- gen gekennzeichnetes Vibrieren mit dem Abdomen ergeben. Die Vibrationen werden auf das Substrat über- tragen und erreichen über dieses andere Individuen, die die spezifischen Signale mit den Subgenualorganen der Beine wahrnehmen. Beide Geschlechter zeigen dieses der Partner-Findung dienende Verhalten, und wenn zwei Phäna mit unterschiedlichen Vibrationsmustern zusammentreffen, kommt es nicht nur nicht zur Paa- rung, in vielen Fällen ist sogar eine aggressive Reaktion die Folge. Die Taxa sind Spezies mit zum Teil unter- schiedlich fortgeschrittener genetischer Isolierung. Sie werden häufig als „song-morphs“ bezeichnet. Unter ex- perimentellen Bedingungen können Paarungen zwischen verschiedenen „song-morphs“ manchmal „erzwungen“ werden. Ch. mediterranea und Ch. lucasina unterscheiden sich morphologisch hinlänglich und konstant von den ande- ren drei Spezies, deren morphologische Unterschiede gering sind. Alle genannten Chrysoperla-Arten über- wintern als Imagines, aber nur drei von ihnen (Ch. car- nea, Ch. pallida, Ch. agilis) verändern im Herbst die Färbung von grün zu rötlich-bräunlich-gelblich. Diese Färbung wird während der gesamten winterlichen Dia- pause bis zum nächsten Frühjahr beibehalten. Für die Deutung der Abbildung der „Hofdame“ (Abb. 8) VON GLEICHENS scheiden also Ch. lucasina und Ch. mediter- ranea jedenfalls aus. Auch die in verschiedenen Teilen des Mittelmeerraums nördlich bis in die Südschweiz nachgewiesene Ch. agi- lis kann ausgeklammert werden, sie ist im übrigen bis- her in Mitteleuropa nicht nachgewiesen worden und kommt mit großer Wahrscheinlichkeit in diesem Teil Europas tatsáchlich nicht vor. Von den verbleibenden zwei Arten — Ch. carnea und Ch. pallida — gilt Ch. pal- lida als ,,Waldart™, die nur selten in Häusern úberwin- tert. Ch. carnea hingegen ist ein ausgeprägter Kulturfol- ger und eine in allen Städten und Dörfern Mitteleuropas häufige Florfliege. Auch in diesem Fall haben wir den führenden Chryso- perla-Spezialisten, Prof. Dr. Peter DUELLI (Birmens- dorf), um seine Beurteilung des kolorierten Kupferstichs gebeten. Seine Antwort: „Die Färbung der Hofdame ist exakt das, was ich für C. carnea s. str. erwarten würde. Auch die Tatsache, dass sie im Januar im Haus anflog, spricht für carnea, C. lucasina ist wegen der Färbung ausgeschlossen, C. pallida (Waldart) kommt selten in die Häuser und ist etwas bráunlich-gelber im Winter“. Wir können also tatsächlich davon ausgehen, dass W. F. VON GLEICHEN die erste kenntliche — und überdies far- bige — Darstellung jener Spezies veröffentlicht hat, die STEPHENS (1836) mehr als 70 Jahre später als Chrysopa carnea nomenklatorisch gültig beschrieben hat und die mit Sicherheit die in der Literatur am meisten behandel- te Neuropteren-Spezies ist. Chrysoperla carnea ist ZU- dem gewiss auch die am intensivsten und unter den ver- schiedensten Gesichtspunkten untersuchte Neuropteren- Art. VON GLEICHENs Abbildung ist — obwohl 240 Jahre alt — keineswegs die erste Abbildung einer Florfliege. Darstellungen von Chrysopiden findet man schon bei MOUFFET (1634), ALDROVANDI (1638), JONSTON (1657), GOEDART (1668), FRISCH (1736), RÖSEL VON ROSENHOF (1755), und natürlich bei REAUMUR (1734- 1742) und bei BONNET (1745) und anderen früheren Autoren. Einige Autoren (GOEDART, ALBIN, RÖSEL VON ROSENHOF) haben sogar kolorierte Abbildungen veröf- fentlicht. Allerdings ist eine sichere Identifizierung der Art in keinem Fall möglich. Man muss jedoch gerech- terweise erwähnen, dass drei Umstände die Identifizie- rung der von VON GLEICHEN dargestellten Florfliege ermöglicht haben: die genaue Kenntnis des Fundorts (Schloss Greiffenstein in Unterfranken), das Funddatum (Januar, aktiv im Haus) und die bräunliche (Ver-) Fär- bung des Tiers. Das Geäder hat VON GLEICHEN zwar sehr schön (und sogar — erstmals in der Literatur — mit Horst AspOCK & Ulrike ASPÓCK: Neuropterologische Beiträge in den Werken von Wilhelm Friedrich von GLEICHEN 19 doppelten Strichen), aber alles andere als richtig ge- zeichnet. Zu jener Zeit wusste man natiirlich noch nichts iiber Geáder-Merkmale von Chrysopiden und deren sys- tematische Bedeutung, weshalb man — das gilt geradezu fiir alle Autoren jener Zeit — auf eine korrekte Wieder- gabe des Geäderverlaufs und der Zahl und Position der Queradern kaum achtete. Tab. 3, S Ja Sp y NS \ S N 93 N N N 27 PITTEEERRLERRRERER N >72 SA E Se AGS ss = 2.CKeller exc. Abb. 8: W. F. VON GLEICHEN (1764b), Tafel zu der Arbeit über „Die Hofdame“ (= Chrysoperla carnea). VON GLEICHEN mutmaßte, dass seine „Hofdame“ „die nemliche ist, welche nach der Beschreibung des Herrn von REAUMURs aus der seidenartigen Puppe hervor- komt“. Das ist sicher nicht richtig, aufgrund der biologi- schen Angaben von REAUMUR kann es sich nicht um ei- ne Chrysoperla-Art gehandelt haben. Auch VON GLEICHENs Vermutung, dass seine „Hofdame‘“ sich aus einem „entkommenen Blatlausfreßer“‘, der im „Zimmer eine ... anständige Speise“ gefunden habe, entwickelt habe und „dass durch die Wärme geheizter Zimmer, wie dies bei allen sich verwandelnden Insekten geschieht, das Wachstum und Auskriechen der Fliege dergestalt befördert worden ist, daß letzteres im Jenner erfolget, da es nach Herrn von REAUMUR bei den Spätlingen erst im Frühjahr geschieht..“ trifft natürlich nicht zu. REAUMUR hat über eine Spezies geschrieben, die nicht als Imago überwintert. Erstaunlich ist indes, dass VON GLEICHEN offenbar erst kurz vor der Veröffentlichung seines Artikels über die „Hofdame“ (er war damals immerhin schon 47 Jahre alt) eine Florfliege beachtet hat, denn nichts in den bei- den Artikeln, die sich mit Chrysopiden befassen, deutet darauf hin, dass VON GLEICHEN diese Insekten vorher gekannt hatte. Kaum ein an der Natur und insbesondere an Insekten interessierter Mensch in Mitteleuropa be- gegnet nicht schon früh, meist irgendwann in der Kind- heit, einer Florfliege und erkennt die charakteristischen, in etwa 30 Arten in Mitteleuropa vorkommenden Neu- ropteren immer wieder. Auch und besonders an den bräunlich verfärbten Florfliegen kann man im Spät- herbst, Winter oder Vorfrühling kaum vorübergehen, weil sie in fast allen Häusern, die von Gärten oder gar Feldern umgeben sind, vorkommen; oft findet man sie — besonders in alten Häusern — zwischen Fenstern, wo sie manchmal in Ritzen überwintern. In allen größeren entomologischen Werken, die vor 1764 existierten, sind auch Chrysopiden behandelt und auch abgebildet worden. Bei aller Würdigung der Verdienste VON GLEICHENS er- staunt es doch, dass ein in einer so sorgenfreien mate- riellen Situation lebender Mensch mit einer so aus- geprägten Neugierde für die Natur nicht über die entomologischen Standard-Werke, die im 17. und 18. Jahrhundert erschienen waren, verfügte. Er muss dem- nach zumindest in den ersten Jahren seiner naturwissen- schaftlichen Studien auf Schloss Greiffenstein nur eine bescheidene Bibliothek besessen haben. 4. 1770: „VERSUCH EINER GESCHICHTE DER BLATLÄUSE UND BLATLAUSFRESSER DES ULMENBAUMS“ (ABB. 9-12) Diese 1770 und in einer 2. Auflage 1787 erschienene, insgesamt 30 Seiten umfassende und mit vier kolo- rierten Kupferstichen ausgestattete Publikation zählt zu den bekanntesten Arbeiten des Autors und ist zumindest von MORTON (1910), WEIDNER (1980), GEUS (1989) und DETTNER (1997) besprochen worden. Im Vorder- grund der Sekundärliteratur steht der Nachweis des Le- benszyklus und insbesondere der Parthenogenese der Ulmenblattlaus, Byrsocrypta ulmi (L.), durch VON GLEICHEN sowie dessen Überlegungen zu diesem Phä- nomen. Im Zuge der Befassung mit den Blattläusen stieß VON GLEICHEN auf einen der Prädatoren unter den Neurop- teren, jene charakteristische Hemerobiiden-Spezies, die LINNAEUS (1758) als Hemerobius phalaenoides be- schrieben hat und die später in das Genus Drepanepte- ryx Leach transferiert wurde. 20 Bonner zoologische Beitráge 53 (2004) Verfu ch einer Gr (hichte ver Batan Blorlausfreifer des Lllmenbaumd nebft vier mit Farben erleudteten Kupfertafeln von Wilhelm Friederich Frevherrn v. Gleichen genannt Nußmworm, Herrn auf Greifenftein, Bonnland und Ejelbad) , Groß: Creuz des Hodfirflidy Branden, burgifchen rothen Adler» Ordens, und Hodfiirftlid) Brandenburg: Culmbachifhen Geheimen» Rath. Mebft einer Vorrede des Herrn Hofraths und Prof. Delius. EES ee A er Tee Sn Kupfer gebraht, und verlegt von Georg Paul Nufbiegel a Múrnberg y gedructe bei Johann Seinridy Gottfried Bicling, 3.7.7.0% 8 by Abb. 9: W. F. VON GLEICHEN (1770), Titelblatt (Bibl. H.& U. ASPOCK) Etwa sieben Seiten Text und zwei kolorierte Kupfersti- che widmet VON GLEICHEN der Beschreibung der Larve und der Imago von D. phalaenoides (Abb. 10 - 12). Die Zeichnungen hat VON GLEICHEN — wie in seinen Wer- ken durchwegs — selbst angefertigt, die Tafeln wurden von seinem Verleger, dem Kupferstecher Georg Paul NUBBIEGEL (um 1713 — 1786), gestochen. VON GLEICHEN beschreibt, wie er in dem Gewimmel der jungen Blattläuse “einige braune Würmer“ erblickt habe, ,,die sehr ämsig waren, und ihre 6. langen Beine mit ungemeiner Geschwindigkeit zu gebrauchen wusten“. Er hielt „sie sogleich für das, was sie waren, nemlich für diejenige Art Blatlausfreßer, die bei dem Herrn von REAUMUR die zwote Claße machen, und wel- chen er den Nahmen Blatlaus-Löwen, Lion de Pouce- rons, gegeben hat“, womit VON GLEICHEN recht hatte. Er beschreibt die Larve von D. phalaenoides sehr genau und gibt eine gelungene Abbildung (Abb.11: Tab. III, Fig. 21-22). Erwähnung verdient auch seine Bemerkung über den Geruch der Larven, er sei „außerordentlich stark, und dabei sehr angenehm, nichts anders als Quän- del und Thimian. Vielleicht ist er die Witterung der Blatläuse die sie locket, ihrem Feinde in ‘der Nähe zu bleiben.“ Tatsächlich haben die Larven von D. phalae- Berfud einer Gefchichte der Blatlaufe und Blatlausfreffer des Ulmenbaumes. aii Der Ulmenbaum, der, wegen feiner did belaubren Zweige, eine Zierbe der Garten Do, ift, gibt einer Art Infeften , die wic Blatläufe nennen, auf feinen breiten Bla: A tern Wohnungen , In folder Menge birnförmiger Blafen, daß fic) y wenn fie in der Mitte des Bradmonate Ihre Bolfommenheit erlanger haben, Xefte un? Zweige davon biegen. Die erfte Figur der erften Tafel zeiget ein folhes Blat, das an der Spike einige dies fer Blafen von verfihledener Größe träge, Die erften habe ich zu Anfang des Manmonats, da fic noch fehr Flein waren, aufgemahr. Id) fand cin einiges braunes, dicflelbiges, ungeflügel, tes, fehr fleincs InfeFt dariunen, welches in allem fo viel ähnliches mit den fo genannten Blatláufen hatte, daß es mir nicht fd)mer fiel, es davor zu erfennen. BLeuwenhoek, Sartfoecker, A de Abb. 10: W. F. VON GLEICHEN (1770), erste Seite (mit der von »J.d.G.R.“ [= J. VON GLEICHEN-RUSSWORM? Ein Verwandter?] gezeichneten und von Johann NUSBIEGEL (1750-1833) gesto- chenen Vignette. noides einen charakteristischen Geruch, den FULMEK (1941) mit dem von Anis- oder Fenchelól vergleicht. VON GLEICHEN beschreibt das Verhalten der Larven sehr treffend und widmet sich dann recht ausführlich den Mundwerkzeugen. Er lobt auch an dieser Stelle zu- nächst die Arbeiten von REAUMUR, schreibt aber dann, dass er — REAUMUR — „gleichwohl bei der Untersuchung erstgedachter Kopfteile des Blausfreßers [sic!], ohnfehl- bar aus Mangel recht brauchbarer Vergrößerungswerk- zeuge, dießmal von seiner gewöhnlichen Bahn abge- kommen“ sei. Und weiter meint er, er habe daher „viel vergebliche Zeit und Mühe angewendet, auch hier [sei- ne] Beobachtungen‘ mit jenen von REAUMUR „zu ver- einigen, ohne die Oefnungen an den Spitzen der Fress- zangen des Blatlausfreßers zu finden“. Darauf zitiert er REAUMUR im Original, der völlig richtig Bau und Funk- tion der Saugzangen beschreibt. Merkwürdig, dass VON GLEICHEN bei der Beschreibung der Mundwerkzeuge von Drepanepteryx phalaenoides nicht auf seine 1764 veröffentlichte Untersuchung über den „Blatlausfreßer . an einem Hanflblat“ und auf das Bezug nımmt, was er dort über REAUMURS Meinung über die „Fresszan- gen“ gesagt hat, nämlich, dass sie nicht zum Fressen, Horst ASPOCK & Ulrike ASPOCK: Neuropterologische Beiträge in den Werken von Wilhelm Friedrich VON GLEICHEN 2] Zz TAB. Mm. MERLO GR pm Abb. 11: W. F. VON GLEICHEN (1770), Tafel IH. mit der Larve des „Ulmenblatlausfressers“ (= Drepanepteryx phalaenoides). TAB.IV rams Abb. 12: W.-F. VON GLEICHEN (1770), Tafel IV: mit Puppen- kokon (Fig. 30) und Imago (Fig. 31-33). sondern zum Saugen dienen und ,,hohl und an den Spit- zen offen sind“ (siehe oben). Vielmehr kam er jetzt zu dem Schluss, dass sie an den Spitzen keine Offnungen haben, weshalb er zwischen den Saugzangen nach der Mundóffnung suchte und schlieBlich eine solche auch fand und sogar abbildete (Abb.11: Tab. III, Fig. 23). Dazu schreibt er: „Da aber das Maul dieses Insekts nur alleine zum Saugen, und nicht zum Nagen gebrauchet wird, so ist auch der genaue Zusammenschlus der Lip- pen eine natúrliche Folge dieser Bestimmung, so, wie der Unmöglichkeit, solches auf andere Art, als durch ei- nen wohl angebrachten Druck des Kopfes zu ófnen und zu sehen.“ Die akribische práparative und mikroskopi- sche Arbeit VON GLEICHENS verdient alle Bewunderung. Die Nahrung wird bei allen Neuropteren-Larven über die Saugzangen aufgenommen, die Mundöffnung ist ei- ne zwischen den Saugzangen quer verlaufende, falzartig verschlossene, kaum erkennbare Spalte ohne jede Funk- tion für die Nahrungsaufnahme. Wie hat sich VON GLEICHEN die Nahrungsaufnahme wohl wirklich vorge- stellt, da er meinte, das die „Fresszangen“ keine Öff- nung haben, das Maul aber zum Saugen dient. Es ist tat- sächlich erstaunlich, mit welcher Bestimmtheit VON GLEICHEN die richtige Deutung der Saugzangen durch REAUMUR verwirft und durch eine durchaus falsche und unlogische Behauptung ersetzt. Dann beschreibt VON GLEICHEN die Entwicklung von D. phalanoides, gibt die Larvalperiode mit 13 bis 14 Tagen an, erwähnt den „Bemerkungswürdige[n] Umstand, bey diesem Insekt, daß es dieses mit den Spinnen gemein hat, aus dem Ende oder der Spize seines Leibes, und nicht, wie andere Insekten aus dem Munde zu spinnen, und zwar nur zu der Zeit, wenn man das Gespinnste zu seiner Verwandlung anfängt, ohne zuvor jemals nur das geringste Merkmal, daß er hiezu geschickt sey, gegeben zu haben.“ VON GLEICHEN gibt eine schöne Abbildung des Kokons (Abb. 12: Tab. IV, Fig. 30) und erwähnt dann, dass zwei Imagines geschlüpft seien. Es folgt eine ausführliche und anschauliche Beschreibung der Imago, ergänzt durch die schöne kolorierte Kupfertafel. Ge- rechterweise muss man aber sagen, dass die Art zwar sofort zu erkennen ist (nicht zuletzt, weil es in Europa keine andere nur halbwegs ähnliche Hemerobiiden- Spezies gibt und die zweite in Europa vorkommende Art des Genus, D. algidus (Erichson in Middendorf, 1851), sich habituell erheblich unterscheidet), dass aber das Geäder recht fehlerhaft gezeichnet ist. Auch hier gilt das bei VON GLEICHENs Abhandlung über die ,,Hofda- me“ Gesagte. Im 18. Jahrhundert unterschätzte man ein- fach die Gesetzmäßigkeiten des Flügelgeäders von In- sekten erheblich und achtete bei bildlichen Wiedergaben vielmehr darauf, den Habitus möglichst typisch zu erfas- sen — was VON GLEICHEN ja auch recht gut gelungen ist. REAUMUR (1737) hat die Larve, den Kokon mit der Puppe und die Imago des 21 Jahre später von LINNAEUS u , ] 5 . i. nig a ae fa ree ö \ a ö “yt n Kor ' y 7 Fi 1 7 1 o 0 Ir a D de A ' wit . a . o \ . od A u no : Pd A 4 . . y [m u ‘ ‘ . \ ] ‘ mm y i 1 . 7 ye ; ne : - 1 . b ' ‘ \ uy . ; : Zoe A 1 a ' y . de \ 0 ] 1 ' N ae A B ö 7 ma \ A 5 y 1 ñ ñ vat A Axel HOFMANN: Neue Zygaena-Taxa aus Siidosteuropa, Vorder- und Zentralasien 91 93 Axel HOFMANN: Neue Zygaena-Taxa aus Südosteuropa, Vorder- und Zentralasien Axel HOFMANN: Neue Zygaena-Taxa aus Súdosteuropa, Vorder- und Zentralasien 95 Axel HOFMANN: Neue Zygaena-Taxa aus Südosteuropa, Vorder- und Zentralasien 97 Bernhard A. HUBER, Abel PEREZ & Renner L. C. BAPTISTA: Leptopholcus in Continental America 10] er d | Figs. 2-13: Habitus and female external genitalia photographs. 2-3. L. brazlandia. 4-5. L. pataxo. 6-7. L. evaluna. 8-9. L. brazlandia, female abdomen, lateral view, and external genitalia, ventral view. 10-11. £. pataxo, female, lateral view, and exter- nal genitalia, ventral view. 12-13. L. evaluna, female abdomen, lateral view, and external genitalia, ventral view. Bonner zoologische Beitráge Band 53 (2004) Heft 1/2 | Seiten 109-110 Bonn, Juni 2005 First Record of Hyles dahlii (Geyer, 1827) (Lepidoptera: Sphingidae) from the African Mainland ' / Anna K. HUNDSDORFER, Dresden 'The hawkmoth species Hyles dahlii (Geyer, 1827) is considered to be restricted to Corsica, Sardinia and the Balearic Islands (REBEL 1934), although there are occa- sional reports from Sicily (PITTAWAY 2004). In addi- tion, it has also been found on the Catalan coast, in northeastern Spain, in 1975 (MASO PLANAS et al. 1979) and on the island of Pianosa, off the Tuscan coast (Italy) in 1998 (DAPPORTO et al. 1999). Here I report the discovery of about 15-20 first instar (L1) larvae of H. dahlii feeding on Euphorbia paralias in northern Tunisia in October 2004. The exact locality is the beach at Residence Shiri, a small hotel and camp- ing site on the coast north of Sejane between Cap Serrat and Bizerte. I collected and reared about a dozen indi- viduals. After two moults it was possible to determine them as A. dahlii (a caterpillar is illustrated in Fig. 1 and an adult in Fig. 2). This species has to date not been reported from Africa. Voucher specimens will be depos- ited at the Zoologisches Forschungsinstitut und Museum Alexander Koenig (Bonn) and the Natural History Mu- seum (London). I doubt that H. dahlii is already established on the Afri- can mainland. I observed very few habitats for spurge- feeding hawkmoths in Tunisia, as I found little food plant in the parts of the country I visited. During my one-week tour (from the northern coast due south near the Algerian border, then back north along the eastern coast), I found herbaceous Euphorbia only along one inland stretch of road; otherwise there was only E. paralias along the coast. In the north, there are few sandy coastlines and these are often short and very iso- lated, with stretches of rocky coast and cultivated pine forest surrounding them. In parts of the eastern coast, the air is so humid that water condenses on the plants and the sand becomes baked onto them, so that they do not appear to be easily eatable by caterpillars. Also, the beaches are often partly or wholly developed for the tourist industry and thus it is possible that the popula- tions of E. paralias have become relictual or destroyed due to human activities. I found no other Hy/es caterpil- lars at the beach of Residence Shiri, or any indications 1 Clas Michael Naumann zu Königsbrück (26.06.1939 — 15.02.2004) zum Gedenken of prior occurrence, such as droppings or nibbled plant sprouts. In contrast, on other beaches in Tunisia, and one especially on the north coast, as well as at one local- ity inland, I found numerous caterpillars of A. tithymali, but I did not find any other A. dahlii caterpillars. There- fore, I consider this first record of H. dahlii from the Af- rican mainland is probably the result of a rare dispersion event, possibly of only a single fertilised female from Sardinia, which lies geographically nearest to the Tuni- sian locality. Fig. 1: Fifth instar larva of Hyles dahlii found on Euphorbia paralias in Tunisia in October 2004 (feeding in captivity on E. myrsinites ). 110 Bonner zoologische Beitráge 53 (2004) Fig. 2: An adult moth of Hy/es dahlii, emerged from the pupa in February 2005. The individual was collected as first instar larva in Tunisia in October 2004 and reared in captivity. Acknowledgements. I would like to thank Michael Korn (Konstanz) for this successful joint collecting trip to Tuni- sia and his specific help in looking out for Euphorbia plants and Hyles caterpillars. Special thanks also go to Paolo Mazzei (Rome), who first determined these larvae as H. dahlii on the basis of the single specimen I gave to him during my return journey. I am indebted to lan J. Kitching (London) for his correction of my language and for very helpful suggestions to this note. I would also like to ex- press gratitude to the late Clas Naumann for his long-term interest and participation in the collection of Hyles samples for our joint studies with Ian J. Kitching and Michael Wink (Heidelberg). REFERENCES DAPPORTO, L., CECCHI, B., LO CASCIO, P. & SFORZI, A. (1999): Contributi alla conoscenza dell'artropodofauna dell'Isola di Pianosa (Arcipelago Toscana). II. Prima nota sui macrolepidotteri (Insecta Lepidoptera). Bol- lettino della Societa Entomologica Italiana 131: 245- 232, MASÓ PLANAS, A., PÉREZ DE-GREGORIO, J. & SIMO CA- STELS, L. L. (1979): Primera cita d'Hyles dahlii (Lep. Sphingidae) al continent Europeu. Treballs de la So- ciétat Catalana de Lepidopterologia 2: 11-17. PITTAWAY, A. R. (2004): Sphingidae of the Western Palae- arctic: Hyles dahlii (Geyer, 1827). http://tpittaway.tri- pod.com/sphinx/h_dah.htm REBEL, H. (1934): Lepidopteren von den Balearen und Pi- tyusen. Deutsche Entomologische Zeitschrift Iris 48: 122-138. Author’s present address: Anna K. HUNDSDORFER, Staatliches Naturhistorische Sammlungen, Museum fúr Tierkunde — Zoological Museum, DNA — Laboratory, Königsbrücker Landstr. 159, D-01109 Dresden, Germa- ny, anna.hundsdoerfer@snsd.smwk.sachsen.de 136 tory Museum (BMNH), London, and the Museum fiir Naturkunde (ZMB), Berlin. Biometric and scalation characters as well as colour pat- tern data were evaluated in both living and preserved specimens. Measurements were taken with a dial calip- per (to the nearest mm). The following abbreviations are used: a.s.l. = above sea level, SVL = snout vent length, TL = tail length, SE = distance from snout tip to front of eye, EE = distance between hindmargin of eye to hind- margin of ear, HW = maximum head width, HH = maximum head height, HL = head length (from tip of snout to hindmargin of ear). The following scale counts were taken: supralabials (SPL), sublabials (SBL), nasals (N); in direction from rostral to labial: nasorostrals, su- pranasals, postnasals, internasals (IN); postmentals (PM), gulars bordering the postmentals (GP), interorbi- tals (IO), ventrals (V), scales around midbody (SB), subdigital lamellae under the first (LT1) and fourth toe (LT4), preanal pores (PP), postanal tubercles (PAT), dorsal scale rows in the middle of the third caudal whorl (S3R). In living specimens symmetrical scale counts were taken only on the lizard's right side. Colouration is described according to the standard plates published by GRALLERT & ROLAND (1960). 3. RESULTS 3.1. Description of Gekko scientiadventura sp. n. 3.1.1. Diagnosis. A small-bodied species of Gekko. The biggest specimen (captive) has a total length of 154 mm, of which 81 mm are made up of the tail. Habitus slen- der, head distinctly broader than neck, head and body slightly depressed. Unregenerated tail always longer than head-body length, not constricted at its base, not thickened, slighty depressed. Lateral fold weak, upper side of head, body and tail without tubercular scales. Digits and toes slightly webbed at the base, only inter- space between toes 4 and 5 not webbed. All digits and toes, except the inner ones, clawed. 14-17 subdigital lamellae below fourth toe. 5-8 preanal pores. Nostril touches rostral. No internasals. Posterior ciliaries spiny. Upper side yellowish to brownish in life. Dorsally seven large light spots that may be expanded to lateral narrow wavy bands. Tail with 7-10 light crossbands. Gular re- gion and parts of venter marbled. From all species of Gekko known to occur in Vietnam (G. badenii, G. chinensis, G. gecko, G. japonicus, G. palmatus, G. ulikovskii), G. scientiadventura sp. n. is invariably distinguished by the lack of middorsal tuber- cles. For more detailed comparison, also with other Gekko-species see below (3.4. and 3.5.). 3.1.2. Holotype. ZFMK 76198, adult male, Figs. 1-3, primary limestone forest, Phong Nha - Ke Bang, sur- roundings of 17.32 N, 106.16 E, ca. 50-150 m a.s.l., Bonner zoologische Beiträge 53 (2004) Quang Binh Province, Vietnam, coll. H.-W. HERRMANN, VU NGOC THAN & T. ZIEGLER, 27-29 Au- gust 2001. Fig. 1: Live holotype of Gekko scientiadventura sp. n. (ZFMK 76198). Phot. T. ZIEGLER 3.1.3. Description of the holotype. Total length 128 mm (for more measurements and indices see Table 1). Rostral twice as broad as high, narrower than mental, without a median suture. 13/12 supra- and 11/10 sublabials. Nostril touches rostral and Ist supralabial. 3/3 nasals, nasorostrals in contact medially. Nasorostrals twice as big as postnasals. Pupil vertical. Tympanum obliquely oval, ca. half of the eye diameter. Interorbital scale count 44. Mental triangular, as broad as high. Two postmentals, twice as long as broad and of subequal length with the Ist supralabial. Postmentals posteriorly touched by six scales. Lateral scales of snout about three times bigger than those on the dorsal side of the snout, roundish, juxtaposed. Head scales granular, ciliaries posteriorly spiny. Gular scales granular, about the same size as upper head scales. Dorsal scales round, smooth, juxtaposed, convex, ca. 1.5 times bigger than head scales. Ventral scales flat, smooth, imbricate, three times as big as dorsals, 41 in an oblique row between the weakly developed lateral folds. Midbody scale count 140. Scales on the upper surface of the upper fore- and hindlimbs flat, smooth and imbricate, granular on the lower. Lower parts of all limbs with granular scales. Fingers and toes basally weakly webbed, only toes 4 and 5 without webbing. 14/12 lamellae under the Ist and 17/15 under the 4th toe. 5 preanal pores arranged in an angularly bent series, 3/3 postanal tubercles. Upper caudal scales flat, subimbricate, as big as the dorsals, ar- ranged in regular transverse rows and weakly expressed whorls, the 3rd whorl being composed of (dorsally) 10 scale rows. Subcaudals flat, smooth, imbricate, 2.5 times wider than high, with a median row of regularly arranged plates; each 3rd plate ca. 3 times as broad as high, marking the beginning of whorl. Laterally, the subcaudals are bordered by alternating scales (2-2-3-2- H. ROSLER, T. ZIEGLER, VU N.T., H.-W. HERRMANN & W. BOHME: A New Gekko Species from Vietnam 157 2-3-2-2 ...) where scales each are bordering the respec- tive broader subcaudal plate. = ia y Fig. 2: Ventral view of the live holotype of Gekko scientiad- ventura sp. n. (ZFMK 76198). Phot. T. ZIEGLER Hemipenial morphology. Right hemipenis (Fig. 3: terminology after ROSLER 1998) 2.2 mm wide, 4.6. mm long, clavate, forked. Pedicel and truncus not dif- ferentiated from each other; basally on the sulcal side is a small wrinkled adminiculum, running laterocranially and not reaching the sperm groove. Above the admini- culum a ribbed area. Apex bilobed, the lobes being sub- equal in size. Sperm groove originates from the latero- caudal side of the organ and runs straight along the truncus, where it divides into two branches that termi- nate on the upper side of each lobe in a slit-like concav- ity. Apically, the sperm groove is shallow and is ac- companied by weakly developed sulcal lips. The sperm groove gets deeper and widens towards the proximal part of the truncus. The epidermal tissue of the lobes appears to be rough, although no calyces can be distin- guished. Colour of preserved holotype. Upper side of head and body dark grey. Head grey-brown, with darker vermicu- Fig. 3: Right hemipenis of the holotype of Gekko scientiad- ventura sp. n. (ZFMK 76198): a) sulcal, b) asulcal view. Dra- wing: H. ROSLER. lations and flecks, additionally small light dots. Dorsum with seven median light flecks that continue laterally into small transverse bands with strongly waved anterior and posterior margins. These transverse bands are ante- riorly bordered by brown-blackish lines and dissolve laterally. Between the single bands light flecks and nu- merous small spots are interspersed. Limbs above dark grey with grey-brown stripes. Throat, chest and belly buff, the throat bearing a reticulate pattern. Belly later- ally with light spots. Forepart of tail above and below dark grey, hindpart grey-black. Nine grey-brown trans- verse bands on the upper side of tail, the hindmost five reaching the lower side of the tail and nearly form closed rings. 3.1.4. Paratypes. ZFMK 76174 (Fig. 4), adult female, ZFMK 76175-179, juveniles, Phong Nha - Ke Bang, surroundings of 17.32 N, 106.16 E, 50-150 m a.s.l., Quang Binh Province, Vietnam, coll. H.-W. HERRMANN, VU NGOC THANH & T. ZIEGLER, 26 August - 1 September 2001. ZFMK 80651-652, adult females, same locality data, coll. VU NGOC THANH & T. ZIEGLER, 29 August - 10 September 2003. Paratypes ZFMK 76179 and 80652 will be permanently transferred to the scientific collection of the Vietnamese National University, Hanoi (HNUV). 3.1.5. Description of paratypes. Individual body meas- urements and proportions can be seen in Table 1. For the latter, the paratypes have the following mean values and standard deviations (n being variable): SVL:TL 0.87+/-0.07, SVL:HL 3.49+/-0.24, HL:HW 1.28+/-0.03, HL:HH 2:28+/-0.03, SE:EE 1.33+/20:09: All paratypes possess invariably 3 nasals, no internasals and 2 postmentals. Further, females and juveniles are lacking preanal pores. The postmentals are differently shaped, the maximal relative length is the left post- mental of ZFMK 76176 which is three times as long as 138 Bonner zoologische Beitráge 53 (2004) Fig. 4: Live female paratype of Gekko scientiadventura sp. n. (ZFMK 76174). Phot. T. ZIEGLER Fig. 5: Portrait of one of the captive males of Gekko scien- tiadventura sp. n. (see also fig. 28 in ZIEGLER et al. 2004) from 22 April 2003, measuring a SVL of 67 mm and a TL of 70 mm at the time; note the spiny posterior ciliaries as well as the bordering nasorostrals. Phot. T. ZIEGLER broad. The right postmental is divided in ZFMK 76176 and 76178. The posterior ciliaries are spiny in all para- types. The remaining scalation characters vary to a small degree only (see Table 2). Range (mean and stan- dard deviation) of the number of supralabials is 12-14 (13.00+/-0.82), of the sublabials 9-13 (9.94+/-1.26), of the scales behind postmentals 5-7 (5.67+/-0.71), of the interorbitals 41-49 (43.75+/-3.59), of the ventrals 38-48 (41.80+/-3.90), of the scales around midbody 118-139 (132.20+/-8.23), of the lamellae under the Ist toe 12-15 (13.86+/-1.03), of the lamellae under 4th toe 14-17 (15.64+/-1.15), of the postanal tubercles 2-3 (2.50+/- 0.53) and of the number of dorsal scales in the 3rd tail whorl 10-11 (10.29+/-0.49). All juvenile paratypes (as well as two embryos, see 3.1.6.) have still two egg teeth which touch each other apically as it is typical for geckos with relative thick and calcified eggshells (ROSLER 2001b). The tooth crown of the egg teeth has a shape similar to that of G. gecko (see SLUITER 1893). Colour of preserved paratypes. The pattern of the pre- served material corresponds to that of the living ani- mals, the overall colouration being more greyish. 3.1.6. Other material. Next to the type series, there are two male embryos (ZFMK 76199 and CPHR 2028) and three eggshells (ZFMK 76180) from the same locality. The embryonic males exibit egg teeth as decribed above, their hemipenes are completely everted, deeply forked and show two subequal apical lobes. In one of them (CPHR 8028) the dark pigments (6 crossbands) start to develop. In addition there are three males and one female (Figs. 5-7, see also fig. 28 in ZIEGLER et al. 2004) that had hatched in September/October 2001 from eggs found in a karst cave at the type locality in August 2001, and which are kept alive in the Cologne Zoo. 3.1.7. Colouration in life. Variable in all adults. Basic colour either more brownish marbled with yellowish flecks, or yellowish with brownish marbling. Upper side of head with large, blackish-brown flecks or spots which may be present also on the snout. Some speci- mens additionally possess some greyish flecks on the same head regions. Ciliary scales black with yellow margins. Iris metallic green-yellowish, pupil with bronze-coloured margin. A row of 7 middorsal grey flecks which may be extended to wavy crossbands of the same colour towards the flanks. These crossbands may be dissolved into flecks. The middorsal flecks are anteriorly bordered with a blackish brown pattern that is w-shaped in the neck region. The grey crossbars on the tail are likewise anteriorly bordered by blackish brown pattern elements. The grey flecks on the limbs are big- ger on the hind legs. Throat and belly with more or less distinct lemon to buff colouration, with more or less dis- tinct brown-olive marbling. Colour pattern of underside of tail variable, with pale or dark grey to ochre or black- ish brown flecks. H. ROSLER, T. ZIEGLER, VU N.T., H.-W. HERRMANN & W. BOHME: A New Gekko Species from Vietnam Fig. 6: Gular region of the male of Gekko scientiadventura sp. n. depicted in Fig. 5 (10 November 2003). Phot. T. ZIEGLER Fig. 7: Captive female of Gekko scientiadventura sp. n. (10 November 2003); note the regenerated tail. Phot. T. ZIEGLER 3.1.8. Etymology. The species name is a patronym for a scientific magazine programme of the German Televi- sion channel ZDF called “Abenteuer Wissen” (adven- ture of knowledge) in order to acknowledge the first documentation of the Phong Nha - Ke Bang National Park biodiversity for the German public. First live pic- tures of Gekko scientiadventura sp. n. were shown on 139 this TV programme. The name is derived from the Latin words “scientia” (knowledge, science) and “adventura” (literally = things that will come = adventure) and is a noun in apposition. 3.2. Distribution So far, Gekko scientiadventura sp. n. is known only from the type locality (Fig. 8). Although currently re- garded as endemic for Vietnam, its occurrence in Laos cannot be excluded, considering the proximity of the Laotian border from the type locality. CHINA VIETNAM Hanoi ® South China Sea CAMBODIA Fig. 8: Position of the type locality within Quang Binh Prov- ince. 3.3. Natural history All specimens of the type series were collected at night in primary forest, either directly on or in the immediate neighbourhood of the karst limestone outcrops that were partly overgrown with vegetation (Fig. 9). The geckos were mostly seen on the vegetation in low height (1.0- 2.5 m) rather than on the bare rocks. In contrast, these bare rocks were mostly populated by the syntopic and only recently described bent-toed gecko Cyrtodactylus phongnhakebangensis which, in turn, tended to avoid 140 Bonner zoologische Beitráge 53 (2004) the vegetational parts of the microhabitat (ZIEGLER et al. 2002). Fig. 9: Habitat of Gekko scientiadventura sp. n. near the type locality. Phot. T. ZIEGLER Apart from the voucher material of our type series no further sightings of Gekko scientiadventura sp. n. were made, with the exception of a mass egg-laying site in a dark, manifold rock crevice system at the type locality. Fig. 10 shows this site (in about 2.5 m height) at the be- ginning of the rainy season (end of August) in 2001. The four geckos cited under “Other material” originate from these eggs; they hatched in September/October 2001 which correlated with the beginning rains. One year later, in August 2003, we found two glued eggs just at this mass egg-laying site. In addition, two further communal egg-laying places were found in this cave system which seemed to be used also already since a longer period; they contained, however, much less eggs and were situated much higher (ca. 5 m high). On August 30, 2003, we measured at the lower egg- laying site, at 14.30 h, a relative humidity of 85% and a temperature of 27.3 °C. At 21.40 h the same day the temperature was 26.3 °C and the relative humidity had Fig. 10: Mass egg-laying site of Gekko scientiadventura sp. n. in a karst rock crevice at the type locality. Phot. T. ZIEGLER increased to 93% (with a recorded minimum of 70% be- tween 14.30 and 21.40 h). In the primary forest, outside the rock crevice system, we measured with a minimum- maximum thermometer on August 28, 2001 24.4-30.2 °C and one day later 24.3-27.8 °C. For further (long- term) climatic data from this area see HERRMANN et al. (2002). One of the authors (TZ) was able to record some behav- ioural and autecological observations of the captive- raised geckos: The growling calls of Gekko scientiad- ventura sp. n. were mostly uttered during dawn and night, but could also be heard in summer afternoons (e.g. 22-26 July, 2002), 1.e. during the light phase of the terrarium. Obviously, also the black-and-white annula- tion of the tail plays a role in the infraspecific commu- nication between both sexes, because in an interacting couple the tails were raised and displayed with undulat- ing movements. A male that was taken out of its terrar- ium used to wag its tail in a much faster manner. Once an acinetic reaction could be observed, a lethisimulation with a recurved tail and extremities that were raised from the ground and standing off from the body. The geckos were fed mainly with crickets of various size classes that were powdered with a vitamin-mineral mix- ture, however, they were repeatedly observed to lick on and even to bite off small pieces of bananas. In 2003, two egg-layings occured but the paired, partly deformed clutches proved to be unfertilised. 3.4. Comparison with other Vietnamese Gekko species Next to the lack of dorsal tubercles, Gekko scientiadven- tura sp. n. can be distinguished from its Vietnamese congeners by the combination of the characters size, number of internasals and preanal pores as well as dor- sal pattern: G. badenii reaches a SVL up to 76.5 mm and has 1-3 in- ternasals; males have 14-18 preanal pores, and the dor- sal pattern consists of 4-8 very narrow and sometimes 142 Bonner zoologische Beitráge 53 (2004) 23 21 subdigital lamellae under 4th toe. Males with 22-23 preanal and femoral pores; subcaudals transversely broadened (BROWN & ALCALA 1962, 1978). Fig. 11: Dorsal view of the holotype of Gekko subpalmatus (BMNH 1946.8.2592). Phot. T. ZIEGLER G. subpalmatus reaches a SVL of 78 mm and has 1-2 internasals. Toes basally narrowly webbed. 7-10 sub- digital lamellae under 4th toe. Males with 5-11 preanal pores. Subcaudals transversely broadened (GUNTHER 1864; POPE 1935; ZHOU et al. 1989; ZHAO et al. 1999). For a better comparison with G. scientiadventura sp. n. we studied the holotype of Gekko subpalmatus (BMNH 1946.8.2592) and the lectotype and the paralectotype of Gekko melli Vogt, 1922 (ZMB 27659 A & B) which is considered to be synonymous with G. subpalmatus (e.g. ZHAO & ADLER 1993) (Figs. 11-13): The rostral of the holotype of G. subpalmatus has a short upper transverse suture, the posterior ciliaries are pointed, and there are five distinct tubercles anterior of the tympanum on the right side of the head. The dorsum is lilac-grey to violet- grey, a light postocular stripe and four longish grey- brown vertebral flecks are barely visible, and the tail has several broad light transverse bands that are bordered by darker pigment. The lectotype and the paralectotype of G. melli lack a rostral suture, the posterior ciliaries are likewise pointed, and pretympanic tubercles are lacking. The dorsum of ZMB 27659 A is olive-brown. There is a dark naso-ocular and a light postocular stripe. Two dark stripes are running from the ear openings to the axillae, and the shoulder region shows two sepia-coloured stripes. The dorsum has three broad, irregular and partly confluent, grey-brown bands with sepia-coloured mar- gins, the tail has four broad pale brown bands that are likewise bordered by darker pigment. The regenerated tail tip has fine darker dots. Apart from their colour pat- tern, the types of both taxa differ also in size and in body proportions (SVL/TL index) and in the number of interorbitals, subdigital lamellae under the 4th toe, and scale rows in the third caudal whorl (see Tab. 3). Pend- ing future studies on more material we consider G. melli therefore as valid. Fig. 12: Portrait of the holotype of Gekko subpalmatus (BMNH 1946.8.2592). Phot. T. ZIEGLER Fig. 13: Dorsal view of the lectotype of Gekko melli (ZMB 27659 A). Phot. H. ROSLER G. tawaensis reaches a SVL up to 70 mm and has inter- nasals. The toes are basally narrowly webbed, and there are 12 subdigital lamellae under the 4th toe. The males have no preanofemoral pores, and the subcaudals are anteriorly mesially divided (OKADA 1956; SENGOKU 1989; UTSUNOMIYA et al. 1996). G. scientiadventura sp. n. differs from the other Gekko species without dorsal tubercles by the combination of following characters: (a) SVL, (b) internasals, (c) exten- sion of webbing between toes, (d) subdigital lamellae under 4th toe, (e) number of preanal pores, and (f) shape of subcaudals. G. scientiadventura sp. n. differs thus from G. athymus in the characters a, b, d, e; from G. subpalmatus in the characters a, b, c, d; and from G. tawaensis in the characters b, e, and f. 4. DISCUSSION The genus Gekko Laurenti, 1768 contains, according to the last overview by KLUGE (2001) 28 species. We add to this number five more species which we consider as valid: G. liboensis Zhou & Li, 1982, G. melli Vogt, 1922, G. mindorensis Taylor, 1919 G. scabridus Liu & Zhou, 1982, and G. scientiadventura sp. n. Alexander SCHINTLMEISTER: Peridea clasnaumanni spec. nov. (Lepidoptera: Notodontidae) aus China 167 >" ee 6 Abb. 1-4: Imagines Peridea. (1) Peridea clasnaumanni — 3, China, NE. Jiangxi, Wuyi Shan, 1600 m, 50 km SE Yingtan, 27°56’N, 117°25’E, April 2002, Holotype. (2) Peridea clasnaumanni — Y, China, Hunan, Cili, 3.ix.1988, Paratype. (3) Peridea aliena — 3, Russia, Primorye, 30 km NE Wladiwostok, 24.vii.1963. (4) Peridea hoenei — Y, China, Sichuan, Daxue Shan, 40 km W Mianning, 2750 m, 7.-8.vii.1998. Abb. 5-6: Genitalapparate Peridea. (5) Peridea clasnaumanni — &, China, Jiangxi, Wuyi Shan, Xipaihe, 27°54’N, 117° 20’E, Ju- ly 2003, 1500-m (GU 79-98), Paratype. (6) Peridea aliena — Ö, N. Korea, Myohang san, Myohang-Shon-valley, 24.-30.vi.1985 (GU 24-73). ul I Ñ 1 + ' Ki ’ ' i La, ot ihe 292 Bonner zoologische Beiträge 53 (2004) Figs. 1-4: SS aedeagus-structure as found in Ambia locuples and the new species seems not advisable. We consider Ambia thyridialis (Lederer, 1855) (fig. 3), a species recorded from the Lebanon and Egypt which also has no saccular tooth as a closely related species. 2. DESCRIPTION OF SPECIES Ambia naumanni Speidel & Stiining, sp. nov. (Fig. 1, 2) Holotype Ö: “Li-kiang. (China). Yuennan. 8. 8. 1934. H. Hóne” Provinz Nord- Paratypes: Males: dto. 1 4 13. 6. 1934; 1 & 19. 7. 1934; MGW 2 81935" 1 S172 719342 1.8.7.8. 193422 SS. 1935; 1 $ 12. 7. 1934: 1 8 7.7.1934, GS-15036 . Ambia naumanni sp. nov., holotype, male (Lijiang, 8. viii. 1934) (wing span 15 mm). . Ambia naumanni sp. nov., paratype, male (Lijiang, 13. vi. 1934) (wing span 16 mm). . Ambia thyridialis (Lederer, 1855), male (Ghazir) (wing span 15 mm). . Ambia naumanni sp. nov., male paratype, lateral view of head. &-SB; 1.6 17.7. 1934; GU 137 & SP; 1 4 21. 7. 1934, Prep. 15378 SB and FU 160 SP; 27. 6. 1934, 1 Y 24. 6. 1935 with genitalia in glycerol; Females: 1 Y 3. 8. 1935; 1 9 8. 9. 1935; 2 9 3. 7. 1934 with FU 159 SP; 1 Y 29. 6. 1934, GU 15063; 1 Y 15. 6. 1935; 1 Y “Li-kiang ca. 2000 m, Prov. Nord-Yuennan, 26. 6. 1934. H. Höne” with genitalia in glycerol. Diagnosis (fig. 1, 2).- Head and body: Antenna com- paratively stout in the males, pubescent. Labial palpi rather long, upcurved, terminal segment almost reaching base of antennae. Frons smoothly scaled, whitish like the palpi. Thorax and abdomen greyish-brown, mixed with white, abdominal tergites with posterior white tranverse bands. Wolfgang SPEIDEL & Dieter STUNING: Ambia naumanni sp. n., a New Species of Musotiminae from Yunnan 223 {mm 5 E | Fig. 5: Ambia naumanni sp. nov., male genitalia. Wings: Wingspan 15-17 mm, forewing length 7,5-8 mm. Wings with termen sinuous as usual in the genus, apices produced. Ground colour yellowish. Transverse fasciae prominent, white, bordered blackish-brown. The basal fascia strongly oblique, broken, the proximal one slightly curved, the distal one outcurved anteriorly and strongly retracted below middle, reaching the inner margin at about one half. The latter fascia is widened to a quadrangular spot at inner margin. Submarginal fascia conspicuous, widening towards tornus. Medial area with a white disco-cellular spot and another larger white spot in the anterior angle of the distal fascia. Marginal area yellow. Hindwing with similar pattern. Proximal fascia wider than in forewing, submarginal fascia strongly widening towards apex, incised at vein M2. Medial area suffused with brownish scales, like in the forewing, with two large white spots at the costal region and a small one at inner margin. Marginal area yellow. The venation (fig. 7) shows no principle difference to the other mem- bers of the genus. 1mm 6 Fig. 6: Ambia naumanni sp. nov., female genitalia. Male genitalia (Fig. 5): Vinculum large, with u-shaped saccus. Tegumen short, uncus with a broadly triangular base, otherwise slender, distinctly flattened dorso- ventrally, curved ventrad, pointed. Gnathos with strong lateral arms, medial process very narrow, slightly curved parallel to the uncus, minutely dentate dorsally, pointed at tip. Juxta large, with two dorsal spines situ- ated laterally at the posterior margin. Valva simple, broadening towards apex, without a saccular spine or other appendages, slightly ciliate. Aedeagus with a slender, sharply bent process, arising just distally to the opening of the ductus ejaculatorius. Shaft of aedeagus strongly incurved beneath this process. A curved ap- pendage originates at the inner side of the sinus. Female genitalia (Fig. 6): Ovipositor comparatively short, with moderately long apophyses. Ostium wide, with a bilobed sclerotized structure inside. Upper third of the ductus bursae sclerotized, with the ductus semi- nalis originating in the center of this region. Ductus bur- sae slightly broadening towards corpus, which is small and with a trace of a signum only. 224 Fig. 7: Ambia naumanni sp. nov., wing venation. HOPE ENT. COLL OXF. UNIY. = a GENITA TEC HOPE ENT. COLL. OXF. UNIY. MUS. NO.: 1300-93 J.Minet Fer 542, a Prep 542 24 549 b HOPE ENT. COLL. OXF. UNIV. MUS. TYPE LEP. ‘Me 90 mbia . 7 fol l, MAT HOPE DEPT.OXFORD Fig. 8: Ambia ptolycusalis Walker, 1859. Hole Sri labels (Sarawak, Borneo). The new species is very similar to the Western Palearc- tic Ambia thyridialis Lederer, which can easily be dis- tinguished by the more brownish ground colour of the wing surface, with yellowish scales only found in the marginal fasciae. Ambia thyridialis Lederer is also simi- lar in the male genitalia, lacking the saccular process of the valvae as well, but the aedeagus is completely dif- ferent. Bonner zoologische Beiträge 53 (2004) os 9 Fig. 9: Ambia ptolvcusalis Walker, 1859. Male genitalia of holotype. Distribution and bionomics: Hitherto only known from the type-locality in SW. China, Yunnan, Yueling- shan near Lijiang, at elevations between 2800 and 3500 m. Flight-period from mid of June until beginning of September. First instars and foodplant unknown, but probably fern-feeding, as most of the Musotiminae are. Acknowledgments and etymology. The new species is named in honour of Prof. Dr. C. M. NAUMANN who im- proved the situation in the entomological department of our museum considerably during his directorship and facili- tated our work in many respects. We are grateful to Prof. Dr. K. Efetof and V. Saenko for the drawings of the genita- lia and to D. J. Mann (Hope Entomological Collections, Oxford) for the loan of the type slides of Ambia ptoly- cusalis. REFERENCES LANGE, W. H. (1956): A generic revision of the aquatic moths of North America (Lepidoptera: Pyralidae, Nymphulinae). Wasmann Journal of Biology 14(1): 59-144. MUNROE, E. & SOLIS, M. A. (1999): The Pyraloidea. pp. 233-256 in KRISTENSEN, N. P. (ed.): Lepidoptera, Moths and Butterflies. 1. Evolution, Systematics, and Biogeography. In FISCHER, M. (Ed.): Handbuch der Zoologie 4. Arthropoda: Insecta (35). Berlin, New York, 491 pp. 228 Bonner zoologische Beitráge 53 (2004) 0,5 cm Fig. 1: Paracymoriza naumanniella sp. n. Holotype (male) (A) and one Paratype (female) (B). Indonesia, North Lombok, Gu- nung Rinjani NP, Air terjun, near village Senaru, ca. 500 - 600 m, 08°18' S / 116°24' E, 28. Dezember 2003, TF, leg. Mei-Yu Chen é Ulf Buchsbaum. Fig. 2a-b: Paratypes of the new species still alive at the type locality in Lombok (see above). Photo: Ulf Buchsbaum. Y genitalia (Fig. 4): Ovipositor short, with long, slen- der, fragile apophyses. Ductus bursae rather long, gradually widening towards corpus bursae, with an un- usual evagination about in its middle. Corpus bursae round, ball-shaped, without a clear signum. Colliculum situated terminally, with the slender ductus seminalis originating inside and in a short distance of the colli- culum. Ostial region strongly sclerotized, large, cup- shaped, narrowing strongly towards the ductus bursae. Egg (Fig. 7a, b): Oval, with micropylar zone at the nar- row end of the egg (size: 310 x 180 pm). Distribution (maps, Figs. 5a and 5b): Hitherto only known from the type locality in Lombok. Genetic data: We sequenced the mitochondrial gene cytochrome oxidase subunit I in order to provide a mo- lecular dataset for subsequent re-identification of the new species as recommended by TAUTZ et al. (2003). This gene has proven its usefulness for DNA taxonomy purposes in many studies (e.g. HEBERT et al. 2003). Genetic characterization: Method: DNA was extracted from abdominal tissue of some of the type specimens of P. naumanniella sp. nov. using Qiagen (Hilden, Ger- many) tissue kit according to the protocol of KNOLKE et al. (2005). Mitochondrial (mtDNA) cytochrome oxi- dase subunit I (COI) gene was amplified with PCR us- ing protocols and primers as in SIMON et al. (1994). Di- rect sequencing of dye labelled templates was carried Wolfgang SPEIDEL, Ulf BUCHSBAUM & Michael A. MILLER: A New Paracymoriza Species from Lombok (Indonesia) 231 A Makassars Ya [«SUNDA-INSELN Javasee GROSSE S UND A - 1 .NOSSERESN PAZIFISCHER OZEAN 4 MAL I lS Cn gees ee 4 ARCHTPEL Lombok > k Le ¡Flores Bali Sumbawa INDISCHER OZEAN I Sorongjukung dra £ A - INDISCHER OZEAN Fig 5: Type locality. 5 a: Enlarged view to show the situation of the Island Lombok (Indonesia) east of Bali; 5 b: type locality of P. naumanniella on Lombok Island near the village Senaru below the mountain Gunung Rinjanı. bo i>) hn Bonner zoologische Beitráge 53 (2004) ¿PA PS UR Fig. 6: Biotope at type locality beside the waterfall. 6 a: Here, most moths rest under the rocks near the water surface. Photo: Buchsbaum; 6 b: One of the authors (U. B.) collecting specimens of the new species. Photo: Mei-Yu Chen. Ulf wit? 236 Bonner zoologische Beitráge 53 (2004) ” e i yo z ‘ i ' , , a ‘ Be 1 Bonner zoologische Beitráge Instructions to Authors Bonner zoologische Beitráge publishes original papers, reviews, and scientific notes covering the whole field of zoology, with em- phasis on biodiversity, biogeography, history of zoology, mor- phology, phylogenetics, and taxonomy. The material must not be submitted for publication elsewhere. All manuscripts will be peer- refereed. Final acceptance of items for publication is the responsi- bility of the corresponding editor dealing with the respective taxon (see list of editors). Manuscripts not compatible with the following guidelines will be returned. Manuscript preparation: Manuscripts should be written in English or German. If you plan to submit a manuscript of more than 80 manuscript pages, please contact the editor-in-chief in advance. 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Box 50007, SE-104 05 Stockholm, E-mail: sven.kullander(@nrm.se Prof Dr. Steven PERRY, Rhemische Friedrich-Wilhelms- Universitat, Institut für Zoologie, Poppelsdorfer Schloss, D-53115 Bonn, Tel. +49 228-73 3807, E-mail: perry(@uni-bonn.de Dr. Wolfgang SCHAWALLER, Staatliches Museum für Naturkunde, Rosenstein 1, D-70191 Stuttgart, Germany, Tel. +49 711-8936 221, Fax +49 711-8936 100, E-mail: schawaller.smns(@naturkundemuseum-bw.de Dr. W. David Sissom. Dept. of Life, Earth and Environ- mental Sciences, W. Texas A. & M. University, WTAMU Box 60808, Canyon, Texas 79016, USA, E-mail: dsissom(@wtamu.edu/ Dr. Miguel VENCES, University of Amsterdam, Zoological Museum. Mauritskade 61, PO Box 94766, NL-1090 GT Amsterdam, The Netherlands, Tel. +31 20-525 7319, E-mail: vences(@science.uva.nl PD Dr. Heike WAGELE, Rheinische Friedrich-Wilhelms- Universitat. Institut für Evolutionsbiologie und Okologie, D-53121 Bonn, Tel.: +49 228-73 5159, Fax: +49 234-322 4114, E-mail: hwaegele(@evolution.uni-bonn.de Dr. Erich WEBER, Eberhard-Karls-Universitat, Zoologische Schausammlung, Sigwartstr. 3, D-72076 Tubingen, Germany, E-mail: erich.weber(@uni-tuebingen.de Bonner zoologische Beitráge Band 54 (2005) Heft 1 Seiten 1-2 | Bonn, April 2006 Andreas Helbig (28.07.1957 — 19.10.2005) Prof. Dr. Andreas Helbig, member of the advisory board of Bonner zoologische Beitráge, died on 19 October 2005, at the age of only 48. He leaves a wife and two sons. His early death is a great loss not only to his fam- ily but to the scientific community, as he was a leading figure in avian science in Europe. ai ie Andreas Helbig was an ornithologist with heart and soul. His interests in birdlife were broad, extending from bird-watching to molecular science and taxonomy, comprising field ornithology, bird migration, orientation and, more recently, molecular phylogeny and population genetics of birds. His merits are in his molecular phy- logenetic publications, especially on Sylvia warblers, but also on birds of prey, shearwaters, gulls and other bird groups. One of his last projects was: The Herring Gull complex as a model group for recent holarctic ver- tebrate radiations. His interests in birds date back to his school days in Enger, Kreis Herford, where Andreas Helbig published his first papers on observations of rare birds. From 1979 onwards he studied Biology at the University of Biele- feld, San Diego State University in California and re- ceived his Diploma on “Zugrichtung tagstiber ziehender Vögel am Teutoburger Wald, NW-Deutschland” [Mi- gration routes of diurnal travelling birds in the Teuto- burg woods, NW-Germany] at the J.W. Goethe- University in Frankfurt in 1983. After a break due to his alternative civilian service in the “Europareservat Dümmer” he continued his studies in Frankfurt. His dis- sertation on “Angeborene Zugrichtungen nachts ziehender Singvógel: Orientierungsmechanismen, geographische Variation und Vererbung” [Inborn migration routes of E night travelling passerines: orientation mechanisms, geographic variation and inheritance] was awarded best dissertation of the yed 1989. In order-to- learn tech- niques and methods f& the application of molecular markers he spent several months at Queens University in Kingston (Canada), funded by the Konrad Adenauer Foundation. For a short period Andreas Helbig contin- ued his studies on inborn orientation mechanisms in the Blackcap, Sylvia atricapilla, as a Postdoctoral Fellow at the Max Planck Institute for Ornithology, Radolfzell. Until 1993 he held a Postdoctoral position at the Insti- tute of Pharmaceutical Biology, Heidelberg, financed by the German Research Foundation (DFG). From this pe- riod onwards Andreas Helbig focussed his research on molecular phylogenetics. Ornithological systematics, species concepts and those species-complexes at the transition between subspecies and species attracted his special interest. From July 1993 until his sudden and unexpected death he held the position as director of the “Vogelwarte Hiddensee” [Bird ringing center Hidden- see] at the Zoological Institute of the Ernst-Moritz- Arndt University of Greifswald. Besides his activities at the ringing station he taught “Evolution and Phylogenet- ics” and “Ornithology”. With his habilitation on “Mole- kulare Phylogenie, Systematik und Evolution der Zweig- sänger (Aves, Sylviidae)” [Molecular phylogeny, systematcs and evolution of Old World Warblers (Aves, Sylviidae] in 1997 he advanced to the leading expert on avian molecular systematics over broad areas of Europe. The German Ornithologists’ Society (DO-G) awarded his research with the “Stresemann-Fórderung” in the same year. Besides his duties as director of the Institute and teacher, Andreas Helbig was active in several other fields: He was also chairman of the Scientific Commit- tee of the DO-G, Secretary General of the European Or- nithologists’ Union for five years, joined the TSC (= Taxonomic Sub-committee (TSC) of the British Orni- thologists’ Union, BOU), and was well-known as a member of the German Rarities Committee for about 20 years. As editor, subject editor or member of the edito- rial board, he contributed to the scientific quality of sev- eral journals; e.g., “Vogelwelt — Beitráge zur Vo- gelkunde”, “Journal of Ornithology“, “Berichte aus der Vogelwarte Hiddensee”, “Limicola”, and “Journal of Evolutionary Biology”. Andreas Helbig also kept contact with our Research Museum. He was on the advisory board of our institu- 2 Bonner zoologische Beitráge 54 (2005) tional journal “Bonner zoologische Beitráge”. He was invited speaker at the Second European Symposium on Bird Collections in November 2001 and at the 44" Phy- logenetisches Symposium on adaptive radiations in 2002. The presentation of his research project on radia- tion in the Herring Gull complex (Larus argentatus- complex) by his wife, Dorit Liebers-Helbig, had to be delayed until February 2006. I still remember the circumstances when I met Andreas Helbig for the first time. It was during the Pan African Ornithological Congress in Nairobi 1988, when every- body was spending his or her spare time in Nairobi National Park for bird-watching. Provided with the in- timate knowledge of local ornithologists where to spot rare birds, we easily arrived at the best places — always to find Andreas Helbig already there. This episode ex- emplifies his attitude towards all fields of ornithology — his alert interest in any actual and novel event con- cerning birds, as well as his competency of recog- nizing these. His passing leaves a great gap in avian science. Renate VAN DEN ELZEN, Zoologisches Forschungsmu- seum Alexander Koenig, Bonn (r.elzen.zfmk@uni- bonn.de) | Bonner zoologische Beitráge Band 54 (2005) Heft 1 | Seiten 3—23 Bonn, April 2006 Butterfly Diversity and Protection in Turkey Sigbert WAGENER' Koblenz, Germany Abstract. The aim of this paper is to demonstrate the outstanding diversity, the endemic and rare taxa, to present a model of evaluation of sites to be protected and to give some recommendations for the protection of butterflies in Tur- key. Key words. Butterfly endemism, rare butterflies, priority sites, protection management. 1. INTRODUCTION What should be protected? — In many countries legal prescriptions exist for the protection of plants and ani- mals. As a rule, all measures taken are directed at the conservation of individuals of a single species. Nine of the butterfly species occurring in Turkey are named in Annex II and Annex IV of the Habitats Directive and the Bern Convention. Background information on these species and their threat situation in Europe is given in VAN HELSDINGEN et al. (1996) and WAGENER (1996). The author presented data on the threat and conservation status for all butterfly species occurring in Turkey in the Red Data Book of European Butterflies (VAN SWAAY & WARREN 1999). In spite of all efforts, the number of butterfly popula- tions incessantly decreases. This experience is depress- ing for any lepidopterist working in the field, and asks for new ways. To protect individuals of mammals and birds with a very low rate of increase surely is worthwhile, but to protect individuals of insects such as butterflies with a generally high maximum rate of fertility is scientifically unjustified and is practically not workable. By nature, insects serve as food for many predators like birds, liz- ards, and spiders. In an undisturbed ecosystem, a butter- fly population usually shows only limited annual undu- lations in their density, because the population is able to compensate such losses through predators and even through unfavourable weather conditions during their life cycle. But such undisturbed habitats become rare more and more by human activities, in Turkey, and elsewhere in Europe and all over the world. The device must call: Protect the populations and their habitats be- fore the last legally protected species has disappeared! That is in favour not only of butterflies but also of other animals and plants, too. If one wants to protect a butterfly species one has at first to know if and where it occurs at all. Secondly one must know the biology of the species and their environmental requirements such as special climatic factors, food plants for the larvae and the butterflies, special habitat and vegetation structure and others. Thirdly, one must know what exactly threatens the existence of the species by man’s activities, which disturb the balance of the ecosystem of which the species is a part of (cfr. WAR- REN 1992). This paper attempts to answer the first question for Tur- key. Satisfying answers to the second and third ques- tions can be given presently for only a very few species. A lot of scientific research work of that kind that TULU- HAN (1998, fig. 2) has shown in his “flowchart of an ef- fective environmental management” has still to come for many years. The only effective protection of Turkish butterfly taxa right away is therefore the conservation of the sites in which they are still found today. The Tables and Maps herein show where to look for such sites. “Diversity”, “endemisms” and “rarity” demonstrate this each in a somewhat different way but the final result is almost the same related to a certain UTM 10 km square. 2. DIVERSITY OF BUTTERFLY TAXA IN TURKEY Species diversity is generally higher in Turkey than in any other country of Europe or the Near East, in plants as well as in animals, not only in butterflies and moths but also in other orders of insects. The first butterflies from Turkey were described by CRAMER (1775) and HERBST (1798) in the late eight- eenth century. Systematic research started in the early nineteenth century, mainly by Hungarian, Austrian and German collectors. STAUDINGER (1878-1881) already could list 199 butterfly species from about 30 localities. In the last time, especially since 1970, the exploration was strongly intensifyed by Turkish and European lepi- dopterists so that today 369 species or, including 79 subspecies, 448 taxa of butterflies are known from more than 2300 localities. In Appendix 1, all recognized spe- cies and subspecies of butterflies occurring in Turkey 4 Bonner zoologische Beitráge 54 (2005) ascertained up to 2003 are listed and shows the number of UTM 10 km squares in which that taxon occurs. The list mainly is based on HESSELBARTH et al. (1995) and represents through corrections and additions the latest stand of faunistic, taxonomic and nomenclatural knowl- edge. The species shared between families of butterflies are shown in Table 1. In Turkey, the family Lycaenidae has the highest number of species and the highest number of endemic species or subspecies. Almost every year new taxa are discovered whilst the number of species and subspecies in the other families now seems to be stable. The taxonomic status and the distribution of some de- scribed taxa, especially in the genus Polyommatus (sub- genus Agrodiaetus) is still uncertain. Many data for spe- cies of Agrodiaetus therefore could not be used in this study. The different forms of Hyponephele lycaon are treated here as a single lycaon-complex, and also Lep- tidea sinapis and L. reali as a single sinapis-complex. To settle these open questions still more research is ur- gently needed. Table 1. Distribution of species among the families of butter- flies in Turkey Family Species Subspecies Total Hesperiidae 4] 5 46 Papilionidae 12 8 20 Pieridae 36 9 45 Lycaenidae 150 26 176 Nymphalidae 125 28 153 (Satyrinae (72) (23) (95) 369 species 79 subspecies 448 taxa The species Catopsilia florella and Hypolimnas missip- pus cannot be deamed as stable elements of the Turkish fauna. They are very rare accidental immigrants from Arabia and are not taken into account in the further con- siderations. To demonstrate the extraordinarily high butterfly taxa diversity in Turkey, Map 1 shows the number of spe- cies for every UTM 10 km square. The squares are enumerated according to their UTM alphanumerical codes from West to East according to the Tactical Pi- lotage Chart. Data for the time between January 1951 and December 2002 were at the author’s disposal for 1846 UTM 10 km squares out of 7998 possible ones for Turkey. These data were taken from HESSELBARTH et al. (1995), the data bank for Turkish Butterflies held by Harry van OORSCHOT and publications since 1994 (see references) as well as from a list of observations between 1995 and 2001 sent to the author by Lutz Lehmann. On the basis'of these data the distribution maps in HES- SELBARTH et al. (1995) have been made topical. The numbers of taxa per 10 km UTM square have been gained by means of a line list. The resulted numbers from 1 up to 170 taxa/square are given in column 3 of the Appendix 2'. To make the map comprehensible the numbers were divided into 10 grades, each of 17 taxa. A certain colour has been assigned to each grade (see “Colour scale for number of taxa” at top of Map 1). White marks squares without any record. By using this scale, the colour grade 1 to 10 can be deemed as a “di- versity weight” in favour of the corresponding square. The squares from which a butterfly taxon is known and the number of taxa observed within one square is not evenly distributed over the country for several reasons: ° The recording level is with 23 % of possible squares moderate. From many squares only one or two spe- cies are known, the highest species number in one square is 170 (South side of Ovit Gecidi, Prov. Erzu- rum). ° The single parts of the country are very different in their natural provision by orography, climate, soil, vegetation, land use and other factors so that only species adjusted to special conditions can survive in certain sites or are, owing to geohistorical processes, restricted geographically to defined regions. + Large parts of mountainous regions are hardly ac- cessible, others suffer through overgrazing, other parts, formerly steppe land, have been converted into large monotonous wheat fields during the last dec- ades, again other sites became lost through artificial lakes, intensification of agriculture or building espe- cially around the large cities of Istanbul, Bursa, Íz- mir, Konya, Adana and along the Mediterranean and Marmara coasts. Therefore, large white spots in the map do not necessarily mean that these areas have not been expiored. In fact, there are large areas in which one may hardly see any butterfly at all. The number of such sites increases from year to year. + Usually researchers, going by car, collected butter- flies mostly by the wayside, rarely 1 to 2 kilometres from a road. Sites further away, accessible only by walking or riding have been mostly neglected. Many squares have been visited only once and shortly, other places repeatedly and at different times for several days. Success of collecting depends often on the weather and to come across just with the right flying period. Large regions in the Southeast of Ana- tolia were prohibited to enter by military. The complete Appendix 2, containing all primary (total species number) and secondary data (rarity, endemicity & other weights) for 1846 UTM 10km squares evalutated can be obtained from the Lepidoptera curator of Museum Koenig. Sigbert WAGENER”: Butterfly Diversity in Tukey OLI-PS TEE FEST-LEI “9ET-07 15 “6TI-EOL » :ZOT-98 M'S8-69 53 “89-25 MIS-SE mM “pE-81 miLI-1 exe) JO J9QUINU Joy 9[eIS-ANOJ0J Map 1: Butterfly taxa diversity in Turkey (taxa/square). 6 The best explored areas can be deamed from West to East: The region of Istanbul, the surroundings of Bursa with the Uludag, Sultan Dagları, North- and South-side of Sertavul Gecidi (provinces Icel/ Karaman), the districts of Ayas, Kızılcahamam and Ankara (province Ankara), the area North of Saim- beyli (province Adana), Ala Dagları (provinces Nigde and Kayseri), the area of Kopdag (provinces Erzururm/Bayburt) and Palandóken Dagi (province Erzurum), Ovit Gegidi (province Erzurum/Rize), the district of Sarikamis, the Aras Valley between Kara- kurt and Tuzluca (provinces Kars and Igdir) and the district of Posof (province Ardahan). Endemic taxa Bonner zoologische Beitráge 54 (2005) 3. ENDEMIC BUTTERFLY TAXA IN TURKEY Under the aspect “what should be protected”, nobody will doubt that a site with high species diversity should have priority. But diversity in itself has no comprehen- sive significance. One must also ask for the quality of a taxon, 1.e. does it occur only in Turkey or also in other countries? The taxa so far known to be endemic for Turkey are listed in Table 2. No differences have been made be- tween species and subspecies. Both are equally impor- tant in terms of evolution. Altogether there are 101 en- demic taxa, 1.e. 22.5 % of 448 known taxa in Turkey. The number of squares in which the taxon has been found gives an idea about the distribution and common- ness of each endemic taxon. Table 2. List of endemic taxa in Turkey and number of squares in which they occur Squares Pyrgus melotis graecus (Oberthúr, 1910) Pyrgus bolkariensis De Prins & van der Poorten, 1995 Pyrgus aladaghensis De Prins & van der Poorten, 1995 Archon apollinus forsteri Kogak, 1977 Parnassius nordmanni thomai de Freina, 1980 Parnassius apollo graslini Oberthür, 1891 Pieris bryoniae turcica Eitschberger & Hesselbarth, 1977 Pieris bryoniae goergneri Eitschberger, 1986 Pieris bowdeni Eitschberger, [1984] Lycaena virgaureae aureomicans (Heyne, 1897) Lycaena euphratica Eckweiler, 1989 Tomares nogelii obscura (Rühl, [1893]) Satyrium marcidum mardinum van Oorschot et al. 1985 Satyrium zabni van Oorschot & van den Brink, 1991 Satyrium myrtale armenum (Rebel, 1901) Cupido minimus albocilia van Oorschot et al., 1984 Pseudophilotes bavius vanicola Kogak, 1977 Glaucopsyche astraea astraea (Freyer, [1851[) Glaucopsyche astraea eckweileri Kogak, 1979 Jolana ¡olas lessei (Bernardi, 1964) Turanana cytis kurdistana Eckweiler, 1984 Plebeius eumedon aladaghensis (Kogak, 1979) Plebeius teberdinus nahizericus (Eckweiler, 1978) Plebeius hyacinthus (Herrich-Schäffer, [1847]) Plebeius torulensis (Hesselbarth & Siepe, 1993) Plebeius isauricus isauricus (Staudinger, 1871) Plebeius isauricus latimargo (Courvoisier, 1913) Polyommatus fatima (Eckweiler & Schurian, 1980) Polyommatus myrrha myrrha (Herrich-Schäffer, [1851]) Polyommatus myrrha hakkariensis (Kocak, 1977) Polyommatus aedon myrrhinus (Staudinger, 1901) Polyommatus aedon araxianus (Kogak, 1980) Polyommatus eros molleti Carbonell, [1994] Polyommatus bollandi Dumont, 1998 Polyommatus buzulmavi Carbonell, [1992] Polyommatus syriacus burak (Kogak, 1992) Polyommatus dezinus (de Freina & Witt, 1983) Sigbert WAGENER”: Butterfly Diversity in Tukey Endemic taxa Squares Polyommatus ossmar ossmar (Gerhard, [1851]) Polyommatus interjectus (de Lesse, 1960) Polyommatus antidolus (Rebel, 1901) Polyommatus kurdistanicus (Forster, 1961) Polyommatus menalcas (Freyer, [1837]) Polyommatus hopfferi (Herrich-Scháffer, [1851]) Polyommatus poseidon poseidon (Herrich-Scháffer, [1851]) Polyommatus putnami Dantchenko & Lukhtanov, 2002 Polyommatus dama dama (Staudinger, 1892) Polyommatus maraschi Forster, 1956 Polyommatus damocles kanduli Dantchenko & Lukhtanov, 2002 Polyommatus cilicius cilicius Carbonell, 1998 Polyommatus cilicius bolkarensis Carbonell, 1998 Polyommatus sertavulensis (Kocak, 1979) (stat. inc.) Polyommatus ernesti Eckweiler, 1989 (stat. inc.) Polyommatus artvinensis (Carbonell, 1997) Polyommatus bilgini Dantchenko & Lukhtanov, 2002 Polyommatus haigi Dantchenko & Lukhtanov, 2002 Polyommatus sigberti Olivier et al., 2000 Polyommatus lycius (Carbonell, 1996) Polyommatus pierceae Lukhtanov & Dantchenko, 2002 Polyommatus erzindjanensis Carbonell, 2002 Polyommatus turcicolus (Kogak, 1977) Polyommatus guezelmavi Olivier et al., 1999 Polyommatus theresiae Schurian et al., 1992 Polyommatus surakovi sekercioglui Dantchenko € Lukhtanov, 2002 Polyommatus carmon carmon (Herrich-Scháffer, [1851]) Polyommatus carmon munzuricus (Rose, 1978 Polyommatus schuriani (Rose, 1978) Polyommatus anticarmon (Kocak, 1983) Polyommatus huberti (Carbonell, 1993) Polyommatus turcicus (Kocak, 1977) Polyommatus merhaba De Prins et. al., 1991 Polyommatus iphicarmon Eckweiler & Rose, 1993 Polyommatus tankeri (de Lesse, 1960) Polyommatus baytopi (de Lesse, 1959) Argynnis paphia delila Róber, 1896 Argymnis aglaja ottomana Róber, 1896 Argymnis adippe taurica Staudinger, 1878 Boloria graeca karina van Oorschot & Wagener, 1990 Coenonympha leander dierli Kogak, 1977 Hyponephele urartua de Freina & Aussem, [1987] Hyponephele naricina naricoides Groß, 1977 Hyponephele kocaki kocaki Eckweiler, 1978 Hyponephele kocaki melesina Weiss & Skala, 2000 Hyponephele cadusia zerneca Skala, 2003 Erebia ottomana ottomana Herrich-Schäffer, [1847] Melanargia syriaca syriaca (Oberthiir, 1894) Melanargia syriaca kocaki Wagener, 1983 Melanargia grumi Standfuss, 1892 Melanargia titea wiskotti Röber, 1896 Melanargia larissa noacki Wagener, 1983 Melanargia larissa massageta Staudinger, 1901 Melanargia larissa taurica Róber, 1896 Satyrus favonius favonius Staudinger, 1892 90 5 18 8 Bonner zoologische Beitráge 54 (2005) Endemic taxa Squares Chazara egina egina (Staudinger, 1892) 11 Chazara egina aisha Rose, 1986 3 Pseudochazara lydia lydia (Staudinger, 1878) 24 Pseudochazara lydia obscura (Staudinger, 1878) 29 Pseudochazara lydia aurora Eckweiler & Rose, [1989] 32 Pseudochazara mamurra mamurra (Herrich-Schaffer, [1846]) 56 Pseudochazara mamurra birgit Groß, 1978 30 Pseudochazara mniszechii mniszechii (Herrich-Schäffer, [1851]) 152 Pseudochazara anthelea selcuki van Oorschot et al., 1987 8 In the fifth column “endemisms” of Appendix 2(see Footnote 1), the number of endemic taxa occurring in the corresponding square is named. The distribution of the endemisms is shown in Map 2 “endemisms/square”. Two squares (FE68 southside of Ovit Gegidi, province Erzurum, and LH31 10 km N Catak, province Van) each bear with 16 taxa the highest number of endem- isms. To make the map more impressive, this number has been divided into 10 colour grades as indicated at the top of Map 2. These colour grades are listed in Ap- pendix 2 (see Footnote 1) in column 6 “weight 2”. Map 2 shows the distribution of squares with endem- isms over the country. Generally, the number of endem- isms per square is higher from West to East. The 101 endemic taxa are spread over Anatolia with concentra- tions in the Southwest, Northeast and Southeast. These concentrations are the result of processes of repeated glacial displacements and postglacial recolonizations and connected with the fact that the endemisms occur today in or nearby former refuges in which they could survive the different pleistocene glacial periods. In the Mediterranean region these refuges are valleys on the southside of the Taurus mountains and the coastal areas along the Mediterranean Sea southward up to Lebanon and Israel. In Northeast, Anatolia postglacial recoloniza- tion has taken place mostly from the kolchic refuge in today’s Republic of Georgia and the climatically fa- voured Coruh Valley. Many of the here existing taxa are rare Caucasian elements not occurring in other regions of Turkey. In Southeast Anatolia, south of Lake Van, the deep gorges of Botan, Harbur and Zab with her rich vertical structures formed a favourable refuge for nu- merous Tertiary relicts and elements of the Turk-Iranian Zagros fauna and flora, which do not exist in other Turkish regions. 4. RARE BUTTERFLY TAXA IN TURKEY With the previous actions only the species diversity has been treated in two different ways. But also the rarity of the taxa must be taken into consideration under the as- pect of the protection of sites. In the list of butterfly taxa occurring in Turkey, there has been said from how many squares the taxon is known. If a butterfly taxon occurs only in up to 50 squares — that are about 10 % of the highest numbers of squares (more than 500) — it can be deemed as rare in Turkey. All taxa with an occurrence in 51 or more squares remain unconsidered, equally valid for endemic as well as non-endemic taxa. In column seven “rare taxa” of Appendix 2 (see Foot- note 1), the number of such rare taxa in the correspond- ing square is given independently from the number of squares in which the taxa have been found elsewhere. But it is easy to understand that there is quite a differ- ence if a taxon occurs only in one square at all or in fifty squares. Therefore a scale has been used to weight the difference in the occurrence of a taxon in one up to fifty squares through 10 grades of “rarity points” according to the scheme for each taxon (see Table 3). Table 3. Scheme for the calculation of “rarity points” Number of Number of squares / Rarity squares / Rarity rare taxon points rare taxon points 50 — 43 1 20-16 6 42-36 2 15-12 7 35-31 3 11-8 8 30 - 26 4 7-4 2 25-21 5 3= 1 10 If a square holds more than one rare taxon it gains for each taxon the rarity points connected to it. The summa- rized rarity points for each square are shown to the left in column eight “weight 3” in Appendix 2 (see Footnote 1). For the demonstration of rarity in Map 3, the sum of rarity points has been attributed to 10 colour grades as indicated in the “color scale for sum of rarity points” at top of Map 3. The numbers gained by that way repre- sent the rarity “weight 3” of each square as shown to the right of column eight in Appendix 2 (see Footnote 1). Sigbert WAGENER”: Butterfly Diversity in Tukey "LT-91M !ST-PIW ‘€1-ZI 11-01 6-8 LI 5 ‘C-p MSC SZ mi] gy :SUUSTUISPUO JO JoquINU 10F I[LIS-INOJO)) Map 2: Geographical and numerical distribution of endemic taxa in Turkey (endemisms/ square). Bonner zoologische Beitráge 54 (2005) 10 ‘O€T-161M@ “061-191 “09T-1p TH :OrI-ITI -OTI-101 m00I-18 rt ‘08-19 @ ¿09-19 Mm “0b-17 m “07-115 :siurod ÁjLIe1 Jo ums IO] aTedS-INO[OD Map 3: Rarity weights/square. Sigbert WAGENER”: Butterfly Diversity in Tukey 11 5. PRIORITY SITES FOR THE PROTECTION OF BUTTERFLIES IN TURKEY In a further step, to summarize the results, “weight 1”, “weight 2” and “weight 3” of Appendix 2 (see Footnote 1) were added and set to the left in column nine “sum” of Appendix 2. To demonstrate the result in Map 4, the gained values have been attributed to 10 colour grades as indicated in the “Colour scale for sum of weight points” at the top of Map 4. The colour grades | to 10 represent the priority of a square for the protection of butterflies in Turkey and are shown to the right in col- umn nine in Appendix 2. Generally, all squares with high values of more than 5 priority-weight-points (Appendix 2: column “sum”, right) can request for priority in the achievement of pro- tection measures. But that level is almost too low for eastern squares and too high for western ones. In west- ern and Central Turkey only a few squares reach such a level of 5 priority points, and endemisms which occur only in one or a few more western squares without a high number of accompanying rare species would not have — on the base of the analysis given above — any chance for protection because they never can stay against squares with a high diversity and a high number of endemisms and rare species in eastern Anatolia. Therefore it is necessary to differentiate in the assess- ment of the value of protection in the different regions of Turkey. Thus it seems to be appropiate to start from a level of 3 priority points in all western und central prov- inces of Turkey in choosing sites for protection. In any case, a site with very rare endemism should have prior- ity even if the square inhabited by that endemism does not even reach the level of 3 points. Every such square and endemic taxon occurring in it asks for highest atten- tion by the provincial and local proper authorities con- cerned with nature protection. Examples for such cases are given below: Archon apollinus forsteri KOCAK, 1977, only occur- rence in square WMS50, province Kastamonu, Odemis south of Küre, priority points: 2; Polyommatus dama dama (STAUDINGER, 1892), only two squares: DC32, province Malatya, south of Yesily- urt, priority points: 2 (see WAGENER in VAN SWAAY & WARREN 2003), and square DC71, province Adıyaman, north-side of Nemrut DaZı, priority points: 2. The sites are mostly not identical with the whole area of a square but only with a part of it. Hence the square code can give only a rough indication that in that square exists one or more sites with a high claim for protection. Therefore, the sites have to be explored and exactly de- fined before any measures can be taken. 6. RECOMMENDATIONS Each butterfly species is embedded in a special envi- ronment to which it is adapted and from which it is more or less strongly dependent. The relations are mostly very complex and different for each species. Most of the butterflies like sheltered open sunny places with a rich structure of edges of forests, coppices, hedgerows, grassland, fields, rides, glades or erosion gullies which offer the opportunity for patrolling, perching or hilltop- ping and mating. A few prefer the shade of a wood. Not so few need a special foodplant for their larvae, others are more stenophagous or even polyphagous. Many nec- tar-offering flowers are vitally necessary for the butter- flies. The males of most species must have the opportu- nity to suck minerals on wet soil to raise their fitness and the fertility of the copulating female. One may assume that the sites in squares with a high species diversity basically still fulfil all these require- ments, but one should not overlook the threats and de- clines of Turkish butterflies, ultimately caused by habi- tat changes or loss of habitats (see WAGENER in VAN SWAAY & WARREN 1999). It may be allowed to the author to name some threats and to give some special recommendations on the base of his own observations. 6.1. To the address of forestry commissions Almost every provincial capital in Turkey has today its own “Milli Park1”, a place of woodland, old ones or new afforestations, controlled by the forestry commission. As a rule these National Parks are valuable sites for but- terflies but they suffer under a high recreation pressure of the nearby population especially on the weekends of summer months through wild picnicing destroying the low vegetation and leaving behind rubbish. On the Kayatepe Gecidi on the border of provinces Kastamonu/Corum between Tosya and Iskilip existed in an oakwood an isolated population of Melanargia larissa lesbina of particular biogeographic interest. In 1988 this oakwood was found clear-cut and the popula- tion through such a radical change of its habitat de- stroyed. To avoid such unintended threats, the forestry commissions should generally use the selection forest method only. Experience shows that with the afforestation of an area the low vegetation, before degraded by overgrazing, de- velops very soon and plentiful which is followed by many butterfly species finding new suitable habitats here. This was observed for example in the new Atatiirk Ormani west of Konya. When the young trees become Bonner zoologische Beitráge 54 (2005) “YI< “pHT-17 > “07-L1 @ III WEITE 01-6 E :8-L “9-S mM ópy-€ 2-18 :siurod Jy3tam Jo UMS JOJ A[BIS-MOJO) Map 4: Priority sites for the protection of butterflies in Turkey. Sigbert WAGENER”: Butterfly Diversity in Tukey 13 higher and grow more densely together the low vegeta- tion disappears more and more and, for many butterfly species slowly but surely comes the end of their exis- tence. The forestry commissions should give the threat- ened butterflies in consequence of the changes in their habitat the chance to survive by not closing small natu- rally developed clearings and, under circumstances, enlarge such clearings or create some more by taking away some trees. This correlation could be well ob- served in the Camlik Milli Parki, south of the town Yozgat, where within the dense oak and pine wood only very few butterfly species could be found whilst a lot of them could be seen on the top of the hill on open grassy places between single young pines. For the conservation of butterflies this is a very important matter. The Pinus sylvestris-wood between Sarikamis and the Aras valley in the province Kars is only so rich in butterfly species, because there are closed pine-stands always between smaller and larger wet or dry meadows. If the proper forestry commission does not wish to destroy this unique species diversity of 144 taxa it should do without afforestation of the open areas and take care that these meadows be cut once every year not before mid of Au- gust. The same should be done to the meadows along the road from Sarikamis to Handere (Karaurgan) (prov- ince Kars), the pine wood north of Hanak (province Ar- dahan) as well as the pine wood along the road to Kütlü southwest of Góle (province Erzurum) and in all other similar places all over the country. It is to be seen with respect to Turkish forestry authorities making every ef- fort for culturing existent woods and afforestation in struggle against soil erosion; it would be very apprecia- tive, if they would do the same for the conservation of butterflies and through that of many other animals and plant species. 6.2. To the address of the proper authorities for land use and agriculture Overgrazing by sheep, goats and cattle is a long known problem in Turkey not only because of the destruction of the vegetation and following soil erosion caused by this, but also because of one of the heaviest threats to butterflies and other insects as well as to the soil fauna especially in higher mountains anyway already with sparse vegetation. If one wishes to protect a site for but- terflies it is unevitable to reduce overgrazing to exten- sive grazing. To stop grazing at all would not be desir- able in any case as vegetation could grow too high, which is not favourable for numerous butterfly species. This must be considered from place to place and from species to species one wishes to protect. Where meadows are cut once a year as a traditional way of land use, this should be maintained. The life cycle of the butterflies is adjusted to such a form of land use and any change would cause heavy damage and loss of spe- cies. It is only to take care that cutting of the meadows does not begin before mid August. That’s when the but- terflies — mostly Blues, Fritillaries and Satyrines — in- habiting such sites have their flying period between 10" July and 10" August. The butterflies otherwise would not find enough flowers for nectar with the consequence of lower fertility and lower egg-laying rate, if cutting occurs too early. Thorn-cushion forming species of the plant genus Ono- brychis, Astragalus and Astracantha are typical of high mountain steppes and are food plants of some Clouded Yellows (Colias) and numerous Blues of the genus Polyommatus especially in southeastern Anatolia. In 1985, it could be observed on the northside of Giizeldere Gec- idi (province Van) at about 2500 m that a large area of the slope covered with that plant formation had been de- stroyed through deep ploughing with a tractor not slope- parallel but from top downhill. Such a dangerous behav- iour has not only taken away the habitat for the plants and the insects living on them, but has also opened the way for severe soil erosion. The thorn-cushion plant- formation might be useless for agricultural purposes, but they are very important for soil preservation, and vital for many very rare insects in Turkey. This ought to be urgently taken into consideration when areas shall be ar- ranged for the conservation of butterflies. In valleys of mountainous regions the slopes on either side of a road are very often preferred habitats for nu- merous plant and insect species. It could be observed several times that with widening out or new construct- ing of such roads soil material, stones and rocks were taken from the slope above the road and tipped on the other side downward the slope. Through such a proce- dure the flora and fauna of both sides of the new road were deleted for a long time and along large distances. Already in the planning stage of a new road it should be deliberated how to minimize the damage and to preserve as much as possible of the present vegetation and the animals living thereon. Otherwise many rare species may be lost from that valley even forever. By such a way the populations of the butterfly Euapatura mirza living on Zelkova carpinifolia, a Tertiary relict, and of the very rare moth Brahmaea ledereri zaba whose cat- erpillars feed on Fraxinus in the Zab valley (province Hakkari) have been heavily threatened in connection with the construction of the new road from Hakkari to Cukurca. Rivulets along the roads should be preserved in any case to ensure that male butterflies can supply their needs for water and minerals. 6.3. Specific localities or regions These rather general recommendations can be strength- ened by aspects considering individual places or areas: 14 Bonner zoologische Beitráge 54 (2005) WAGENER (in VAN SWAAY & WARREN 2003) has identi- fied five areas for eight target-species occuring in Tur- key out of 34 in Europe dealt with in the Project “Prime Butterfly Areas in Europe”: Pyrgus cirsii, Zerynthia caucasica, apollo, Lycaena ottomanus, Maculinea arion, Maculinea nausithous, Polyommatus dama, and Euphydryas aurinia. The five areas are: a site 10 km SSE of Yesilyurt, prov. Malatya; Dimcay Valley near Alanya, prov. Antalya; Kopdagı Gegidi, prov. Bayburt and Erzurum; Palandóken, prov. Erzurum; O- vitdagı Gecidi, Prov. Erzurum and Rize. For the last three areas also see below. Parnassius Belgrad ormanı. Thanks to the forestry authorities the Belgrad ormani in todays Greater Istanbul is the only place where nature has still a chance to survive (square PF65 in Appendix 2). But even here some rare butterfly species like Pyronia cecilia and Nemeobius lucina seem to have become extinct during the last thirty years. The understandable search for recreation of the town’s peo- ple should be directed and controlled to less vulnerable parts of the wood. Protection should also include the conservation of small grassy open places with only one cut of the meadows in late summer. Uludag near Bursa (2543 m). It is recommended to enlarge the Milli Park1 to the whole area of the Uludag comprising the squares PE73, PE74, PE82 and PE83 (see Appendix 2) including Sogukpinar to a nature re- serve. To satisfy recreation pressure, the place between the forester’s lodge and the sanatorium at about 1550 m NN on the westside, the plateau north of the summit nearby the funicular from Bursa and the sporting centre as well as the immediate surrounding of the hotels at about 2000-2100 m NN should be excepted. From a lepidopterological point of view the closed 4bies wood is not such a valuable place for butterflies except small clearings for Parnassius mnemosyne, but the summit re- gion of Uludag above treeline with the Nardus stricta meadows on the northside and the rocky and stony parts on the southside claim for absolute protection and should be held free from recreation during the snowless time of the year. Here are the habitats of the rare butter- fly species, Pieris bryoniae turcica, Plebeius pyrenaicus dardanus, Plebeius hyacinthus (type locality), Erebia ottomana ottomana (type locality), Pseudochazara beroe beroe (type locality), Parnassius apollo and other butterflies. But even the area between Osman Gazi and Sogukpinar (500 — 1400 m) on the westside of Uludag bears numerous other rare butterfly species and is the only locality in Turkey for Pseudochazara mniszechii tisiphone. To save the nature of Uludag with its unique relict fauna, permission to build more hotels and ski- lifts for winter sports should not be granted. Sultan Dagları. The Sultan Dagları, a mountain up to 2610 m NN situated in the provinces Afyon, Konya and Isparta between the towns Cay and Doganhisar offers themselves to the creation of a Nature Reserve. Espe- cially the valleys open to the plain on the north- and eastside and the summit region, including the pass road from Aksehir/Engilli to Bagkonak, are inhabited by a very rich flora and fauna with several endemisms and rare butterfly species (see squares UH26, UH32, UH36, UH45, UH46, UH50, UHS3, UH54, UH63, UH64 in Appendix 2). The mountain is nearly unsettled. Settle- ments, small villages and towns follow the foot of the mountain to the plain of Konya. During summer the zone above treeline serves sheep as pasture. Grazing should be restricted before damage to the vegetation and soil becomes apparent. Abant Gölü southwest of Bolu. Abant Gölü (UK59) and Abant valley (UL60 in Appendix 2) in the province Bolu are two sites, geographically connected with each other, with a number of rare plant and butterfly species. The place is threatened by a high recreational pressure already apparent by an alarming decline of several spe- cies like Zerynthia caucasica, Pieris bryoniae turcica, Erebia aethiops aethiops and others. The forestry au- thorities have done well to concentrate the search for recreation, especially for children, to a restricted area. The danger is that along the valley and around the lake might be set up a lot of small “lokantast” (restaurants) — as has already happened in the Dimgay valley near Alanya (province Antalya) — and that some more hotels might be erected. It would mean the death of the par- ticularly delightful scenery of this area and the extinc- tion of more than one rare plant and animal species. That should be avoided in any way. Sertavul Gecidi. This pass on the road from Karaman to Silifke (border of the provinces Karaman and Icel) (WF28 in Appendix 2) offers habitats for 9 endemic and 12 rare butterfly taxa and claims for protection with pri- ority. Meadows especially on the Karaman side of the pass should by no way be changed into arable land. Ex- tensive grazing or grass cutting once a year in late summer would be enough to preserve this place. Pine wood north of Saimbeyli. About 10 to 15 km north of Saimbeyli (province Adana) a large pine wood extends on both sides of the road to Tufanbeyli, 1400— 1550 m NN, which is lepidopterologically already a his- torical place under the name “Hadjin”; it is still today a valuable habitat for 11 endemic and 15 rare butterfly taxa (see square BC41 in Appendix 2) and the only lo- cality for Polyommatus theresiae. The site does not seem to be threatened by maintaining the small open places and present use as selection forest and occasional extensive grazing by sheep. But it is to be assumed that in the course of time recreational pressure will occupy the wood. The forestry authorities should recognize this Sigbert WAGENER”: Butterfly Diversity in Tukey | threat in good time and direct the pressure to the most southern part of the wood. Kopdag. The Kopdag (2963 m NN) on the border be- tween the provinces Bayburt and Erzurum (square FE23 in Appendix 2) gleams through high diversity (10 en- demic and 37 rare butterfly taxa). The whole area should get the status of a nature reserve upwards an alti- tude of about 1750 m NN because it is extraordinary rich in wet and dry ecological structures with different plant associations and insect communities. To preserve this unique place, the traditional land use through graz- ing by sheep on the southern slopes (prov. Erzurum) and cutting on western slopes (prov. Bayburt) should be maintained, grazing by cattle and the change to arable land should strongly be forbidden. Sheep grazing should not be intensified rather extensified in regard to the pre- sent state. Ovit gecidi. The whole valley from Yukari Ozbag in the Coruh valley upward to the provincial border of Erzu- rum recommends itself for nature reserve (squares FE68, FE69, FES9 in Appendix 2). It possesses the highest number of butterfly taxa and endemisms in Tur- key with 33 rare species. The slopes are downward from the village Cayirózú so steep that they can hardly be used for any purpose. Settlements are nearly missing, the village Ozliice is situated on a plateau near the upper edge of the valley. Most of the species are concentrated on the slopes near Ozliice. Of another species composi- tion is the butterfly fauna in the pass region (2600 m) al- ready in the province Rize and from the pass downward to Sivrikaya and Dereköy on the northside of the pontic mountain with several alpine Caucasian elements (espe- cially of the genus Erebia). This part of the mountain on both sides of the road to Rize should be included into the nature reserve (squares FE59, FE49, FF40 in Ap- pendix 2). Both sides of the pass have a different cli- mate and therefore a different flora and fauna which makes this region a very interesting one under ecologi- cal and biogeographical aspects. The valley on the southside of the pass does not need any management, if the present state will be maintained. The very productive meadows on the northside above Sivrikaya in the province Rize could be cut once a year in late summer. But the yaylası are problematical by too much cattle grazing on the slopes and the settlements of the herdsmen with their families especially if the places are repeatedly changed. That way of land use conducts to fertilizing and in consequence to a commutation in the plant composition of the meadows wherefrom the disappearance of the foodplants of rare butterfly species unevitably follows and finally the disappearance of the butterflies themselves. Palandöken Daéglari. Palandóken Dagları (about 3100 m NN) about 10 km south of Erzurum and traversed by un the road from Erzurum to Tekman are mountains differ- ent to the Kopdag and Ovit Gegidi in its natural outfit but also rich in structures. The species composition therefore is not the same but in diversity, number of en- demisms and rare species also very high (see squares FE90, FE91 in Appendix 2). There are long- and shoft- grassy slopes, rocky and stony ones and, in the upper parts, erosion gullies and ridges with well developed thorn-cushion-formations. Most of the butterflies fly be- tween 2200 and 2500 m NN, others only above that line. Threats go out to the butterflies from overgrazing and in the lower parts especially from recreation pressure (pic- nicing in summer and winter sports). Just in the sur- roundings of Kayak Tesisleri the number of butterflies 20 years ago was still plentiful, today 1t has obviously declined. It should be taken into consideration, if it is not possible to reconcile conservation of nature and jus- tified recreation through a sensible planning and man- agement in cooperation with the relevant sections of the University. Aras valley. Between Karakurt (1510 m NN, province Kars) and Tuzluca (1075 m NN, province Igdir) the river Aras has carved out a deep valley in volcanic lay- ers and created a very impressive landscape. The valley is the area of lowest precipitation within Turkey. But the higher slopes of the numerous side-valleys and the ero- sion gullies profit from the melting covering of snow and allow a diverse shrub vegetation, whilst the herba- ceous steppe vegetation on the ground of the valley al- ready dies by the end of June from the lack of rain. These differences in short distances create a lot of dif- ferentiated habitats and allows for very high diversity of endemic and rare butterfly taxa (see square LK53, south of Akcay). Here is the only occurrence of Lycaena phoenicurus, Satyrium hyrcanicum cyri, Plebeius chris- tophi transcaucasicus in Turkey, and the only locality of Polyommatus aedon araxianus at all (all Turk-Iranian faunal elements). As the area is almost uninhabited nothing stands in the way to declare the valley with its side-valleys as a nature reserve. However, care should be taken that grazing does not get out of control. Region south of Lake Van. The provinces south of Lake Van (Bitlis, Van, Siirt, Sirnak and Hakkari) may be deamed as the treasure chamber of nature for Turkey. Plateaus between 2000 and 3000 m protruded by moun- tains up to 4135 m covered from eternal snow, deep ra- vines of streams tributary to Tigris and populated by a relatively very thin human population. Especially some high passes like Kuzgunkiran (LH04), Kurubas (LH55), Giizeldere (MHO2), Siivarihalil (LG34, LG44, LG45, LG58), Tanintanin (LG14, LG25, LG35) and the deep valleys and ravines are very rich in endemic taxa and rare plants and insects. Most of these squares are trav- ersed by the routes from Bitlis to Van, from Catak to Van, from Van to Ytiksekova — Semdinli, from Van to 16 Bonner zoologische Beitráge 54 (2005) Hakkarı — Cukurca and from Sırnak to Hakkari which are the best explored. But without doubt there are many other places with a valuable flora and fauna, only these are not accessible. The whole region is predestined for a large National Park like the big ones in the United States of America. It would not even be needed to reset- tle people as the small towns and villages could remain and some could be included in a comprising manage- ment plan for the whole region, which conserves the present state of nature and gives some of the people an occupation. The small irrigated fields in the surround- ings of the settlements are valuable habitats for butter- flies and therefore should be preserved but not enlarged. Grazing in defined limits and horse breeding could be carried on in the traditional way. A weak and controlled mountaineering tourism might be of interest to the population to raise their income. But the main aim should be to conserve the unique nature, flora and fauna of the whole region accompanied by scientific explora- tion and research work — after it has been pacified. Let us hope! Acknowledgements. The author wishes to give his cordial thanks to Bernd Beermann, Miinster, for computerizing the maps, Hans Duffels, Amsterdam, for fruitful discussions, Harry van Oorschot, Badhoevedoorp, for help and the data from his databank, and Hikmet Ozbek, Erzurum, for the exact topography of some localities. REFERENCES AKIN, €. 2000. Edirne kelebekleri hakkinda notlar (Lepi- doptera). CESA Miscellaneous Papers 63: 5-8. Ankara 15.04.2000. Avcl, U. & ÖZBEK, H. 1996. Erzurum ve bazi cevre illerin Lycaenidae (Lepidoptera) tiirleri üzerinde faunistik calismalar. Türkish Journal of Zoology 20(1996): 73— 87. TUBITAK. Avcı, U. & ÖZBEK, H. 1996. Contribution to the knowl- edge of the Rhopalocera (Lepidoptera) fauna of East- ern and Northeastern Anatolia of Túrkey (Families: Papilionidae, Pieridae, Nymphalidae, Satyridae and Hesperiidae). Türkish Journal of Zoology 20(1996): 361-373. TUBITAK. BARANIAK, E., BAKOWSKI, M. & NOWACKI, J. 1994. A con- tribution to the knowledge of the Lepidoptera of Euro- pean Turkey Part I Macrolepidoptera. CESA Miscella- neous Papers 19: 1-5. Ankara 22.03.1994. CARBONELL, F. 1996. Contribution a la connaissance du genre Agrodiaetus HUBNER (1822): Agrodiaetus lycius n. sp. en Turquie méridionale, (Lepidoptera: Lycaeni- dae) (avec la collaboration de Luc Manil). Linneana Belgica 15(7): 281-285. CARBONELL, F. 1997. Contribution a la connaissance du genre Agrodiaetus HUBNER, (1822). Agrodiaetus actis artvinensis n. ssp. en Turquie orientale (Lepidoptera Lycaenidae). Linneana Belgica 16(4): 140-141. CARBONELL, F. 1998. Contribution a la connaissance du genre Agrodiaetus HUBNER, (1822). Position taxi- nomique d’Agrodiaetus anticarmon KOCAK, 1983 (Lepidoptera: Lycaenidae). Linneana Belgica 16(7): 263-265. CARBONELL, F. 1998. Contribution a la connaissance du genre Agrodiaetus HUBNER, (1822). Agrodiaetus cilicius n. sp. en Turquie méridionale (Lepidoptera Lycaenidae). Linneana Belgica 16(8): 343-349. CARBONELL, F. 2002. Agrodiaetus erzindjanensis n. sp. de Turquie (Lep., Lycaenidae). Bulletin de la Société en- tomologique de France 107(5): 524. CRAMER, P. 1775. De uitlandsche Kapellen voorkommende in de drie waereld-deelen Asia, Africa en America. S. J. Balde, Amsteldam, and Barthelemy Wild, Utrecht. Vol. 1(1—7): xxx+16+132 pp., pls 1-84. DANTCHENKO, A. V. & Lukhtanov, V. A. 1994. New taxa of the subgenus Agrodiaetus HUBNER, (1822) from Caucasus (Lepidoptera, Lycaenidae). Atalanta (Wiirz- burg) 25(1/2): 207-213, colour pl. 8. DOGANLAR, M., OZBEK, H., ECEVIT, O. & YURKSEL, H. 1982. Dogu Anadolu bölgesinin bazı Lepidopter'leri. Bitki Koruma Búlteni 21(3): 155-172. DUMONT, D. 1998. Une nouvelle espece de Lycaenidae du sud de la Turquie: Polyommatus bollandi n. sp. Linne- ana Belgica 16(8): 335-338. HANUS, J. & HOAREAU, D. 1998. Polyommatus (Agrodi- aetus) dama (STAUDINGER, 1892) on the Nemrut dagi in Turkey (Lepidoptera: Lycaenidae). Nachrichten des entomologischen Vereins Apollo, N. F. 18(4): 416. HELSDINGEN, P. J. VAN, WILLEMSE, L. & SPEIGHT M.C.D. (eds.) 1996. Background information on invertebrates of the Habitats Directive and the Bern Convention Part I — Crustacea, Coleoptera and Lepidoptera. Nature and environment 79. 280 pp. Council of Europe, Stras- bourg. HESSELBARTH, G., VAN OORSCHOT, H. & WAGENER, S. 1995. Die Tagfalter der Tiirkei unter Beriicksichtigung der angrenzenden Lander. Vol. 1 + 2: 1354 pp. Vol. 3: 847 pp. Selbstverlag Sigbert Wagener, Bocholt. HERBST, J. F. W. 1798. Natursystem aller bekannten in- und auslandischen Insekten. Schmetterlinge 9, 206 pp., 30 pls. Pauli, Berlin. HUSEYINOGLU, Y. 2000. Sincanli (Afyon) Papilionoidea ve Hesperioidea faunası üzerine arastirmala (Lepidopte- ra). CESA Miscellaneous Papers 60: 1-8. Ankara 25.12.1999, ICZN 2000. Opinion 1944. Papilio sylvanus ESPER, 1777 (currently known as Ochlodes sylvanus or O. venatus faunus; Insecta, Lepidoptera): specific name con- served. Bulletin Zoological Nomenclature 57(1): 56- a7. Kocak, A. O. 1999. Orta Anadolu'da yeni bir Agrodiaetus Hbn. türünün tanımı (Lepidoptera, Lycaenidae). CESA Miscellaneous Papers 59: 3—7. Ankara 14.11.1999. Kocak, A. O. 2000. Revisional notes on the taxonomy of “eurypilos GERHARD” and “sukarovi DANTCHENKO & LUKHTANOV” in the genus Polyommatus Latreille (Ly- caenidae, Lepidoptera). CESA Miscellaneous Papers 65/66: 8-10. Ankara 25.05.2000. Kocak, A. O. & Seven, S., 1996. Tokat Papilionoidea ve Hesperioidea faunası ve ekolojisi hakkinda. Priamus 7(3): 93-116. Ankara 15.05.1996. Kocak, A. O. 8 Seven, S. 1998. A tentative list of the threatended butterflies in Turkey. CESA Miscellane- ous Papers 52: 3-8. Ankara 28.07.1998. Kocak, A. O. & SEVEN, S. 1998. Karyagdi Daginin Papil- ionoidea ve Hesperioidea fauznası hakkinda (Lepidop- Sigbert WAGENER”: Butterfly Diversity in Tukey 17 tera). CESA Miscellaneous Papers 53/54: 1-12. Anka- ra 28.08.1998. LUKHTANOV, V. A. & DANTCHENKO, A. V. 2002. Descrip- tions of new taxa of the genus Agrodiaetus HUBNER, [1822] based on karyotype investigation (Lepidoptera, Lycaenidae). Atalanta (Würzburg) 33(1/2): 81-107, colour pl 1. MAIRIAUX, J. & HUTSEBAUT, J. 1997. The identification of Apatura metis and Apatura ilia and their distribution in Greece and Turkey (Lepidoptera: Nymphalidae). Phegea 25(2): 65-79. OLIVIER, A. 2000. Discovery of the types of Polyommatus (Agrodiaetus) actis and its taxonomic consequences (Lycaenidae). Nota lepidopterologica 23(2): 86-118. OLIVIER, A. 2000. Christian Friedrich FREYER’s ,,Neuere Beitráge zur Schmetterlingskunde mit Abbildungen nach der Natur“: an analysis, with new data on its pub- lication dates (Insecta, Lepidoptera). Beiträge zur En- tomologie. Berlin 50(2000) (2): 407-486. OLIVIER, A., VAN DER POORTEN, D. 8 DE PRINS, W. 1998. Rhopalocera and Grypocera of Turkey 16. Taxonomic notes on Plebeius christophi and Plebeius idas in northeastern Turkey (Lepidoptera, Lycaenidae). Phe- gea 26(3): 87-102. OLIVIER, A., PUPLESIENE, J., VAN DER POORTEN, D., W. DE PRINS & WIEMERS, M. 1999. Revision of some taxa of the Polyommatus (Agrodiaetus) transcaspcius group with descriptions of a new species from Central Ana- tolia (Lepidoptera: Lycaenidae). Phegea 27(1): 1-20. OLIVIER, A., VAN DER POORTEN, D., PUPLESIENE, J. & DE PRINS, W. 2000. Polyommatus (Agrodiaetus) artvinensis (CARBONELL, 1997), stat. nov. and P. (A.) sigbern sp. nov., two vicariant species known so far only from Tur- key (Lepidoptera: Lycaenidae). Phegea 28(2): 59 — 66. PAGES, J. & CHARMEUX, J.-F. 1997. Apatura ilia, “nou- velle” espece pour la faune turque (Lepidoptera, Nym- phalidae). Alexanor 19(8)1996(1997): 454. SCHURIAN, K. G. & ECKWEILER, W. 1997. Wiederfund von Polyommatus (Agrodiaetus) dama STAUDINGER, 1892 in der Tiirkei (Lepidoptera: Lycaenidae). Nachrichten des entomologischen Vereins Apollo, Frankfurt a. M., Suppl. 16: 49-52. SEVEN, S. 1994. Kizilcahamam Kocacay vadisi'nin diurnal Lepidopterlerinin ekolojisi ve faunası. Priamus 7(1/2): 3-62. Ankara 20.01.1994. SEVEN, S. 1995. Trakya Lepidoptera faunasına katkılar. CESA Miscellaneous Papers 23/24: 1-13. Ankara 15.01.1995. SEVEN, S. & BAKOWSKI, M. 1996. Soguksu Milliparkı (Ankara) Lepidoptera faunasına katkılar. Priamus 7(4): 156-170. Ankara 20.08.1996. SEVEN, S. 1996. Türkiye Lepidoptera faunası ve ekolojisi üzerine arastırmalar — I. Priamus 8(1/2): 1-52. Ankara 20.09.1996. SEVEN, S. 2000. Kirikkale Lepidoptera faunas: ekoloji ve taksonomisis üzeerine arastirmalar (Lepidoptera). Pri- amus 10(1/2): 1-101. Ankara 25.08.2000. SEVEN, S. & KEMAL, M. 2000. Additions to the local fau- nas of the diurnal Lepidoptera in the province of An- kara (North Turkey). CESA Miscellaneous Papers 62: 1-7. Ankara 15.02.2000. SEVEN, S. & KEMAL, M. 2000. Additions to the diurnal Lepidoptera fauna of the districts Cubuk and Kalecik (Ankara Prov., North Turkey). CESA Miscellaneous Papers 63: 1-5. Ankara 15.04.2000. SEVEN, S., HUSEYINOGLU, Y., KEMAL, M. & OZDEMIR, M. 2000. Recent collections of the diurnal Lepidoptera of the Soguksu N. P. and Isik Dag (Kizilcahaman, An- kara province, N. Turkey). CESA Miscellaneous Pa- pers 64: 1-6. Ankara 25.05.2000. : SKALA, P. 2003. New taxa of the genus Hyponephele Muschamp, 1915 from Iran and Turkey (Lepidoptefa, Nymphalidae). Linneana Belgica, 19(1): 41-50. STAUDINGER, O. 1878-1881. Lepidopteren-Fauna Kleina- sien’s. Horae Societatis Entomologicae Rossicae 14: 129-320, pls. 1-2 (01.11.1878); 321-482, pls. 3-4 (15.05.1879); Nachtráge: 16: 65-135 (1881). TULUHAN, Y. K. 1998. Ecological diversity of the Eastern Mediterranean region of Turkey and its conservation. Biodiversity and Conservation 7(1): 87-96. VAN OORSCHOT, H. & WAGENER, S. 2000. Zu Tomares in der Türkei. Ergänzungen und Korrekturen zu HESSEL- BARTH, VAN OORSCHOT & WAGENER 1995. Die Tag- falter der Türkei. 3 (Lepidoptera). Phegea 28(3): 87— 147. VAN SWAAY, C. & WARREN, M. (eds.) 1999. Red Data Book of European Butterflies (Rhopalocera). Nature and Environment 99: 1-220+8 pp., Council of Europe, Strasbourg. WAGENER, S. 1996. Apatura metis (FREYER, 1829). Pp. 85— 89 in: VAN HELSDINGEN, P. J., WILLEMSE, L. & SPEIGHT, M. C. D. (eds.) Background information on invertebrates of the Habitats Directive and the Bern Convention Part I — Crustacea, Coleoptera and Lepi- doptera. Nature and Environment 79. Council of Europe, Strasbourg. WAGENER, S. 2003. Turkey. Pp. 600-610 in: VAN SWAAY, C. A. M. & WARREN, M. S. (eds.) Prime Butterfly Ar- eas in Europe: Priority sites for conservation. National Reference Centre for Agriculture, Nature and Fisher- ies. Ministry of Agriculture, Nature Management and Fisheries, The Netherlands. WAGENER, S. & VAN OORSCHOT, H. 1998. Rhopalocera and Grypocera of Turkey. Zur Identitát von Callo- phrys mystaphia MILLER, 1913 (Lepidoptera. Lycae- nidae) Phegea 26(2): 61-67, | coloured map. WARREN, M. S. 1992. The conservation of British Butter- flies. Pp. 246-274 in: DENNIS, R. L. H. (ed.) The Ecol- ogy of Butterflies in Britain. Oxford University Press, Oxford. 14+354 pp. WEISS, D. & SKALA, P. 2000. A new subspecies of Hy- ponephele Muschamp, 1915 from Western Turkey (Lepidoptera, Satyrıdae). Atalanta (Würzburg) 32(2000)(1/2): 129-133, colour pls. 8, 9. Author’s address: Dr. P. Sigbert WAGENER', Kapuziner- platz 34, D-56077 Koblenz. Sadly, Dr. Wagener died prior to the publication of his manuscript. All corre- spondence concerning this paper should be directed to the corresponding editor: Dr. D. Stiining (E-mail: d.stuening.zfmk@uni-bonn.de). Received: 15.10.2003 Revised: 11.02.2004 Accepted: 15.04.2004 Corresponding editor: D. Stüning IS Bonner zoologische Beitráge 54 (2005) APPENDIX 1 List of butterfly taxa occurring in Turkey with the num- ber of 10 km UTM squares in which they have been ob- served Hesperiidae Pyrginae Erynnis Schrank, 1801 tages tages (Linnaeus, 1758) 241 marloyi marloyi (Boisduval, [1834]) 122 Carcharodus Hübner, [1819] alceae alceae (Esper, [1780]) 302 lavatherae tauricus Reverdin, 1915 137 flocciferus (Zeller, 1847) 19 orientalis Reverdin, 1913 205 stauderi ambiguus (Verity, 1925) 27 Spialia Swinhoe [1912] phlomidis (Herrich-Schäffer, [1845]) 102 osthelderi osthelderi (Pfeiffer, 1932) 3 orbifer (Hübner, [1823]) 426 Muschampia Tutt [1906] proto aragonensis (De Sagarra, 1924) 22 proteides (Wagner, 1929) 68 poggei poggei (Lederer, 1858) 46 plurimacula (Christoph, 1893) 6 tessellum tessellum (Hübner, [1803]) 87 tessellum nomas (Lederer, 1855) 66 tessellum tersa Evans, 1949 20 Pyrgus Hübner [1819] carthami carthami (Hübner, [1813]) l sidae sidae (Esper, [1784]) 184 malvae malvae (Linnaeus, 1758) 18 melotis melotis (Duponchel, [1834]) 17 melotis ponticus (Reverdin, 1914) 189 melotis graecus (Oberthiir, 1910). 5 serratulae major (Staudinger, 1878) 164 serratulae balcanicus (Warren, 1926) l cirsii (Rambur, [1839]) 24 cinarae cinarae (Rambur, [1839] 38 armoricanus persicus (Reverdin, 1913) 154 aleus alveus (Hübner, [1803]) 40 jupei (Alberti, 1967) 7 bolkariensis De Prins & v. d. Poorten, 1995 e) aladaghensis De Prins & v. d. Poorten, 1995 4 Heteropterinae Heteropterus Duméril 1806 morpheus (Pallas, 1771) 3 Carterocephalus Lederer, 1852 palaemon (Pallas, 1771) | Hesperiinae Eogenes Mabille, [1909] alcides alcides (Herrich-Schäffer, [1852]) 19 lesliei elama Wiltshire, 1941 2 Thymelicus Hübner, [1819] lineola (Ochsenheimer, 1808) 236 sylvestris syriacus (Tutt, [1905]) 387 novus (Reverdin, 1916) acteon acteon (Rottemburg, 1775) hyrax (Lederer, 1861) Hesperia Fabricius, 1793 comma pallida (Staudinger, 1901) Ochlodes Scudder, 1872 sylvanus (Esper, 1777) Gegenes Hiibner, [1819] pumilio (Hoffmansegg, 1804) nostrodamus (Fabricius, 1793) Pelopidas Walker, 1870 thrax (Hübner, [1821]) Papilionidae Parnassiinae Zerynthia Ochsenheimer, 1816 polyxena gracilis Schultz, 1908 cerisy cerisy (Godart, 1824) cerisy ferdinandi Stichel, 1907 deyrollei (Oberthiir, 1869) caucasica (Lederer, 1864) Archon Hübner, 1822 apollinus apollinus (Herbst, 1798) apollinus forsteri Kogak, 1977 apollinus thracicus (Buresch, 1915) apollinaris (Staudinger, [1892]) Parnassius Latreille, 1804 mnemosyne caucasica Verity, [1911] mnemosyne sheljuzhkoi Bryk, 1914 mnemosyne nubilosus Christoph, 1873 nordmanni nordmanni [Ménétriés], [1850] nordmanni thomai de Freina, 1980 apollo graslini Oberthür, 1891 apollo tirabzonus Sheljuzhko, 1924 Papilioninae Iphiclides Hübner, [1819] podalirius podalirius (Linnaeus, 1758) Papilio Linnaeus, 1758 machaon syriacus Eller, 1936 alexanor eitschbergeri Bollino & Sala, 1992 alexanor orientalis Romanoff, 1884 Pieridae Pierinae Anthocharis Boisduval et al., [1833] cardamines cardamines (Linnaeus, 1758) cardamines phoenissa Kalchberg, 1895 damone eunomia (Freyer, 1851) gruneri gruneri Herrich-Schäffer, [1851] gruneri armeniaca Christoph, 1893 Zegris Boisduval 1836 eupheme menestho (Ménétrés, 1832) Euchloe Hübner, [1819] belemia belemia (Esper, [1800]) ausonia taurica Rober, [1907] Elphinstonia Klots, 1930 penia (Freyer, 1851) Aporia Hübner, [1819] Sigbert WAGENER”: Butterfly Diversity in Tukey crataegi (Linnaeus, 1758) Pieris Schrank, 1801 brassicae brassicae (Linnaeus, 1758) krueperi krueperi Staudinger, 1860 mannii hethaea Pfeiffer, 1931 rapae rapae (Linnaeus, 1758) ergane detersa Verity, [1908] napi pseudorapae Verity, [1908] bryoniae turcica Eitschb. & Hesselbarth, 1977 bryoniae goergneri Eitschberger, 1986 bowdeni Eitschberger, [1984] Pontia Fabricius, 1807 callidice chrysidice (Herrich-Schäffer, [1844]) daplidice (Linnaeus, 1758) edusa (Fabricius, 1777) chloridice chloridice (Hübner, [1813]) Colotis Hübner, [1819] fausta fausta (Olivier, [1804]) Coliadinae Colias Fabricius in Illiger, 1807 erate erate (Esper, [1805]) crocea (Fourcroy, 1785) chlorocoma Christoph, 1888 aurorina aurorina Herrich-Scháffer, [1850] caucasica caucasica Staudinger, 1871 thisoa thisoa Ménétriés, 1832 thisoa shakuhensis Sheljuzhko, 1935 hyale (Linnaeus, 1758) alfacariensis Ribbe, 1905 Catopsilia Hiibner, [1819] florella (Fabricius, 1775) Gonepteryx Leach, [1815] rhamni rhamni (Linnaeus, 1758) rhamni miljanowskii Nekrutenko, 1966 rhamni transiens Verity, 1913 rhamni kurdistana de Freina, 1990 farinosa farinosa (Zeller, 1847) farinosa turcirana de Freina, 1983 cleopatra cleopatra (Linnaeus, 1767) cleopatra taurica (Staudinger, 1881) Dismorphiinae Leptidea Billberg, 1820 sinapis sinapis (Linnaeus, 1758) reali Reissinger, 1989 duponcheli lorkovici (Pfeiffer, 1932) Lycaenidae Riodininae Hamearis Hiibner, [1819] lucina (Linnaeus, 1758) Lycaeninae Cigaritis Donzel, 1847 acamas acamas (Klug, 1834) cilissa Lederer, 1861 maxima Staudinger, 1901 Lycaena Fabricius in Illıger, 1807 phlaeas phlaeas (Linnaeus, 1761) dispar rutila (Werneburg, 1864) virgaureae virgaureae (Linnaeus, 1758) virgaureae aureomicans (Heyne, 1897) ottomanus (Lefebvre, 1830) tityrus tityrus (Poda, 1761) alciphron melibaeus (Staudinger, 1878) candens candens (Herrich.Schaffer, [1844]) thersamon (Esper, [1784]) lampon lampon (Lederer, 1870) thetis thetis Klug, 1834 asabinus (Herrich-Schäffer, [1851 ]) ochimus ochimus (Herrich-Schäffer, [1851] phoenicurus (Lederer, 1870) euphratica Eckweiler, 1989 Thecla Fabricius, 1807 betualae betulae (Linnaeus, 1758) Neozephyrus Sıbatanı & Ito, 1942 quercus quercus (Linnaeus, 1758) Tomares Rambur, 1840 nogelii nogelii (Herrich-Schäffer, [1851]) nogelii obscura (Rühl, [1893]) nesimachus (Oberthúr, 1893) callimachus (Eversmann, 1848) romanovi romanovi (Christoph, 1882) Callophrys Billberg, 1829 rubi (Linnaeus, 1758) chalybeitincta Sovinsky, 1905 mystaphia Miller, 1913 suaveola (Staudinger, 1881) Satyrium Scudder, 1876 w-album (Knoch, 1782) spini (Fabricius, 1787) marcidum marcidum (Riley, 1921) marcidum mardinum van Oorschot et al. 1985 ilicis ilicis (Esper, [1779]) zabni van Oorschot & van den Brink acaciae (Fabricius, 1787) abdominalis (Gerhard, [1850]) myrtale armenum (Rebel, 1901) ledereri ledereri (Boisduval, 1848) hyrcanicum cyri (Nekrutenko, 1978) Lampides Hiibner, [1819] boeticus (Linnaeus, 1767) Leptotes Scudder, 1876 pirithous (Linnaeus, 1767) Tarucus Moore, [1881] balkanicus (Freyer, [1843]) Zizeeria Chapman, 1910 karsandra karsandra (Moore, 1865) Cupido Schrank, 1801 minimus minimus (Fuessly, 1775) minimus albocilia van Oorschot et al., 1984 osiris (Meigen, [1829]) argiades (Pallas, 1771) alcetas (Hoffmansegg, 1804) decoloratus (Staudinger, 1886) 20 Bonner zoologische Beitráge 54 (2005) Celastrina Tutt, 1906 argiolus argiolus (Linnaeus, 1758) Pseudophilotes Beuret, 1958 vicrama schiffermuelleri (Hemming, 1929) bavius egea (Herrich-Schäffer, [1852]) bavius eitschbergeri Kocak, 1975 bavius vanicola Kocak, 1977 Scolitantides Hübner, [1819] orion orion (Pallas, 1771) Glaucopsyche Scudder, 1872 alexis alexis (Poda, 1761) astraea astraea (Freyer, [1851[) astraea eckweileri Kocak, 1979 Jolana Bethune-Baker, 1914 iolas lessei (Bernardi, 1964) Maculinea van Eecke, 1915 alcon monticola (Staudinger, 1901) arion arion (Linnaeus, 1758) nausithous (Bergstrasser, 1779) Lachides Nekrutenko, 1984 galba (Lederer, 1855) Turanana Bethune-Baker, 1916 cytis kurdistana Eckweiler, 1984 endymion endymion (Gerhard, [1851 ]) Chilades Moore, [1881] trochylus trochylus (Freyer, 1844) Plebeius Kluk, 1780 pylaon sephirus (Frivaldszky, [1835]) pylaon turcmenicus (Forster,1936) pylaon nichollae (Elwes, 1901) argus aegidion (Meisner, 1818) idas baldur (Hemming, 1934) idas altarmenus (Forster, 1936) christophi transcaucasicus (Rebel, 1901) argyrognomon caspicus (Forster, 1936) argyrognomon subspecies nova ? alcedo (Christoph, 1877) rosei (Eckweiler, 1989) morgianus morgianus (Kirby, 1871) loewii loewii (Zeller, 1847) eurypilus eurypilus (Freyer, 1851) pyrenaicus dardanus (Freyer, [1843] eumedon eumedon (Esper, [1780]) eumedon aladaghensis (Kogak, 1979) eumedon modestus (Nekrutenko, 1972) agestis ([Denis & Schiffermüller], 1775) artaxerxes macedonicus (Verity, 1936) artaxerxes sheljuzhkoi (Obraztsov, 1935) teberdinus nahizericus (Eckweiler, 1978) hyacinthus (Herrich-Schaffer, [1847]) torulensis (Hesselbarth & Siepe, 1993) isauricus isauricus (Staudinger, 1871) isauricus dorsumstellae (Graves, 1923) isauricus latimargo (Courvoisier, 1913) anteros anteros (Freyer, [1838]) anteros crassipunctus (Christoph, 1893) bassoni (Larsen, 1974) Polyommatus Latreille, 1804 semiargus bellis (Freyer, [1842]) semiargus antiochenus (Lederer, 1861) coelestinus (Eversmann, 1843) diana (Miller, 1913) fatima (Eckweiler & Schurian, 1980) escheri (Húbner, [1823]) dorylas dorylas ([Denis & Schiffermiiller], 1775) dorylas armenus (Staudinger, 1871) amandus amandus (Schneider, 1792) thersites (Cantener, [1835]) myrrha myrrha (Herrich-Schaffer, [1851]) myrrha hakkariensis (Kocak, 1977) aedon myrrhinus (Staudinger, 1901) aedon araxianus (Kocak, 1980) cornelia (Gerhard, [1851]) eros yildizae Kocak, 1977 eros molleti Carbonell, [1994] eroides eroides (Frivaldszky, 1835) ciloicus de Freina & Witt, 1983 bollandi Dumont, 1998 icarus (Rottemburg, 1775) buzulmavi Carbonell, [1992] daphnis daphnis ([Denis & Schiffermiiller], 1775) daphnis versicolor (Heyne, [1895]) daphnis elamita (Le Cerf, 1913) bellargus (Rottemburg, 1775) syriacus burak (Kogak, 1992) dezinus (de Freina & Witt, 1983) ossmar olympicus (Lederer, 1852) ossmar ossmar (Gerhard, [1851]) corydonius caucasicus (Lederer, 1870) coridon coridon (Poda, 1761) alcestis alcestis (Zerny, 1932) alcestis karacetinae Lukhtanov & Dantchenko, 2002 interjectus (de Lesse, 1960) (stat. inc.) demavendi (Pfeiffer, 1938) ripartii (Freyer, 1830) erivanensis (Forster, 1960) admetus (Esper, 1783) mithridates (Staudinger, 1878) antidolus (Rebel, 1901) kurdistanicus (Forster, 1961) menalcas (Freyer, [1837]) hopfferi (Herrich-Scháffer, [1851]) poseidon poseidon (Herrich-Schaffer, [1851]) putnami Dantchenko & Lukhtanov, 2002 caeruleus caeruleus (Staudinger, 1871) dama dama (Staudinger, 1892) wagneri (Forster, 1956) maraschi Forster, 1956 (stat. inc.) cilicius cilicius Carbonell, 1998 Sigbert WAGENER”: Butterfly Diversity in Tukey cilicius bolkarensis Carbonell, 1998 firdussii (Forster, 1956) artvinensis (Carbonell, 1997) bilgini Dantchenko & Lukhtanov, 2002 actis (Herrich-Schäffer, [1851 ]) sigberti Olivier et al., 2000 pseudactis (Forster, 1960) (stat. inc.) haigi Dantchrenko & Lukhtanov, 2002 sertavulensis (Kocak, 1979) (stat. inc.) ernesti Eckweiler, 1989 (stat. inc.) erzindjanensis Carbonell, 2002 altivagans altivagans (Forster, 1956) damocles kanduli Dantchenko & Lukhtanov, 2002 lycius (Carbonell, 1996) pierceae Lukhtanov & Dantchenko, 2002 elbursicus (Forster, 1956) turcicolus (Kocak, 1977) ninae (Forster, 1956) aserbeidschanus (Forster, 1956) guezelmavi Olivier et al., 1999 theresiae Schurian et al., 1992 carmon carmon (Herrich-Schaffer, [1851 ]) carmon munzuricus (Rose, 1978) anticarmon (Kocak, 1983) schuriani schuriani (Rose, 1978) surakovi sekercioglui Dantschenko&Lukhtanov, 2002 huberti (Carbonell, 1993) turcicus (Kocak, 1977) cyaneus cyaneus (Staudinger, 1890) merhaba De Prins et. al., 1991 iphigenia iphigenia (Herrich-Schäffer, [1847]) iphicarmon Eckweiler & Rose, 1993 tankeri (de Lesse, 1960) baytopi (de Lesse, 1959) damon kotshubeji (Sovinsky, [1916]) phyllis vanensis (de Lesse, 1957) Nymphalidae Libytheinae Libythea Fabricius in Mliger, 1807 celtis (Laicharting, 1782) Heliconiinae Argynnis Fabricius in Illiger, 1807 paphia paphia (Linnaeus, 1758) paphia delila Róber, 1896 pandora pandora (|Denis & Schiffermiiller], 1795) aglaja aglaja (Linnaeus, 1758) aglaja ottomana Röber, 1896 adippe adippe ({Denis & Schiffermüller], 1775) adippe taurica Staudinger, 1878 niobe orientalis Alphéraky, 1881 Issoria Hübner, [1819] lathonia (Linnaeus, 1758) Brenthis Hübner, [1819] oN 159) m= WR WHA FW N WWOA PR — 100 90 14 326 117 23 58 16 238 368 ino schmitzi Wagener, 1983 daphne daphne (Bergsträsser, 1780) hecate hecate (|Denis & Schiffermüller], 1775) hecate transcaucasica Wnukowsky, 1929 mofidii Wyatt, 1969 Boloria Moore, [1900] euphrosyne euphrosyne (Linnaeus, 1758) dia (Linnaeus, 1767) caucasica (Lederer, 1852) graeca karina van Oorschot & Wagener, 1990 Nymphalinae Vanessa Fabricius in Iliger, 1807 atalanta atalanta (Linnaeus, 1758) cardui (Linnaeus, 1758) Inachis Hiibner, [1819] io io (Linnaeus, 1758) Aglais Dalman, 1816 urticae turcica (Staudinger, 1861) Polygonia Hübner, [1819] c-album (Linnaeus, 1758) egea (Cramer, [1775]) Araschnia Hiibner, [1819] levana (Linnaeus, 1758) Nymphalis Kluk, 1780 vaualbum vaualbum (| Denis & Schiff.], 1775) polychloros polychloros (Linnaeus, 1758) xanthomelas fervescens (Stichel, [1908]) antiopa antiopa (Linnaeus, 1758) Hypolimnas Hübner, [1819] misippus (Linnaeus, 1764) Apaturinae Apatura Fabricius in Illıger, 1807 ilia ({Denis & Schiffermiiller], 1775) metis metis Freyer, 1829 Euapatura Ebert, 1971 mirza Ebert, 1971 Thaleropis Staudinger, 1871 ionia (Eversmann, 1851) Charaxinae Charaxes Ochsenheimer, 1816 jasius jasius (Linnaeus, 1767) Limenitidinae Limenitis Fabricius in Illiger, 1807 camilla camilla (Linnaeus, 1764) reducta herculeana Stichel, [1908] Neptis Fabricius in Mliger, 1807 rivularis rivularis (Scopoli, 1763) Melitaeinae Euphydryas Scudder, 1872 iduna inexspectata (Sheljuzhko, 1934) orientalis (Herrich-Schäffer, [1845] aurinia bulgarica (Fruhstorfer, 1917) Melitaea Fabricius in Illiger, 1807 cinxia cinxia (Linnaeus, 1758) phoebe phoebe ([Denis & Schiffermiiller], 1775) punica telona Fruhstorfer, 1908 154 22 Bonner zoologische Beitráge 54 (2005) collina collina Lederer, 1861 collina lokmani van Oorschot & v. d. Brink, 1994 arduinna (Esper, [1783]) trivia trivia ([Denis & Schiffermiiller], 1775 didyma didyma (Esper, [1778]) persea persea Kollar, 1849 interrupta interrupta Kolenati, 1846 diamina diamina (Lang, 1789) aurelia ciscaucasica Rjabov, 1926 caucasogenita Verity, 1930 athalia athalia (Rottemburg 1775) Danainae Danaus Kluk, 1780 chrysippus chrysippus (Linnaeus, 1758) Satyrinae Kirinia Moore, 1893 roxelana (Kramer, [1777]) Esperarge Nekrutenko, 1988 climene valentinae (Miller, 1923) Pararge Hiibner, [1819] aegeria aegeria (Linnaeus, 1758) aegeria tircis (Godart, 1821) Lasiommata Westwood, 1841 megera transcaspica (Staudinger, 1901) petropolitana petropolitana (Fabricius, 1787) maera (Linnaeus, 1758) menava Moore, 1865 Ypthima Hiibner, 1818 asterope asterope (Klug, 1832) Coenonympha Hibner, [1819] arcania (Linnaeus, 1761) glycerion glycerion (Borkhausen, 1788) leander leander (Esper, [1784]) leander dierli Kocak, 1977 saadi mesopotamica Heyne, [1894] symphyta Lederer, 1870 pamphilus (Linnaeus, 1758) Triphysa Zeller, 1850 phryne phryne (Pallas, 1771) Pyronia Húbner, [1819] tithonus (Linnaeus, 1767) cecilia (Vallantin, 1894) Maniola Schrank, 1801 jurtina (janira) phormia (Fruhstorfer, 1909) megala (Oberthür, 1909) telmessia (Zeller, 1847) halicarnassus Thomson, 1990 Hyponephele Muschamp, 1915 wagneri wagneri (Herrich-Schäffer, [1846] urartua de Freina & Aussem, [1987] naricina naricoides Grok, 1977 cadusia zerneca Skala, 2003 kocaki kocaki Eckweiler, 1978 kocaki melesina Weiss & Skala, 2000 Ivcaon ?collina (Rober, 1897) (stat. inc. nom. inc.) ?lycaon (?species nova, nom. inc.) 34 7 68 295 411 9] oe 11 12 6 94 15 199 148 lycaon bugrai Kogak, 1989 (stat. inc.) lycaon (Gürün-Form, stat. inc., nom. inc.) lupina intermedia (Staudinger, 1886) Proterebia Roos & Arnscheid, 1980 afra afra (Fabricius , 1787) Erebia Dalman, 1816 aethiops aethiops (Esper, [1777]) aethiops melusina Herrich-Schaffer, [1847] medusa euphrasia Fruhstorfer, 1917 hewitsonii Lederer, 1864 ottomana ottomana Herrich-Schaffer, [1847] graucasica transcaucasica Warren, 1950 iranica dromulus Staudinger, 1901 melancholica Herrich-Schaffer, [1846] Melanargia Meigen, 1828 russiae russiae (Esper, [1783]) galathea satnia Fruhstorfer, 1917 syriaca syriaca (Oberthúr, 1894) syriaca kocaki Wagener, 1983 syriaca karabagi Kocak, 1976 hylata hylata (Ménétriés, 1832) grumi Standfuss, 1892 titea titania Calberla,,1891 titea standfussi Wagener, 1983 titea wiskotti Röber, 1896 larissa lesbina Wagener, 1976 larissa astanda (Boisduval, 1848) larissa noacki Wagener, 1983 larissa massageta Staudinger, 1901 larissa taurica Rober, 1896 Satyrus Latreille, 1810 favonius favonius Staudinger, 1892 parthicus Lederer,1869 ferula (Fabricius, 1793) amasinus amasinus Staudinger, 1861 amasinus iranicus Schwingenschuss, 1939 Minois Hiibner, [1819] dryas dryas (Scopoli, 1763) Hipparchia Fabricius in Illıger, 1807 syriaca syriaca (Staudinger, 1871) senthes (Fruhstorfer, 1908) volgensis delattini Kudrna, 19775 pellucida pellucida (Stauder, 1924) mersina (Staudinger, 1871) statilinus (Hufnagel, 1766) fatua fatua Freyer, [1843] parisatis (Kollar, 1849) Arethusana de Lesse, 1851 arethusa ([Denis & Schiffermüller], 1775) Brintesia Fruhstorfer, [1911] circe (Fabricius, 1775) Chazara Moore, 1893 briseis meridionalis (Staudinger, 1886) persephone transiens (Zerny, 1932) bischoffii (Herrich-Schäffer, [1846]) egina egina (Staudinger, 1892) 231 105 Sigbert WAGENER”: Butterfly Diversity in Tukey 23 egina aisha Rose, 1986 Pseudochazara de Lesse, 1951 beroe beroe (Freyer, 1843) Iydia Iydia (Staudinger, 1878) lydia obscura (Staudinger, 1878) lydia aurora Eckweiler & Rose, [1989] geyeri (Herrich-Scháffer, [1846]) mamurra mamurra (Herrich-Schäffer, [1846]) mamurra birgit Groß, 1978 mamurra schahrudensis (Staudinger, 1881) schahkuhensis brandti (Holik, 1949) pelopea persica (Christoph, 1877) mniszechii mniszechii (Herrich-Schäffer, [1851]) mniszechii caucasica (Lederer, 1864) 3] mniszechii tisiphone Brown, 1980 | anthelea anthelea (Hübner, [1824]) 273 anthelea selcuki van Oorschot et al., 1987 8 thelephassa (Geyer, 1827) 81 ft The sign “ft” with four taxa (Polvommatus escheri, Polvommatus ossmar olympicus, Polyommatus caeruleus caeruleus, Erebia iranica dromulus) symbolizes their extinction in Turkey, because they could no more be ob- served during the last fifty years. “?” for taxa whose taxonomic status and distribution is not fully under- stood. . . 7 Moa ® oo. . . | I f A o rn o iy ; a : t . i sf i i si ; N > = m i ' a + . Seiten 25-95 Bonner zoologische Beiträge Band 54 (2005) | Heft | | Bonn, April 2006 A Revision of Western Palaearctic Medon: the Species of the Atlantic Islands, the Western Mediterranean, and Europe, Except for the Southeast A (Insecta: Coleoptera: Staphylinidae: Paederinae) Volker ASSING Hannover, Germany Abstract. Types and additional material of Medon Stephens are revised. 29 valid species and subspecies, including three doubtful species, are recognized in the region comprising the Western Mediterranean (including Italy and Tunisia), the Atlantic Islands, as well as Western, Central, and Northern Europe. Medon oromii sp. n. (Canary Islands: Gran Ca- naria), M. antricola sp. n. (Canary Islands: El Hierro), and M. kabylicus sp. n. (Algeria) are described. Twenty-three synonymies, two of them pertaining to the previously revised species of the Eastern Mediterranean region, are estab- lished: Medon apicalis (Kraatz, 1857) = Medon orgianus Bordon1, 1980 syn. n.; Medon perniger Coiffait, 1978 = Me- don perniger fraudulentus Coiffait, 1978 syn. n.; Medon dilutus (Erichson, 1839) = M. dilutus meridionalis Bordoni, 1980 syn. n.; M. dilutus cephalus Koch, 1938 = M. dilutus boeticus Jeannel & Jarrige, 1949 syn. n.; Medon dilutus quadriceps (Wollaston, 1864) = Medon vitalei Bernhauer, 1936 syn. n. = M. dilutus eremicus Koch, 1939 syn. n. = M. marocanus Coiffait, 1970 syn. n. = M. mateui Coiffait, 1973 syn. n.; Medon rufiventris (Nordmann, 1837) = M. burdi- galensis Coiffait, 1970 syn. n. = M. aquitanicus Coiffait, 1970 syn. n. = M. siculus Coiffait, 1970 syn. n. = M. sicilianus Coiffait, 1970 syn. n.; Medon africanus (Fauvel, 1872) = M. bodemeyeri Bernhauer, 1915 syn. n.; Medon piceus (Kraatz, 1858) = M. murciensis Coiffait, 1970 syn. n.; Medon cauchoisi Jarrige, 1949 = M. oculatior Peyerimhoff, 1949 syn. n. = M. lusitanicus Coiffait, 1970 syn. n. = M. parviphallus Coiffait, 1973 syn. n. = M. mazices Coiffait, 1973 syn. n. = M. fongondi Coiffait, 1980 syn. n. = M. perraulti Coitfait, 1980 syn. n.; Medon mirei Coiffait, 1980 = M. giachinoi Bordoni, 1988 syn. n.; Medon semiobscurus (Fauvel, 1875) = M. ruber Sahlberg, 1908 syn. n. = M. sahlbergi Scheer- peltz, 1933 syn. n. Medon procerus (Perez Arcas, 1874) and M. sericellus (Fairmaire, 1860), previously regarded as synonyms of M. castaneus (Gravenhorst, 1802) and M. apicalis, respectively, are revalidated. A neotype is designated for Lithocharis sericella Fairmaire, 1860. Lectotypes are designated for Lithocharis oppidana Kraatz, 1857, L. picea Kraatz, 1858, Paederus castaneus Gravenhorst, 1802, Lithocharis africanus Fauvel, 1872, Medon bodemeyeri Bern- hauer, 1915, M. bucharicus Bernhauer, 1902, and M. ruber Sahlberg, 1908. Medon pythonissa (Saulcy, 1864) is re- garded as a subspecies of M. dilutus. All the species not treated in the first part of the revision are described and illus- trated. Bionomic and zoogeographic data are compiled, and the distributions of most species are mapped. A key to the Medon species of the Western Mediterranean region and the Atlantic Islands is provided. Medon bucharicus Bernhauer, 1902 (Middle Asia) is redescribed and moved to Medon again (from Sunius Curtis). Some additional records of species from other regions are presented. The previously published revision of Eastern Mediterranean Medon contains several errors, which are corrected in a separate section. These corrigenda include the following synonymies: M. lydicus Bor- doni, 1980 = M. mimulus Fagel, 1970, syn. n. = M. rhodicus Franz, 1987, syn. n. = M. choparti Coiffait, 1987, syn. n. Key words. Taxonomy, revision, new species, new synonyms, lectotype designations, neotype designation INTRODUCTION The species had never been thoroughly revised and nu- merous species have been described especially from France, the Iberian Peninsula, and from Northwest Af- rica. COIFFAIT (1984) lists 36 species and subspecies for the Western Mediterranean (including Italy, the Atlantic Islands, and Northwest Africa eastwards to Tunisia); three additional species were later described by SERRANO (1993), BORDONI (1988), and ASSING (1998). However, a preliminary examination of abundant material, which had been accumulated during various field trips in the past decades, suggested a significantly lower species di- versity in this region. Many Medon species are inhabi- tants of the leaf litter and occur at relatively high densi- ties, so that it seemed rather unlikely that a considerable | 1. In the recently published first part of the revision of Western Palaearctic Medon and a subsequent supple- ment (ASSING 2004a, 2004b), the species of the Eastern Mediterranean and adjacent regions were treated. This project had been initiated especially by the difficulties | encountered when trying to identify specimens from this | area; attributing names to them was often little more | than a good guess. This observation gave rise to the | suspicion that many names were synonymous and/or misinterpreted, and, as was expected, one of the main | results of this study was a significant reduction of the number of valid names. For several reasons, the situation was hypothesized to be similar for the species of the Western Mediterranean. number of species had just not been found. Moreover, the revision of the Eastern Mediterranean representa- 26 Bonner zoologische Beitráge 54 (2005) tives of the genus had shown that the extent of the dis- tributions and the range of intraspecific variation had generally been underestimated by previous authors. Also, some species, like Medon burdigalensis Coiffait and M. aquitanicus Coiffait, had never been recorded again after their description, although they had been de- scribed from regions where the presence of local en- demics is unlikely; consequently, the absence of records Map 1: The study region. could not be put down to extremely restricted distribu- tions. These considerations, as well as the difficulty en- countered when trying to identify Medon material from the study area suggested that the previous taxonomic concept of the Medon fauna of the Western Mediterra- nean was at least in part inadequate and eventually initi- ated the present revision. Volker ASSING: Western Palaearctic Medon 27 The revision focuses on the species of the Atlantic is- lands and the western parts of the Mediterranean east- wards to Tunisia and Italy. However, since the ranges of several widespread species also extend into Western, Central, Northern, and even Eastern Europe, these re- gions are included, too (Map 1). The recently published revision of Eastern Mediterra- nean Medon is based on an old manuscript version and contains some nomenclatural errors. | use the opportu- nity to correct them in a separate section at the end of this paper. 2. MATERIAL AND MEASUREMENTS Types and additional material from the following insti- tutions and private collections were examined: DEI Deutsches Entomologisches Institut, Eberswalde (L. Zerche) DZUL Departamento de Biologia Animal — Zoologia, Universidad de La Laguna (P. Oromi) IRSNB Institut Royal des Sciences Naturelles de Bel- gique, Bruxelles (D. Drugmand) FMNH Field Museum of Natural History, Chicago (via L. H. Herman) MCNT Museo de Ciencias Naturales de Tenerife MHNG Muséum d Histoire Naturelle, Geneve (G. Cuccodoro) MNHNP Muséum National d Histoire Naturelle, Paris (N. Berti) MNHUB Museum für Naturkunde der Humboldt- Universitat, Berlin (J. Frisch, J. Willers) MRSNT Museo Regionale di Scienze Naturali, Torino (M. Daccordi) NHMB _ Naturhistorisches Museum Basel (E. Sprecher) NHMW Naturhistorisches Museum Wien (H. Schill- hammer) SMNS Staatliches Museum für Naturkunde, Stuttgart (W. Schawaller, K. Wolf-Schwenninger) UCBA — Universita di Catania, Dip. di Biologia Animale (A. Adorno, G. Longo) ZMH Zoological Museum Helsinki (J. Muona) cAss author's private collection cBor private collection A. Bordoni, Firenze cErb private collection D. Erber, Giessen cEss private collection J. Esser, Berlin cFel private collection B. Feldmann, Münster eGil private collection G. Gillerfors, Varberg cGon private collection A. Gontarenko, Odessa cKap private collection A. Kapp, Rankweil cKóh private collection F. Kóhler, Bornheim / cOro private collection P. Oromí, La Laguna cOwe private collection J. A. Owen, Epsom cRou private collection G. de Rougemont, Londinieres cSch private collection M. Schülke, Berlin cTro private collection M. Tronquet, Molitg-les-Bains cWun private collection P. Wunderle, Mónchen- gladbach cZan private collection A. Zanetti, Verona Head length was measured from the anterior margin of the clypeus to the posterior margin of the head, elytral length along suture from the apex of the scutellum to the elytral hind margin. 3. THE MEDON FAUNA OF THE WESTERN MEDITERRANEAN Based on an examination of 5600 specimens of Medon, including the accessible types, 26 valid species and sub- species are recognized, among them two revalidated and three previously undescribed species. The identities (and validity) of three additional species, M. augur Fau- vel, M. subterraneus Coiffait, and M. spelaeus (Scriba), remain doubtful; M. augur is represented only by a fe- male type specimen, and neither types nor non-type ma- terial of M. subterraneus and M. spelaeus were avail- able. As expected, the true diversity of the Medon fauna in the Western Mediterranean was found to be distinctly lower than indicated by COIFFAIT (1984). Altogether 21 new synonymies were established for the Medon fauna of the study area. This enormous degree of synonymy — involving ten now valid species and subspecies, one of them with as many as six new synonyms — explains the difficulties previously encountered when trying to iden- tify and interpret Medon species. In the Western Mediterranean, as defined above, the ge- nus 1s represented by at least four species groups. Since a phylogenetic analysis, which would require an inclu- sion also of Eastern Palaearctic species, has not yet been attempted, the species groups as outlined below must be considered tentative. The Medon apicalis group in- cludes four species (M. apicalis, M. perniger, M. sericellus, M. oromii) and is characterized by a head with relatively ill-defined puncturation, more or less pronounced microculpture, and without distinct shine, a microsculptured pronotum with fine puncturation, rather long elytra, a male sternite VII with weakly modified 28 Bonner zoologische Beiträge 54 (2005) marginal setae (no palisade setae), and either by an api- cally more or less incised or bifid ventral process (usual condition) or by an apically elongated apex (apomor- phic in M. oromii and M. sericellus) of the aedeagus. In the Eastern Mediterranean and Middle Asia, this species group is represented by the widespread M. maronitus (Saulcy), by M. beydaghensis Fagel (southwestern Ana- tolia), and M. bucharicus Bernhauer (Middle Asia). The only Western Mediterranean species of the group whose range extends into the west of the Eastern Mediterra- nean is M. apicalis. The species group previously referred to as the M. fer- rugineus group (ASSING 2004a) is represented in the Western Mediterranean by ten species, all of which are characterized by more or less dilated protarsi (at least in the male), the presence of two combs of palisade setae at the posterior margin of the male sternite VII, by an angled (ventral view) or a more or less deeply incised (ventral view) apex of the aedeagus. This species group includes Medon dilutus, M. subcoriaceus, M. antricola, M. castaneus, M. procerus, M. pocofer, M. rufiventris, M. africanus, M. vicentensis, and M. augur. Of these species, only the distributions of M. dilutus (only the subspecies M. dilutus pythonissa), M. pocofer, and M. rufiventris extend into the Eastern Mediterranean re- gion. Five species of the M. brunneus group occur in the Western Mediterranean: M. brunneus, M. piceus, M. cauchoisi, M. despectus, and M. mirei. Only one of them, M. brunneus, also occurs in the Eastern Mediter- ranean. The species of this group are characterized by a usually well-defined puncturation of the head and pronotum, unmodified protarsi, the presence of two combs of palisade setae, and by an apically truncate or rounded, unincised aedeagus (ventral view). The M. fusculus group, the species group with the high- est diversity in the Eastern Mediterranean, is repre- sented in the Western Mediterranean by only four spe- cies, the widespread Ponto-Mediterranean M. fusculus, the widespread Atlanto-Mediterranean M. ripicola, M. sardous from Sardinia, and M. kabylicus from Algeria. The systematic position of M. ripicola was previously unclear (ASSING, 2004a), but the respective character states in the evidently closely related M. sardous and M. kabylicus suggest that the distinctive morphology of the aedeagus represents a derived state of the condition usu- ally found in the M. fusculus group and the absence of additional long marginal setae at the posterior margin of the male sternite VII is a secondary reduction. Conse- quently, M. ripicola 1s here attributed to the M. fusculus group. SERRANO (1993) erroneously attributed also M. vicentensis to this group, which is characterized espe- cially by the shape and chaetotaxy of the. male sternite VII, the broad semicircular excision of the male sternite VIII, as well as by the morphology of the aedeagus; for details see ASSING (2004a). Two endemic species of the Atlantic Islands — M. indi- gena and M. feloi — may be monophyletic, both of them sharing the absence of palisade setae on the male ster- nite VII. This hypothesis would be supported also by a similar morphology of the aedeagus. Moreover, Medon indigena and M. feloi share a very similar shape of the male sternite VII, as well as a head and pronotum with sparse fine puncturation and with distinct microsculp- ture. Eleven species have very restricted distributions. These include the endemics of the Canary Islands and Madeira — M. indigena, M. subcoriaceus, M. antricola, M. vicen- tensis, M. feloi, and possibly also M. oromii —, M. sar- dous from Sardinia, the microphthalmous M. mirei from northern Algeria, the troglobiont M. procerus from cen- tral Spain, possibly also M. kabylicus and M. sericellus from Algeria. The question whether or not M. subterra- neus, M. spelaeus, and M. augur are valid endemic spe- cies will remain unanswered until material of these spe- cies becomes available for study and their taxonomic status is clarified. Of the remaining species, two are confined to Tunisia and Algeria: M. africanus and M. despectus, with the latter also occurring in the Italian is- land Pantelleria. As was to be expected, most of the more widespread (sub-)species of the Western Mediterranean are Atlanto- Mediterranean elements sensu LATTIN (1967): M. cau- choisi, M. ripicola, M. apicalis, M. ripicola, M. dilutus cephalus, M. piceus, and M. castaneus. Three (sub-) species are Adriato-Mediterranean elements (M. perni- ger, M. brunneus, and M. dilutus dilutus), and two have a Ponto-Mediterranean distribution (M. fusculus, M. rufiventris). More data are needed to decide if M. po- cofer represents an Atlanto- or an Adriato-Mediterranean element. Ecologically, the Medon fauna of the study area is highly diverse. Several species — all the species of the M. brunneus group, plus M. subcoriaceus, M. sardous, M. fusculus, and M. perniger — apparently inhabit the litter layer and are only occasionally encountered also in other habitats; the same may be true of several other species, but too little is known about their ecology. Some species such as M. castaneus and M. ripicola are evidently associated with subterranean mammal nests. Medon rufiventris is usually found in or under dead wood, though its real habitat is unknown. Others — M. procerus, M. feloi, M. antricola, M. vicentensis — are doubtlessly troglobionts. The habitats of several species, among them the widespread M. apicalis, which is usu- ally collected on the wing, are subterranean, but their exact nature is unknown. Volker ASSING: Western Palaearctic Medon 29 Below, only those species that have not already been treated by ASSING (2004a) are redescribed and illus- trated. Additional material of Medon species exclusively occurring in the region treated in the first part of the re- vision is compiled in a separate section at the end of the paper. In order to avoid redundancy, specimens already listed by ASSING (1998, 1999a, 2000, 2004a, 2004b) and ASSING & WUNDERLE (1999) are omitted from the material sections. In the references to the original de- scriptions, the original combination is given in the original spelling even when the gender of an adjectival name is incorrect (e. g. Medon pernigrum instead of M. perniger). 3.1. Medon apicalis (Kraatz, 1857) (Map 2) Lithocharis apicalis Kraatz, 1857 (KRAATZ 1857: 7151). Medon orgianum Bordoni, 1980 (BORDONI 1980: 83) syn. n. Types examined: L. apicalis: see ASSING (2004a). M. orgianus: Holotype 3: ITALIA CENTR. To: Or- gia (Si.) f. Merse, leg. Bordoni, 1X.1974 / HOLOTY- PUS / Medon orgianum n. sp. Det. A. Bordoni 1975 / Medon apicalis (Kraatz) det. V. Assing 2003 (cBor). Paratype Y: mounted on same pin as holotype with extra label “Allotype ...” (cBor). Comments: The holotype of M. orgianus is doubtlessly conspecific with M. apicalis. The synonymy of both names was independently recognized also by A. BOR- DONI (Firenze, pers. comm.) and A. ZANETTI (Verona, pers. comm.). Additional material examined (total: 461 exs.; for ad- ditional material see ASSING (2004a)): Morocco: 2 exs., Haut Atlas, NE Tizi-n-Test, 30°54N, 08°19W, 1710 m, edge of stream, 29.X11.2002, leg. As- sing, Wunderle (cAss, cWun); 1 ex., NE Tizi-n-Test, 30°55N, 08°17W, 1150 m, 29.X11.2002, leg. Wunderle (eWun); 1 ex., 40 km N Oulmes, 27.11.1983, leg. Le- doux (cTro). Portugal: | ex., Lisboa, Serra do Sintra, 24.V.1992, leg. Wunderle (cWun); | ex., Algarve, N Tavira, Alca- ria do Cume, 37°15N, 07°44W, 440 m, 12.1V.2002, leg. Meybohm (cAss); | ex., Algarve, Serra de Mon- chique, NW Foia, 37°19N, 08°36W, 780 m, 15.IV. 2002, leg. Meybohm (cAss); 3 exs., Algarve, Caldas de Monchique, V.-VI.1960, leg. Fagel (IRSNB). Ma- deira: | ex., Canico de Baixo, window, 20.- 27.1X.1990, leg. Pieper (cAss); 5 exs., same data, but 7.-20.V.1992 (cAss); 1 ex., same data, but 14.- 20.1X.1989 (cAss); 3 exs., Santo da Serra, 2. VIII.1975, leg. Vit (MHNG); 2 exs., Camacha, Lavada da Serra, 13.V111.1975, leg. Vit (MHNG). Spain: Galicia: | ex., Lugo, Sierra de Ancares, Tres Obispos, VI.1997 (cFel). Asturias: 12 exs., W Puerto de Pajares, Puerto de la Cubilla — Sotiello, 500 - 1400 m, car-net, 8.V1.1991, leg. Wunderle (cWun). Canta- bria: 1 ex., Espinama, 27.V11.1972, leg. Lohse (cBor). Cataluña: | ex., Gerona, leg. Franz (NHMW); 1 ex., Gerona, Playa de Aro, 25.V.-5.V1.1971 (cBor); 12 exs., Seo de Urgel, Arfa, V.-VI.1962, leg.Fagel (IRSNB); 3 exs., Seo de Urgel, Sierra del Cadi, 1000 m, V.- V1.1962, leg. Fagel (IRSNB). Aragón: | ex., WNW Te- ruel, Sierra de Albarracín, NE Noguera, 40°30N, 01°35W, 1625 m, 11.1V.2003, leg. Assing (cAss). Ma- drid: | ex., Sierra de Guadarrama, El Escorial, leg. Franz (NHMW). Valencia: 10 exs., Alicante, Rio Sella, 25.111.1978 (cAss); 1 ex., Alicante, El Penon, leg. Franz (NHMW); 2 exs., Castellón, Cálig near Benicarlo, 6.V.1966, leg. Besuchet (MHNG). Castilla-La Man- cha: 12 exs., SE Manzanares, Alhambra, 2.V.1925, leg. Scheerpeltz (NHMW); 1 ex., Albacete, Villaverde, 13.1V.1959, leg. Besuchet (MNHG). Andalucía: | ex., Sierra Nevada, Rio Genil, 37°08N, 03°24W, 1000 m, 24.11.2000, leg. Lompe (cAss); 2 exs., Sierra Nevada, Lanjaron, 26.1V.-18.V.1961, leg. Fagel (IRSNB); 1 ex., Puerto de Galis, 36°32N, 05°38W, 400 m, 20.11.2000, leg. Meybohm (cAss); | ex., Granada, Cenes near Gra- nada, 24.V.1966, leg. Besuchet (MHNG ); | ex., Algeci- ras env., Rio Guadarranque, 36°15N, 05°26W, flood plain forest, 21.11.2000, leg. Meybohm (cAss); 1 ex., Algeciras env., Sierra de Luna, 350 m, 28.111.1994, leg. Assing (cAss); 6 exs., Algeciras env., Sierra de Fates, 350 m, 28.111.1994, leg. Assing, Wunderle (cAss, cWun); 2 exs., Sierra de Bermeja (CA), Jubrique, 500 m, 26.111.1994, leg. Assing, Wunderle (cAss, cWun); | ex., Sierra de Bermeja (MA), Puerto de Peñas Blancas, 1200 m, 26.111.1994, leg. Wunderle (cWun); | ex., Cerro de la Novia, barranco at km 6 A-375, 36°37N, 05°30W, 700 m, 19.11.2000, leg. Lompe (cAss); 1 ex., Algeciras, Cer- ro de Mirador, leg. Franz (NHMW); | ex., Vejer de la Frontera (CA), 27.111.1987, leg. Zoia (cZan); 2 exs., Cordoba, Villaviciosa, 29.V1.1969, leg. Comellini (MHNG); 1 ex., Andújar, V.1959, leg. Comellini (cAss); | ex., E Jaén, SE Mancha Real, Sierra Almadén, ca. 37°45N, 03°33W, 1450-1850 m, ©. ilex litter, 26.X11.2003, leg. Assing (cAss). Baleares: 2 exs., Ibiza, Sierra Grosa, leg. Franz (NHMW, cAss); 1 ex., Ibiza, locality not specified, leg. Franz (NHMW). Islas Cana- rias: Gran Canaria: | ex., Bco. de la Miña, 1200 m, 21.X11.1997, leg. Assing (cAss); 1 ex., Bco. de la Vir- gen, 500 m, 20.X11.1997, leg. Wunderle (cWun); 2 exs., N Teror, Osorio, 600 m, 20.X11.1997, leg. Wunderle (cWun). Tenerife: 3 exs., Teno, Las Portelas, 700 m, 30.X1.1996, leg. Schiilke (cSch, cAss). France: Aquitaine: | ex., Landes, Saint-Sever-sur- l"Adour, 11.1X.1915 (cSch); 1 ex., Gironde, Léognan, VIII.1943, leg. Tempere (MHNG); 8 exs., Cap-Ferret, 30 Bonner zoologische Beitráge 54 (2005) 17.1X.1928, leg. Tempere (MHNG); 4 exs., Gironde, Cambes, 17.11.1939, leg. Tempere (MHNG); 4 exs., Cambes, 10.1V.1939, leg. Giraud (MHNG); 1 ex., Gi- ronde, Pessac, 5.V1.1928, leg. Tempere (MHNG). Midi- Pyrénées: | ex., Ariege, Foix, 3.1V.1953, leg. Tempere (MHNG); 4 exs., Hautes-Pyrenées, locality not speci- fied, leg. Pandellé (NHMW). Picardie: | ex., Laigne- ville (Oise), VII.1908 (MHNG). lle-de-France: 2 eXS., Cheptainville, 5.VHI.1945, leg. Levasseur (MHNG). Centre: | ex., Orléans (cBor); | ex., Perrusson, V.1905, leg. Mequignon (MHNG); 8 exs., Perrusson, XII.1903 (MHNG); | ex., same data, but 111.1906 (MHNG); 11 exs., Perrusson, leg. Mequignon (MHNG); 2 exs., Loches (MHNG). Rhóne-Alpes: | ex., Vaugneray (MHNG). Provence: 9 exs., Var, Tanneron, Auribeaux — Lac de S. Cassien, car-net, 30.-31.V.1991, leg. Wunderle (cWun); l ex., Var, Tanneron, car-net, V1.1988, leg. Wunderle (cWun); 5 exs., Var, Plan du Var [43°52N, 7°12E] (MHNG ); 4 exs., Plan du Var, 1.1954 (MHNG); 1 ex. [teneral], Ollioules, 15.X.1973 (MHNG); 1 ex., Var inundation (W Nice), 111.1951 (MHNG); 1 ex., Alpes Maritimes, Sophia Antipolis near Antibes, haystack, 16.X.1991, leg. Schiilke (cSch); 1 ex., Alpes Maritimes, Breil-sur-Roya, 1.V1.1955 (MHNG); | ex., Alpes Mari- times, “Gr. Dozol”, X.1937 (MHNG); 1 ex., Alpes Ma- ritimes, St.-Vallier-de-Thiey, V.1975, leg. Toumayeff (MHNG); 4 exs., Var, Bras, IV.1971, leg. Tronquet (cTro); 7 exs., Var, Gonfaron, chestnut litter, 1V.1971, leg. Tronquet (cTro); 1 ex., Var, Vésubie inundation [near Lantosque], V.1961 (MHNG); 4 exs., Vaucluse, Mont Ventoux, 17.V1.1977, leg. Tronquet (cTro). Cor- sica: | ex., SE Corte, 200 - 900 m, car-net, 8.1V.1990, leg. Assing (cAss); 3 exs., Cursegliente-Pietroserana- Corte, 8.1V.1990, leg. Wunderle (cWun); | ex., Erbajo- lo-Bastinaco-Curtisi, car-net, 12.1V.1990, leg. Wunderle (cWun); 1 ex., Valle di Restonica, 800 - 1000 m, 6.- 13.1V.1990, leg. Wunderle (cWun); 2 exs., NW St. Flo- rent, Désert des Agriates, car-net, 21.1V.2001, leg. Za- netti (cZan); 1 ex., Casta, Campu Castingu, 300m, 18.V11.1976, leg. Sette (cZan); 1 ex., San Nicolao, 270 m, leaf litter, 18.V11.1994, leg. Zerche (DEI); 4 exs., Porto Vecchio, caught flying, V.1971, leg. Tronquet (cTro); 12 exs., Bocognano, caught flying, 28.V.1972, leg. Tronquet (cTro); 1 ex., Bocognano, caught flying, 26.V.1987, leg. Tronquet (cTro); 1 ex., Bocognano, 1905, leg. Leonhard (DEI); 2 exs., Cateraggio, caught flying, 1.V1.1987, leg. Tronquet (cTro); 1 ex., Francar- do, VII.1954 (MHNG); 1 ex., Porto, V.1966 (MHNG); l ex., E Col de Vergio, Ft. Valdo-Niello [42°17N, 08%54E], caught flying, 25.V.1972, leg. Tronquet (cTro); 2 exs., locality not specified (DEI). Locality ambiguous or not specified: | ex., Prunelli, V.1960 (MHNG); 1 ex., “France”, leg. Mulsant (MHNG). England: | ex., Norwich, Norfolk, reed litter, 12.1V.1989, leg. Owen (cOwe). Sweden: Gotland: | ex., Mästerby, 21.V1.1965, leg. Gillerfors (cGil); 1 ex., Vall, 8.VIII.1976, leg. Gillerfors (cGil); 3 exs., Buttle, 7.V111.1976, leg. Gillerfors (cGil):; 4 exs., Fide, 1.VIII.1975, leg. Gillerfors (cGil). Skane: 4 exs., Stenshuvud, 19.111.1977, leg. Gillerfors (cGil). Jämtland: | ex., Svenstavik, 22.V1.1989, leg. Gillerfors (cGil). Halland: 1 ex., Hunnestad, 5.V.1978, leg. Gil- lerfors (cGil); 2 exs., Hunnestad, 4.V1.1988, leg. Giller- fors (cGil); 1 ex., Dagsás, 6.V1.1971, leg. Gillerfors (cGil); 1 ex., Dagsás, 9.1X.1977, leg. Gillerfors (cGil); 2 exs., Dagsás, 1.V1.1979, leg. Gillerfors (cGil). Sma- land: 1 ex., Ruggstorp, 11.V111.1978, leg. Gillerfors (Gil). Pite Lappmark: 3 exs., Moskosel, 13.V1.1976, leg. Gillerfors (cGil). Lule Lappmark: | ex., Messaure, 23.V11.1978, leg. Gillerfors (cGil). Germany: Nordrhein-Westfalen: | ex., Kreis Stein- furt, NSG Heiliges Meer, car-net, 8.V.1998, leg. Feld- mann (cFel); I ex., Kreis Steinfurt, NSG Emsdettener Venn, car-net, 7.V1.1996, leg. Terlutter (cFel); 2 exs., Brüggen, Holter Heide, car-net, 19.V1.1995, leg. Wun- derle (cWun); 3 exs., Düren, Gürzenich-Schevenhütte, car-net, 26.V.1989, leg. Wunderle (cWun); 3 exs., Jü- lich, Hambach, Lindenberger Wald, 2.VII.1991, leg. Köhler (cKóh); 4 exs., Bonn, Róttgen, Kottenforst, 3.VII.1991, leg. Köhler (cKöh); 2 exs., Brühl, Staats- forst Ville, 30.1V.1991, leg. Köhler (cKöh); 2 exs., Ei- fel, Gemünd, Der Kermeter, 6.V.1990, leg. Köhler (cKóh). Rheinland-Pfalz: 2 exs., Ulmen-Vollmerath, car-net, 9.V1.1983, leg. Wunderle (cWun); 1 ex., Stau- dernheim, 24.1.1995, leg. Köhler (cKöh); 2 exs., Mon- zingen, 18.-19.V.1991, leg. Köhler '(cKöh); 1. -ex.; Schaidt, Bienwald, NWR Stuttpferch, 30.V.1996, leg. Köhler (cKöh); 2 exs., Bacharach, Steeg, 5.VI.1991, leg. Köhler (cKöh); 2 exs., Schloßböckelheim, 31.V.1996, leg. Köhler (cKöh); 1 ex., Taben/Saar, car- net, 6.-8.V1.1996, leg. Köhler (cKöh). Schleswig- Holstein: | ex., Eutin, 1.V1.1982, leg. Lohse (MHNG); l ex., Eutin, 1.V111.1984, leg. Lohse (MHNG); 1 ex., Tensfeld, leg. Lohse (MHNG); 1 ex., Güster, 24.V.1958, leg. Lohse (MHNG); 1 ex., Güster, 7.V11.1977, leg. Lohse (MHNG); 2 exs.“ Farchau” (?), 13.V.1981, leg. Lohse (MHNG); 1 ex., Hornbek, 21.V1.1979, leg. Lohse (MHNG ); l ex., Koberg, 14.V.1979, leg. Lohse (MHNG); 6 exs., Sachsenwald, 23.V1.1977, leg. Lohse (MHNG); 2 exs., Sachsenwald, 30.VII.1961, leg. Lohse (MHNG). Ham- burg: | ex., Ohlstedt, 3.V1.1961, leg. Lohse (MHNG). Niedersachsen: 1 ex., Neustadt/Rbg., Himmelreich, window trap, 7.VII.1989, leg. Schmidt (cAss); 5 exs., Hannover, Eilenriede, car-net, 28.-29.V.1991, leg. As- sing (cAss); 1 ex., same data, but 8.VII.1991 (cAss); 1 ex., Liineburger Heide, Schneverdingen, Tiitsberg, pit- fall trap, 17.V1.1997 (cAss); 2 exs., Kreis Lüneburg, Amt Neuhaus/Elbe, car-net, 6.VII.1995, leg. Renner (cFel); 5 exs., Liichow-Dannenberg, car-net, 1.V1.1983, leg. Renner (cWun); 1 ex., Lüchow-Dannenberg, Gar- Volker ASSING: Western Palaearctic Medon 3] tower Forst, 13.V1.1987, leg. Wunderle (cWun); 1 ex., Göhrde, 15.V.1983, leg. Lohse (MHNG); 4 exs., Göhr- de, 8.V1.1977, leg. Lohse (MHNG); 2 ex., Göhrde, 21.V.1981, leg. Lohse (MHNG); 4 exs., Forst Göhrde, 1.V1.1983, leg. Lohse (MHNG); 4 exs., Forst Göhrde, 26.V11.1980, leg. Lohse (MHNG); 8 exs., Forst Göhrde, 6.V1.1980, leg. Lohse (MHNG); 1 ex., Wirl near Gar- tow, 6.V11.1983, leg. Lohse (MHNG); 1 ex., Lüchow- Dannenberg, Höhbeck, 1.V.1978, leg. Lohse (MHNG); l ex., same data, but 26.V.1979, leg. Lohse (MHNG); 1 ex., Lüchow-Dannenberg (cKöh); 2 ex., Uelzen, Bram- bostel, 2.V1.1991, leg. Lohse (MHNG). Hessen: 2 exs., Vogelsberg, Ehringhausen, car-net, V.1987, leg. Wun- derle (cWun); 1 ex., Dautphe-Buchenau, Katzenbachtal, car-net, 26.1V.1987, leg. Wunderle (cAss). Ber- lin/Brandenburg: 19 exs., Berlin, Hohen Neuendorf (DEI). Sachsen: 3 exs., Guttau, car-net, 25.V.1985, leg. Schúlke & Heinig (cSch); 3 exs., Oberlausitz, Daubitz, haystack, 23.1V.1994, leg. Schülke (cSch, cAss); 1 ex., Weißwasser, car-net, 17.V.1984, leg. Apel (DEI). Austria: Niederósterreich/Wien: 11 exs., several loca- lities in the surroundings of Wien (NHMW); 7 exs., Wienerwald, Hadersdorf, leg. Skalitzky (MHNG, NHMW); 5 exs., Wien, Pötzleinsdorfer Park (MHNG). Kärnten: | ex., Gurktaler Alpen, Falkert, car-net, 24.V11.1991, leg. Assing (cAss); 4 exs., Eisenkappel, car-net, 26.V1.-4.V11.1987, leg. Siede & Wunderle (cWun). Steiermark: | ex., Pleschkogel near Rein, 590 m, sifted forest litter, 20.V1.1995, leg. Zerche & Behne (DEI). Italy: Liguria: 1 ex., San Remo, leg. Schneider (DEI); 2 exs., Genova, Premanico, 17.X11.1972, leg. Bartoli (UCBA); 2 exs., Casanova di St. Olcese (GE), 26.11.1972, leg. Bartoni (UCBA). Friuli-Venezia Giu- lia: 1 ex., Rupin Grande (TS), car-net, 15.V.1994, leg. Zanetti (cZan). Veneto: | ex., Tregnago — Vestenavec- chia, car-net, 20.1V.1996, leg. Zanetti (cZan). Toscana: 8 exs., E Arezzo, Lippiano near Padonchia, X1.1924, leg. Andreini (UCBA, cBor); | ex., San Rossore (Pisa), 30.1.1969, leg. Santini (cBor); 1 ex., Padule di Fucce- chio, leg. Castellini (cTro). Elba: 3 exs., leg. Paganetti (DEL cAss). Sardegna: | ex., NE Lula (NU), 600 m, 23.1V.1992, leg. Scheuern (cWun); 1 ex., NE Lula, 600 m, 23.1V.1992, leg. Schawaller (cSch); 1 ex., 10 km NNW Nuoro, Grotta Cuccu, Monte Tiria, 27.V111.1994 (cBor); 3 exs., 5 km SW Galtelli (NU), light source, 16.V11.1999, leg. Zanetti (cZan); 1 ex., Galtelli, car-net, 17.V11.1999, leg. Zanetti (cZan); 1 ex., locality not spe- cified (DEI). Sicilia: 1 ex., M. Iblei, Cava Gissara, Me- lilli (SR), 300 m, 6.VI.1991, leg. Sabella (UCBA); 2 exs., Taormina (ME), Isola Bella, 8.X.2002 (UCBA); 4 exs. same locality, 7.X1.2002 (UCBA, cAss). Slovenia: | ex., Savina, leg. Paganetti (MHNG). / Locality not indicated: 3 exs. (MHNG). Diagnosis: See ASSING (2004a) and the key in section 4. Distribution and bionomics: Medon apicalis is wide- spread in the western parts of the Western Palaearctic region (Map 2). It occurs in the Canaries (Tenerife and Gran Canaria), Madeira, and Morocco; in continental Europe, its distribution ranges from the Iberian Penin- sula in the southwest to England and Scandinavia in the north, and to the northern Balkans in the southeast (ma- terial examined). It has also been reported from the Azores (BERNHAUER 1940; GILLERFORS 1988; LUND- BLAD 1958; SMETANA 1970). In Scandinavia it is the most widespread and least rare species of the genus; in Sweden it has been recorded as far north as Lule Lappmark and Norrbotten (material examined; LUNDBERG 1995). Medon apicalis is, however, unknown from the Baltic countries (SILFVERBERG 1992). In the British Isles, only few localities are known from southern England (FOWLER 1888; JoY 1932). NORMAND (1935) records M. apicalis also from Tunisia, but these records may re- fer to similar species (M. ripicola, M. kabylicus, M. sericellus); | have seen no material from there. For a se- lection of additional literature records see BOHAC (1985), BRUGE et al. (2001), DRUGMAND (1989), FOW- LER (1888), M. HANSEN (1996), HANSEN et al. (1994, 1995), V. HANSEN (1964), HORION (1965), HU- GENTHOBLER (1966), KOCH (1968), KÓHLER (2000), KOHLER € KLAUSNITZER (1998), LUNDBERG (1995), SILFVERBERG (1992), TERLUTTER (1995), and TRON- QUET (2001). In the southern Balkans, Anatolia, and the Caucasus region, M. apicalis is replaced by M. maroni- tus (Saulcy) (ASSING 2004a). In Italy, where the closely related M. perniger is much more abundant, M. apicalis has been found in the north, in Toscana, and in the is- lands Elba, Sardegna, and Sicilia. In some Italian locali- ties and in the Alpes Maritimes, M. apicalis and M. per- niger were apparently collected together: Toscana (Lippiano, San Rossore), Liguria (Premanico, Casanova di St. Olcese (GE)), Sicilia (Isola Bella), Alpes Mari- times (Antibes). All previous literature records of M. apicalis from Italy must be considered doubtful due to probable confusion with M. perniger. Bonner zoologische Beitrage 54 (2005) aa / Map 2: Distributions of Medon apicalis (Kraatz) (filled circles: revised records; open circles: selected unexamined records) and M. sericellus (Fairmaire) (filled squares, examined records). Volker ASSING: Western Palaearctic Medon 33 The species has been collected in various biotopes and habitats. In the material examined, the vast majority of specimens collected in spring and early summer (April through July) were taken on the wing (mostly by car- net, but also by flight interception traps). Ilpo RUTANEN (Hyvinkaa, pers. comm. 2003) communicated to me 22 records from southern Finland, 21 of which were made with a car-net (June through August). The remainder of the specimens examined were found in various types of woodland, at the edge of streams, under haystacks, or at light sources. Some of the records from early spring are apparently from flood debris. The reproduction habitat of M. apicalis is cryptic and unknown. According to HORION (1965) and KOCH (1968), the species is possi- bly associated with nests and burrows of voles and mice. NOWOSAD (2000), however, examined 7000 nests of various small mammals and birds, but found M. api- calis only occasionally. All the records of M. apicalis from mole nests by OSELLA & ZANETTI (1974) refer to M. perniger. Adult beetles have been recorded through- out the year; teneral specimens were observed in au- tumn (October). DRUGMAND (1989) states that the spe- cies is bivoltine, but there is no evidence supporting this. Based on the material examined here, M. apicalis is univoltine. 3.2. Medon perniger Coiffait, 1978 (Figs. 1-7, Map 3) Medon apicalis var. pernigrum Coiffait, 1970a (COIF- FAIT 1970a: 717); unavailable name (infrasubspecific). Medon pernigrum Coiffait, 1978 (COIFFAIT 1978: 268f). Medon pernigrum fraudulentum Coiffait, 1978 (COIF- FAIT 1978: 269f) syn. n. Types examined: M. perniger: Holotype $: Emilie, Parme / MUSEUM PARIS COLL H. COIFFAIT / HO- LOTYPE / Medon pernigrum H. COIFFAIT 1963 / Ho- lotypus Medon perniger Coiffait rev. V. Assing 2003 (MNHNP). M. perniger fraudulentus: Paratype d: d / Sicilien, Messina, April 43. / leg. Wolfrum / Medon apicalis det. E. Ulbrich 1966 / MUSEUM PARIS COLL H. COIF- FAIT / PARATYPE / Edeage casse / Medon pernigrum fraudulentum H. COIFFAIT 1978 [?] / Medon perniger det. V. Assing 2003 (MNHNP). Comments: The holotype of M. perniger is a relatively dark specimen. The main difference between M. perni- ger and M. p. fraudulentus indicated by COIFFAIT (1978) is the lighter coloration of the latter. This colora- tion, however, is clearly within the range of intraspecific variation throughout the range of the species. No sig- nificant differences could be found in the male sexual characters, so that M. p. fraudulentus is here placed in the synonymy of M. perniger. Additional material examined (total: 298 exs.): France: | ex., Alpes Maritimes, Sophia Antipolis near Antibes, haystack, 18.X.1991, leg. Schúlke (cSch); 1 ex., Var, Argens inundation [43°27N, O06°30E], 17.1.1955 (MHNG); 1 ex., Var, Vésubie inundatign [near Lantosque], V.1961 (MHNG); | ex., Alpes Mari- times, “St. Pons”, VI.1956 (MHNG); 1 ex., Alpes Mari- times, Menton, inundation, V.1952 (MHNG). Switzerland: 2 exs., Ticino, Locarno, VIII.1944 [“Ty- pus Medon linderi O. Scheerpeltz”; manuscript name] (NHMW). Italy: Trentino-Alto Adige: | ex., Trento, Villa Laga- rina, 950 m, 29.1V.1962, leg. Petrolli (cBor). Piemonte: 2 exs., Cuneo, Colle San Bernardo, N Erli, 600 m, 12.X.1997, leg.Wolf (cSch); 1 ex., locality not speci- fied, leg. Baudi (DEI). Lombardia: | ex., Alpi Orobi, Morbegno, 350 m, deciduous forest, 18.VIII.1994, leg. Assing (cAss); 1 ex., Zecchone near Pavia, 17.X.1977, leg. Kanaar (cSch); 2 exs., Rivolta d’Adda, 23.X.1959, leg. Rosa (NHMW, cAss); 1 ex., Ostiglia (MN), Palude del Busatello, 7.111.1982, leg. Zanetti (cZan); 10 exs., Mantova, Bosco Fontana, 27.1.1980 (5 exs.), 7.V.1989 (1.6%) 11.91.1989. (1%%.), 12.X, 1989 (3 eXs:), leg; Za- netti (cZan); 1 ex., Torbiere d'Iseo (BS), 28.111.1982, leg. Zanetti (cZan); 4 exs., Castellaro Lagusello (MN), 2.X11.1990, leg. Zanetti (cZan); 1 ex., Pozzolo (MN), 1:IV.1990, leg. Zanetti (cZan); 1 ex., Oltre il Colle, 18.V1.1914 (DED; 1 ex., Corana (PV), 12.1V.1970, leg. Briganti (UCBA); 1 ex., Bereguardo, 2.V.1943, leg. Paganetti (MHNG). Veneto: 1 ex., Arcole (VR), 12.111.1971, leg. Pace (cBor); 3 exs., Sona (VR), 1.111.1970, leg. Zanetti (cZan); 1 ex., Valli Grandi Ve- ronesi, La Torretta [45°06N, I1°16E], leg. Zanetti (cZan); 14 exs., San Giovanni Lupatoto (VR), mole nests, 28.1X.1973, leg. Zanetti (cZan, cAss); 3 exs., sa- me data, but 16.X1.1973 (UCBA, cZan); 1 ex., same lo- cality, “base di Salix”, 2.11.1975, leg. Zanetti (cZan); 4 exs., NW Villafranca di Verona, Custoza, mole nests, 10.11.1974, leg. Zanetti (cZan); 1 ex., Custoza, 24.X1.1974, leg. Zanetti (cZan); 1 ex., Bussolengo — Sommacampagna (VR), car-net, 24.V1.1989, leg. Zanet- ti (cZan); 2 exs., Verona, P.te Saval, 26.11.1974, leg. Zanetti (cZan); 3 exs., Montecchia di Crosara (VR), mo- le nest, 22.11.1976, leg. Zanetti (cZan); 2 exs., Soave — Montecchia d. C., car-net, 20.1V.1996, leg. Zanetti (cZan); 1 ex., San Martino Buon Albergo (VR), Musella Ferrazzetta, mole nest, 28.11.1991, leg. Zanetti (cZan); 1 ex., Pastrengo (VR), 20.1V.1984, leg. Zanetti (cZan); 1 ex., Pellegrina (VR), mole nest, 3.11.1980, leg. Zanetti (cZan); 1 ex., Tregnago (VR), 26.X11.1976, leg. Zanetti (cZan); 2 exs., Tregnago — Vestenavecchia, car-net, 20.1V.1996, leg. Zanetti (cZan); 1 ex., Negrar (VR), Montericco, light source, 1V.2002, leg. Zanetti (cZan). Emilia-Romagna: 4 exs., Sant Agostino (FE), Bosco 34 Bonner zoologische Beitráge 54 (2005) Panfilia, 20 m, nest of Apodemus, 23.11.1997, leg. Ta- ghapietra & Zanetti (cZan); 2 exs., Valle Scoltenna (N Sestola) (MO), car-net. 1.V1.1996, leg. Zanetti (cZan). Friuli-Venezia Giulia: | ex., Udine, Chiarpena, 200 m, 20.X.2001, leg. Kahlen (cAss). Liguria: | ex., ca. 30 km NW La Spezia, Zignago, V11.1962 (cBor); 3 exs., La Spezia, Le Grazie, 5.X11.1982, leg. Zoia (cZan); 5 exs., Genova, Ruta, leg. Dodero (NHMW, cAss); 1 ex., Ge- nova, Premanico, 17.X11.1972, leg. Bartoli (UCBA); 1 ex., Neirone (GE), S. Marco d’Ursi, 30.XII.1982, leg. Zoia (cZan); | ex., Bordighera (DEI); 1 ex., La Spezia, Déiva Marina near Piazza, 5.1V.1970, leg. Briganti (U- CBA); 2 ex., Graveglia (GE), Pian di Fieno, 19.11.1976, leg. Zoia (UCBA); 6 exs., Casanova di St. Olcese (GE), 18.1V.1973, leg. Bartoli (UCBA); | ex., same data, but 26.111.1972 (UCBA y; 34 exs., Genova (citta), Villetta Di Negra, 1.&8.X.1972, leg. Bartoli (UCBA); 2 exs., same data, but 7.V.1975 (UCBA); | ex., 10 km NE Genova, Creto, 2.V.1977, leg. Parodi (UCBA); I ex., Genova, V.1920, leg. Mancini (UCBA); 1 ex., Genova, leg. Nat- terer (MHNG); 3 exs., Arenzano (GE), 6.X1.1977, leg. Gardini (UCBA); | ex., Ameglia, 1X.1913, leg. Mancini (UCBA). Toscana: | ex., Livorno, Tenuta di Tombola, 11.1V.1992, leg. Schawaller (SMNS); 1 ex., SW Firen- ze, Figline d’Arno, 200 m, 14.V1.1992, leg. Assing (cAss); | ex., Vallombrosa, Regelle, 750 m, 28.1V.1991, leg. Wunderle (cWun); 7 exs., Grassina (FI), 350 m, flood debris, 3.V.1991, leg. Wunderle (cWun, cAss); 2 exs., Mt. di Calvanı (FI), Mungona, 650 m, 1.V.1991, leg. Wunderle (cWun); 1 ex., Cecina, 4.1X.1972, leg. Pace (cBor); 3 exs., San Rossore (PI), 22.&30.1.1969, leg. Santini (cBor); 2 exs., Tana Termini (?) (Lucca), V1.1930, leg. Andreini (cBor); 5 exs., Arezzo, inunda- tion of Arno river, X1.1926, leg. Andreini (cBor, cAss); 1 ex., Arezzo, Palazzo del Pero, 9.1V.1961 (cBor); 5 exs., E Arezzo, Lippiano near Padonchia, X1.1924, leg. Andreini (UCBA, cBor); 4 exs., Firenze, 21.1.1962, leg. Castellini (cBor); 7 exs., 15 km ESE Massa, Volegno, V.1967, leg. Bordoni (cBor); I ex., Piteccio, 21.1V.1969, leg. Failla (cBor); 1 ex., N Borgo San Lorenzo, Grezza- no, 27.X.1974, leg. Rocchi (cBor); 2 exs., Ponteginori (SD), 4.1V.1994, leg. Gardini (cZan); 1 ex., Uccellina, Collelungo [42°37N, 11%05E], light-source, 12.V1.1977, leg. Zanetti (cZan); 4 exs., Antona, 1V.1911, leg. Man- cini (UCBA); I ex., Rignano sull’Arno, V.1974, leg. Castellini (cTro). Umbria: | ex., Sigillo (PE), M. Cuc- co, 20.1V.1975, leg. Rossi (cZan). Lazio: 2 exs., Mte. Circeo, 300 - 450 m, Quercus ilex forest, 27.X11.1994, leg. Assing (cAss); 4 exs., Mte. Circeo, mixed forest, 29.X11.1994, leg. Assing (cAss); 1 ex., Itri, 400m, 25.1V.1977, leg. Zampetti (cZan). Abruzzo: | ex., Cap- pelle sul Tavo (PE), 13.V11.1974, leg. Zanetti (cZan); | ex., Fiume Sinello (CH), Bosco di Don Venanzo [41°55N, 14°26E], car-net, 23.V11.1998, leg. Zanetti (cZan); 1 ex., Maiella, Madonna della Mazza, 1020 m, 28.V.2003, leg. Kapp (cKap). Puglia: | ex., Gargano, Forest Umbra, 600 m, beech forest, 30.XII.1994, leg. Assing (cAss); 2 exs., Foresta Umbra (FG), 800m, 6.V.1982, leg. Angelini (cZan). Campania: 10 exs., Napoli, Sorrent, S. Agata su due Golfi, 40°37N, 14°23E, 430 m, chestnut forest, 12.V.2002, leg. Wun- derle (cAss); 2 exs., 5 km NE Salerno, Mte. Stella, leg. Liebmann (cBor). Basilicata: 2 exs., Lucania, Nova Siri (MT), 500m, 29.1V.1979, leg. Angelini (cRou); 1 ex., E Lagonegro, Mt. Sirino, 40°06N, 15°51E, 7.V.2002, leg. Wunderle (cWun); 4 exs., Maratea, 40°01N, 15°44E, 485 m, 12.V.2002, leg. Wunderle (cWun); 1 ex., Rivel- lo, 400 m, oak forest, 6.V.2002, leg. Wunderle (cWun); 1 ex., Taranto, Policoro near Scanzano/Jonico, 22.X.2000, leg. Wolf (cSch); 1 ex., Policoro (MT), 1.V11.1982, leg. Angelini (cZan). Calabria: 10 exs., Aspromonte, dint. San Luca, 300 m, 24.V1.1987, leg. Angelini (cAss); 1 ex., S. Eufemia d’Aspromonte, 31.V11.1975, leg. Pace (cBor); 1 ¡ex AS Eufemia: 10.V11.1997, leg. Adorno (UCBA); 2 exs., S. Eufemia, 31.V1.1993 (UCBA); 1 ex., S Cittanova, Zómaro, 12.V11.1988 (cZan); 2 exs., Antonimina, 1.V1.1993 (U- CBA); 2 exs., Antonimina, mixed forest, 3.V.1993 (U- CBA). Sardegna: 1 ex., Siniscola, light source, 19.1X.1987 (cAss); 2 exs., Salto di Quirra, 400 m, 9.X.1989, leg. Wunderle (cWun); 2 exs., Iglésias, Grotta del Torpedo (?), 18.X1.1970 (cBor). Sicilia: 8 exs., San Fratello, 750 m, oakwood, 30.V1.1978, leg. Pace (cZan, cAss) 2 exs., Lascari — Gratteri (PA), car-net, 7.V11.2000, leg. Zanetti (cZan); 1 ex., Noto Antica, 16.- 17.V11.1991, leg. Zanetti (cZan); 1 ex., Macchia (CT) (UCBA); l ex., Aci Sant’Antonio (CT), 14.V.1992 (U- CBA); 1 ex., Peloritani, Foresta di Ferra, Santa Lucia del Mela (ME), 570 m, 6.V1.1996, leg. Adorno (U- CBA); 3 exs., Messina, leg. Vitale (UCBA); 3 exs., Messina, 6.111.1943 (NHMW); 1 ex., locality illegible, leg. Ragusa (UCBA); I ex., N Mongiuffi Melia, Límina, 14.V.1993 (UCBA); 1 ex., Gerace, Zomino, 15.X1.1993 (UCBA); 1 ex., Taormina (ME), Isola Bella, 8.X.2002 (UCBA); 3 exs., same locality, 7.X1.2002 (UCBA, cAss); 1 ex., M. Iblei, Cassaro (SR), bank of Anapo ri- ver, 5.111.1997 (UCBA); 1 ex., Buccheri (SR), Bosco di Accoro, Valle Cupa, 1.X1.1996 (UCBA). Locality doubtful, illegible, or not indicated: 4 exs., “Alta Valle Tosco-Umbra”, inundation of Tevere river, 18.X1.1935, leg. Andreini (cBor); 1 ex., leg. Parodi (UCBA); 2 exs., without locality (UCBA); 2 exs. [con- fusion of locality labels?], Turkmenistan, Lotfabad [“Ljutfabad”] (MNHUB, cAss). Diagnosis: In general appearance highly similar to M. apicalis (Fig. 1), distinguished as follows: Coloration on average (!) slightly darker. Puncturation of head and pronotum on average slightly coarser and slightly more well defined. Volker ASSING: Western Palaearctic Medon 35 +: posterior margin of sternite VII in the middle with dis- tinct convex projection (Figs. 2-3) (in M. apicalis weakly convex) and, especially in the middle, with darker, stouter, and longer modified setae; sternite VIII as in Fig. 4; aedeagus with ventral process of distinctive morphol- ogy: deeply bifid, apices widely separated, and subapi- cally dentate (best visible in lateral aspect) (Figs. 5-7). Intraspecific variation and comparative notes: Apart from the coloration, intraspecific variation is not pro- nounced. From all its congeners, M. perniger is readily distinguished by the distinctive morphology of the aedeagus, which is most similar to that of M. maronitus (Saulcy) from the Eastern Mediterranean and M. bu- charicus Bernhauer from Middle Asia. For separation from the similar M. apicalis, whose distribution partly overlaps with that of M. perniger see diagnosis above. Distribution and bionomics: Medon perniger is evi- dently an Adriato-Mediterranean element, whose distri- bution is confined to Italy, including Sicily and Sar- dinia, to southeastern France (Provence) and southern Switzerland (Map 3). It is here recorded from France and Switzerland for the first time. An occurrence in Turkmenistan (see material examined) would be far out- side this range and the record should be considered doubtful until it is confirmed; quite possibly, the speci- mens were mislabelled. Apparently, the species is ab- sent from Corsica and Elba, where M. apicalis has been collected frequently; however, both species have been found in the same locality on various occasions (see the distribution section below M. apicalis). Map 3: Distribution of Medon perniger Coiffait in Italy and adjacent regions, based on revised records. 36 Bonner zoologische Beitráge 54 (2005) Unlike M. apicalis, most of the specimens listed above were not caught on the wing, but found in the leaf litter of various kinds of forests (oak, chestnut, etc.), under hay, or in nests of Talpa and Apodemus. OSELLA & ZANETTI (1974) frequently recorded the species (as M. apicalis) from mole nests, often in large numbers (up to 72 specimens per nest). Interestingly, they observed that M. perniger and M. ripicola appear to exclude each other in mole nests, with the former preferring drier situations and the latter occurring in damp habitats. The labels attached to the examined specimens indicate rela- tively low altitudes below 1000 m. With few exceptions, all the nest records examined are from the winter months. Flying specimens were collected in spring and summer (April through July, and September). On two occasions, the species was found at light sources (April, September). Three times in May and once in November it was collected from flood debris. Adult beetles were taken throughout the year, though only relatively few specimens were found in July and August. 3.3. Medon sericellus (Fairmaire, 1860) (Figs. 8-12, Map 2) Lithocharis sericella Fairmaire, 1860 (FAIRMAIRE 1860: 1591). Type: Neotype JS, here designated: Gde Kabylie: Ya- kouren, forét Beni-Ghobri, 800m, — V.1953, G. Fagel / G. Fagel det., 1954: Medon apicalis Kr / Neotypus @ Li- thocharis sericellus Fairmaire desig. V. Assing 2003 / Medon sericellus (Faimaire), det. V. Assing 2003 (IRSNB). Comments: Lithocharis sericellus was described by FAIRMAIRE (1860) based on material from Bone, Alge- ria. Shortly after the description, the name was placed in the synonymy of Medon apicalis, where it has remained ever since (see e.g. FAUVEL 1886; BERNHAUER & SCHUBERT 1912). The type material was looked for, but not found, in the collections of the MNHNP (person- ally) and of the IRSNB by the curator in charge (D. DRUGMAND, Bruxelles, pers. comm. 2003). Thus, it must be regarded as lost. Due to the similarity of the Algerian species redescribed in this section and of Me- don apicalis, as well as the fact the the sexual characters of Medon species were usually neither illustrated nor described in detail in the 19" century, the original de- scription can be attributed either to Medon apicalis or to the material listed in this section. I have not seen any specimens of M. apicalis from Algeria (or Tunisia), so that the presence of this species in Algeria is doubtful. Consequently, the material from Grande Kabylie is here treated as conspecific with the type material of M. sericellus and, in the interest of the stability of nomen- clature, a neotype 1s designated. Additional material examined (total: 22 exs.): Algeria: 13 exs., Grande Kabylie, Yakouren, forét Be- ni-Ghobri, 800 m, V.1953, leg. Fagel (IRSNB, cAss); 3 exs., Grande Kabylie, forét de Akfadou, Tala Kitan, 1100 m, 18.V.1953, leg. Fagel (IRSNB); 2 exs., Grande Kabylie, Foret d’Akfadou, 9 km W Adekar, 1300 m, 17.V.1988, leg. Besuchet, Löbl & Burckhardt (MHNG, cAss); 2 exs., Grande Kabylie, Djebel Bou-Berak, 350 m, 19.V.1988, leg. Besuchet, Lóbl & Burckhardt (MHNG); 1 ex., Atlas de Blida, Chrea, Les Glaciéres, 1100 m, 3.V.1988, leg. Besuchet, Löbl & Burckhardt (MHNG). Tunisia: | ex., 25 km W Jendouba, ca. 10 km N Ghar- dimaou, Ain Soltane, 36°29N, 08°19E, 670 m, litter of oak forest, 27.X11.2004, leg. Assing (cAss). Diagnosis: In external characters indistinguishable from M. apicalis (Figs. 8-9); separated from that species only by the male primary and secondary sexual characters: ¢: posterior margin of sternite VII in the middle con- cave (Fig. 10); aedeagus of highly distinctive morphol- ogy, apically very long and acute, both in lateral and in ventral view (Figs. 11-12). Comparative notes: From other species of the M. api- calis group, M. sericellus is reliably distinguished only by the shape and chaetotaxy of the male sternite VII and by the distinctive morphology of the aedeagus. Distribution and bionomics: Medon sericellus has be- come known only from central northern Algeria and from northwestern Tunisia (Map 2), where the species was collected at altitudes of 350-1300 m in May and December. 3.4. Medon oromii sp. n. (Figs. 13-16) Holotype $: Gran Canaria, 22-V-91, Bco. del Risco, P. Oromi / Holotypus Y Medon oromii sp. n. det. V. Assing 2003 (DZUL). Diagnosis: 3.9 mm. Coloration: head dark brown; pronotum reddish brown; elytra and antennae rufotesta- ceous; abdomen reddish brown with the apex lighter; legs testaceous. Head weakly oblong (Fig. 13); eyes slightly more than half the length of postocular region in dorsal view; puncturation relatively dense, but shallow, relatively fine, ill defined, and not very conspicuous in the pro- nounced microreticulation; dorsal surface matt. Volker ASSING: Western Palaearctic Medon 37 Figs. 1-16. Medon perniger Coiffait (1-7), M. sericellus (Fairmaire) (8-12), and M. oromii sp. n. (13-16): Habitus (1, 8); forebody (9, 13); male sternite VII (2); posterior margin of male sternite VII (3, 10, 14); male sternite VIII (4); aedeagus in lateral and in ventral view (5-7, 11, 12, 15, 16). Scale bars: 1, 8, 9, 13: 1.0 mm; 2-7, 10-12, 14-16: 0.2 mm. 38 Bonner zoologische Beitráge 54 (2005) Pronotum slightly narrower than head and approxi- mately as wide as long (Fig. 13); puncturation ex- tremely fine, barely noticeable; surface matt due to pro- nounced microreticulation. Elytra large and long, approximately 1.25 times as wide and at suture 1.15 times as long as pronotum (Fig. 13); puncturation dense and slightly granulose, much more distinct than that of pronotum; surface more shining than that of head and pronotum. Hind wings fully de- veloped. Abdomen with fine and dense puncturation; posterior margin of tergite VII with palisade fringe. ©: protarsomeres I - IV not dilated; posterior margin of sternite VII of similar morphology and chaetotaxy as that of M. apicalis (Fig. 14) Aedeagus of highly distinc- tive morphology, with long, slender, and apically dilated ventral process (Figs. 15-16). Etymology: The species is dedicated to Dr. Pedro Oromi, La Laguna, who collected the holotype and who has contributed so much to the current state of knowl- edge of Canarian Coleoptera. Comparative notes and systematics: Among its West- ern Palaearctic congeners, oromii is readily identi- fied especially by the extremely fine puncturation and pronounced microreticulation of the pronotum and by the distinctive morphology of the aedeagus. The mor- phology and chaetotaxy of the male sternite VII sug- gests that it belongs to the M. apicalis group. The aedeagus is most similar to that of M. sericellus. The only West Palaearctic congeners with a similarly pronounced microsculpture on the pronotum are M. feloi from La Palma, M. antricola from El Hierro, the Ca- narian endemic M. subcoriaceus, and the Madeiran en- demic M. vicentensis. All these species have strongly reduced eyes, except for M. subcoriaceus, which is dis- tinguished from M. oromii by an oblong head, relatively dark coloration, and completely different male sexual characters. Distribution and bionomics: The long wings and fully developed eyes would suggest that M. oromii is a wide- spread species, but currently it is known only from Gran Canaria, Canary Islands. Nothing is known about its natural history. Strangely, no further material of this species has become available, despite the fact that the staphylinid fauna of the Canaries is comparatively well studied. 3.5. Medon dilutus (Erichson, 1839) In the Western Mediterranean, Medon dilutus 1s repre- sented by three morphs, two of more or less uniformly rufous to ferrugineous coloration with shorter elytra and slightly smaller eyes and one usually with a coloration similar to that of M. castaneus (i.e. head, central part of pronotum, and abdominal segments blackish; elytra, pronotal margins, abdominal apex, and appendages lighter), with longer and broader elytra, and with slightly larger eyes. Based on slight differences in the shape of the aedeagal apex, the light-coloured short- winged morph can again be subdivided into fractions: one with an broader aedeagal apex with subparallel sides, distinct apico-lateral angles, and a small medio- apical emargination (Figs. 28-29), and one with curved sides, indistinct apico-lateral angles, and a large medio- apical emargination (Figs. 22-23), a condition also found in the long-winged morph. I have been unable to find other constant differences in the primary and sec- ondary sexual characters. Since these morphs show a zoogeographically plausible allopatric distribution pat- tern, their previously proposed subspecific status is here maintained. As no material of Lithocharis spelaea Scriba, 1870 or its synonym Medon dilutus breuili Jeannel, 1921 was available for examination, their status still requires clarification. The types of both names were collected in caves in Valencia (Spain). JEANNEL & JAR- RIGE (1949) provide a very detailed illustration of a male of M. dilutus breuili, demonstrating that the eyes are considerably smaller than in M. dilutus. This and the fact that the type locality is within the distribution range of M. dilutus cephalus suggests that M. spelaeus (Scriba) represents a distinct species rather than a sub- species. Remarkably, a comparison of the Western Mediterra- nean subspecies of M. dilutus with material of the wide- spread M. pythonissa (Saulcy, 1864) from the Eastern Mediterranean yielded no constant differences. In exter- nal appearance the latter is difficult to separate from M. d. quadriceps. It usually has an uniformly blackish pronotum, darker elytra, a somewhat coarser and sparser puncturation on the head and pronotum, slightly larger eyes, often an infuscated basal antennomere, but there are also transitions. The male secondary sexual charac- ters are identical to those of the western subspecies, and the aedeagal apex is of similar morphology as that of M. d. dilutus. These observations suggest that M. python- issa is conspecific with M. dilutus and that it should be regarded as a subspecies of this polytypical species, which apparently has a circum-Mediterranean distribu- tion: M. dilutus dilutus, M. d. cephalus, and M. d. quad- riceps in the northwest and southwest, respectively, of the Mediterranean basin and M. d. pythonissa represent- ing the species in the eastern parts of the Mediterranean (Map 5). Volker ASSING: Western Palaearctic Medon 39 3.5.1. Medon dilutus dilutus (Erichson, 1839) (Figs. 24-29, Maps 4-5) Lithocharis diluta Erichson, 1839 (ERICHSON 1839: 514). Lithocharis oppidana Kraatz, 1857 (KRAATZ 1857: 71 1f) synonymy confirmed. Medon dilutus meridionalis Bordoni, 1980 (BORDONI 1980: 98) syn. n. Medon dilutus: GUSAROV 1992: 22f; invalid lectotype designation. Types examined: L. diluta: Holotype Y: 6364 / diluta Er. Berol. Er. / Lectotypus Y Lithocharis diluta Erich- son V. Gusarov des. 1991 / Medon dilutus (Er.) Y Gusarov det. 1991 / Holotypus Lithocharis diluta Erich- son labelled by Zool. Mus. Berlin, 7.2003 / Hist.-Coll. (Coleoptera) Nr. 6364 Lithocharis diluta Er., Berlin, Erichson, Zool. Mus. Berlin (MNHUB). L. oppidana: Lectotype © [remounted, aedeagus dis- sected], present designation: Berol. / Mayer / oppidana mihi / Syntypus / Coll. Kraatz / & / Coll. DEI Eberswalde / Lectotypus @ Lithocharis oppidana Kraatz desig. V. Assing 2003 / Medon dilutus (Erich- son) det. V. Assing 2003 (DEI). Paralectotype Y: Ber- lin, Calix / oppidana Kr. * / Syntypus / Coll. Weise / Coll. DEI Eberswalde / Paralectotypus Y Lithocharis oppidana Kraatz desig. V. Assing 2003 / Medon dilutus (Erichson) det. V. Assing 2003 (DEI). M. dilutus meridionalis: Holotype ©: prov. di Bari, grot- ta di Cassano, 14.X1.1908, Dr. A. Andreini / HOLO- TYPUS / Medon dilutus meridionali ssp. n. det. Bordoni 1980 / Medon dilutus meridionalis Bord. Gusarov det. 1991 / Medon dilutus (Erichson) det. V. Assing 2003 (cBor). Paratypes: 14 [teneral], 329: same data as holotype (cBor). Comments: The original description of Lithocharis di- luta is based on “ein einziges weibliches Exemplar” from the Erichson collection, which consequently has holotype status. Thus, the lectotype designation by GUSAROV (1992) is invalid. The holotype represents the light-coloured morph (similar to material from Sardinia and Spain). According to KRAATZ (1857), there are several syntypes of Lithocharis oppidana, which were collected “unter feuchtem Laube von den Herren Mayer und Calix”. Two of these syntypes were found in the Kraatz collec- tion; they are conspecific with the holotype of M. dilu- tus (Erichson). In order to secure the present interpreta- tion and the long-standing synonymy with Medon dilutus, the male syntype is here designated as the lecto- type. According to BORDONI (1980), M. dilutus meridionalis is characterized by large body size, light coloration, elongate appendages, less convex eyes, and the different shape of the apex of the aedeagus. Regarding both ex- ternal characters and the shape of the aedeagus the types are indistinguishable from Central European M. dilufus dilutus (see Figs. 25-26, 28-29), so that the hypothesis that the population in southern Italy represents a distinct subspecies is not supported and M. d. meridionalis is considered a junior synonym of M. d. dilutus. Additional material examined (total: 25 exs.): Germany: Berlin/Brandenburg: 1 ex., Brieselang (DEI); 4 exs., Eberswalde [“Neustadt”] (DEI, NHMW); l ex., Berlin env. (DEI); 3 exs. [2 with worker of Lasius fuliginosus attached to the pin], Forst Dubrow, coll. Ne- resheimer (DEI). Sachsen: | ex., Rauden (DEI). Austria: Niederösterreich/Wien: 6 exs., Wien, Lobau, mole nest, 21.111.1926, leg. Beier (NHMW, cAss). Bur- genland: 2 exs., Zurndorf, leg. Franz (NHMW) Czech Republic: 2 exs., Praha (NHMW); 1 ex., Bran- dys n. L. (NHMW). Hungaria: | ex., NW Budapest, Mt. Pilis (NHMW). Locality not identified or not specified: 1 ex., without locality label, coll. Erichson (MNHUB); 2 exs. (DEL, NHMW). Diagnosis: Since the nominal subspecies and M. d. cephalus are externally indistinguishable, the following description of characters, except for that of the aedeagus, refers to both taxa: 4.0-6.0 mm. Coloration more or less uniformly rufous to ferrugineous. Head of very variable size and shape, weakly transverse to oblong (Figs. 17-21, 25-26), in small specimens so- metimes not wider than pronotum (Fig. 18), in large specimens 1.2 times as wide as pronotum (Figs. 19, 26); eyes barely projecting from lateral outline of head, in dorsal view 0.35 - 0.45 times the length of postocular region; puncturation relatively fine, dense, well-defined, non-areolate, and usually more or less evenly distributed on whole dorsal surface (1. e. as dense in central dorsal area as in lateral areas), but density of puncturation rather variable (Figs. 17-21, 25-26); microsculpture ab- sent or very shallow. Pronotum usually with puncturation similar to that of head, with or without narrow unpunctured midline. Elytra 1.10 - 1.15 times as wide as pronotum, at suture approximately as long as, sometimes slightly longer than pronotum (Figs. 17-21, 25-26). Abdomen with very dense and fine puncturation; posterior margin of tergite VII with palisade fringe. 40 Bonner zoologische Beitráge 54 (2005) Map 4: Distributions of Medon dilutus dilutus (Erichson) (open circles; examined records), M. d. cephalus Koch (filled circles; examined records), and of M. d. quadriceps (Wollaston) (filled squares; examined records and literature records). Grey circles: li- terature records, referring to either M. d. dilutus and M. d. cephalus. ¢: protarsomeres I - IV distinctly dilated; sternite VII lateral angles, and a small medio-apical emargination posteriorly moderately excavate, with two combs of (ventral view) (Figs. 27-29). palisade setae, margin between these combs straight and without conspicuous tufts of long dark setae (Fig. 24); sternite VIII with relatively small posterior excision; Intraspecific variation: Both the nominal subspecies aedeagal apex with subparallel sides, pronounced apico- and M. d. cephalus are extremely variable. This particu- Y: protarsomere I - IV not dilated. Volker ASSING: Western Palaearctic Medon 4] larly applies to the coloration, body size, the size and shape of the head, and the density of the puncturation (Figs. 17-21, 25-26). The head is usually relatively lar- ger (e. g. in relation to the pronotum) and more trans- verse in larger than in smaller specimens (compare Figs. 18 and 19). Comparative notes: Among Western Mediterranean Medon species, M. dilutus dilutus is readily identified by the dense, well defined, relatively fine, and usually more or less evenly spaced puncturation of head and pronotum, by the uniformly light coloration, and by the primary and secondary sexual characters. The somewhat similar M. pocofer has a flatter head, a more irregularly spaced puncturation of the head and pronotum, a rela- tive wide impunctate midline of the pronotum, and shorter and broader pro-, meso-, and metatarsomeres. From M. d. quadriceps and M. d. pythonissa it is distin- guished especially by the uniformly light coloration, the smaller eyes, and the shorter and narrower elytra, from M. d. quadriceps also by the different shape of the aedeagal apex. Distribution and bionomics: Medon dilutus dilutus 1s apparently an Adriato-Mediterranean element (Maps 4- 5), its distribution ranging from southern mainland Italy in the south at least to southern Central Europe (Poland, Austria, Germany, Hungary; material examined). The literature records from the western parts of Germany and from the Swedish island Gotska Sandón (LUND- BERG 1978) may refer to either this subspecies or to M. d. cephalus; the corresponding material was not avail- able for examination. For additional literature records see also BOHAC (1985), HORION (1965), JANAK (1993), KOCH (1968), KOHLER & KLAUSNITZER (1998), KOLBE (1918), KORGE (1989), LUIGIONI (1929), and LUNDBERG (1995). The species is unknown from Belgium, Den- mark, and the Baltic countries (BRUGE 2001; DRUG- MAND 1989; HANSEN 1996; SILFVERBERG 1992). I know of only one recent record from Germany (KORGE 1989); from the Rhineland area, only two old records are known (KOCH 1968 1974), but they may refer to M. d. cephalus. In Italy, it is known from three localities in Campania and Puglia in the south of the mainland (BORDONI 1980; LUIGIONI 1929; SCHEERPELTZ 1958). It has also been recorded from Corsica (LUIGIONI 1929; SAINTE-CLAIRE DEVILLE 1906), but this record, too, probably refers to M. d. cephalus. The record of M. dilu- tus from the Crimea by GUSAROV (1989) was later cor- rected into M. mersinus Bordoni by GUSAROV (1992); M. mersinus is now a synonym of M. d. pythonissa (see ASSING 2004a). On the whole, M. dilutus dilutus has been collected very rarely. On several occasions, the subspecies was found at the bases or under bark of old or dead oak trees, un- der bark of pine trees, in nests of various mammals (mole, mouse/vole, fox), in a nest of the ant Lasius fu- liginosus, and in caves (material examined, and BOHAC (1985); BORDONI (1980); FRANZ (1938); HORION (1965); KORGE (1989); STROUHAL & BEIER (1928)). On one occasion, STROUHAL & BEIER (1928) found as many as 10 specimens in one apparently abandoned mole nest in Wien-Lobau, but no further specimens were recorded from a total of 188 nests examined. All these data suggest that the reproduction habitat is somewhat cryptic, subterranean, but essentially un- known. Flying specimens were recorded in May and June (HORION 1965). Adult beetles have been found al- most throughout the year, but most records are from spring and autumn. 3.5.2. Medon dilutus cephalus Koch, 1938 (Figs. 17-23, Maps 4-5) Medon dilutus cephalus Koch, 1938 (KOCH 1938: 140f) Medon dilutus boeticus Jeannel & Jarrige, 1949 (JEAN- NEL & JARRIGE 1949: 361) syn. n. Types examined: M. dilutus cephalus: Holotype ©: LULA, Sard., 7.111.1912, A. Dodero / Medon dilutus / spec. nov. det. C. Koch / Type / Mikr. Pr. 104 / M. dilu- tus var. cephalus Koch det. C. Koch / Medon dilutus (E- richson) det. V. Assing 2003 (NHMB). M. dilutus boeticus: Syntype Y: Cueva de la D. Trinidad / Biospeologica 932 III 18 / Jeanneli m. det. Dr. Ram- bousek / Muséum Paris Ex Collection J. JARRIGE1976 / COTYPE / d. baeticus [sic] Jean. & Jar. / Medon dilu- tus (Erichson) det. V. Assing 2003 (MNHNP). Comments: The holotype of M. dilutus cephalus was dissected prior to the present study. The aedeagus and apical abdominal segments are missing; as can be in- ferred from one of the labels attached to the specimen, they were mounted separately for microscopic examina- tion probably by Koch himself. The type of M. dilutus boeticus represents a — probably submacropterous — uniformly ferrugineous specimen with relatively short elytra. In all respects it is within the range of intrasubspecific variation of M. dilutus cepha- lus, hence the synonymy indicated above. Additional material examined (total: 48 exs.): Spain: Cataluña: | ex., Malgrat de Mar, 3.V.1959, leg. Liebmann (NHMW). Estremadura: | ex., Cáceres, Gata, 3.V.1957, leg.González (MHNG). Andalucia: 5 exs., Sierra de Alhamilla (AL), 1000 m, N-slope, 20.11.1994, leg. Assing, Wunderle (cAss, cWun); 2 exs., Sierra Ne- vada, Rio Genil, 37%0826N, 03°24’01W, 1000 m, 24.11.2000, leg. Lompe (cAss); 1 ex., Sierra de Cordoba, leg. Franz (NHMW); 1 ex., San Roque (CA), leg. Co- mellini (MHNG). 42 Bonner zoologische Beitrage 54 (2005) France: 2 exs., Lyon env. (DEI); | ex., Fontainebleau, 23.V.1900 (MHNG); 1 ex., La Bonde, 10.XII.1903, leg. Fagniez (MHNG); 2 exs., Gironde, Biscarosse Plage, 25.-28.V11.1980, leg. Kiener (MHNG). Corsica: | ex., locality not specified (NHMW). England: | ex., London, Richmond, 19.X.1998, leg. Owen (cOwe). Italy: Sardegna: | ex., Mte. Chiesa (NHMB); 2 exs., Aritzo (NHMW); 1 ex., Aritzo, Mte. Genargentu, leg. Krüger (DEI); 2 exs., Golfo Aranci, 111.1907, leg. Dode- ro (DEI); 1 ex., Seneghe, 2.V.1891, leg. Dodero (DEI); 1 ex., Seu (NHMB); | ex., Seui, 7.V.1902, leg. Dodero (NHMW ); 2 exs., Seui, leg. Dodero (NHMW, cAss); 1 ex., N Bolotana (SS), 850 m, 13.1V.1992, leg. Scheuern (cAss); 1 ex., Onani (NU), 450m, 24.X1.1992, leg. Me- loni (cZan); 1 ex., Lanusei (NU), 12.V.1980, leg. Tor- chia (UCBA); 1 ex., Padru (SS), XII.1999, leg. Fancello (cZan); | ex., Iglesias (CA), 8.111.2001, leg. Fancello (cZan); 1 ex., Sorgono, leg. Krausse (DEI); 1 ex., Cam- peda, 28.1V.1908, leg. Dodero (cAss); 1 ex., NE Sadali (NU), 850 m, 20.1V.1992, leg. Schawaller (SMNS). Diagnosis: For a description of external and secondary sexual characters see the diagnosis of M. d. dilutus above. ÁS: aedeagal apex in ventral view with curved sides, in- distinct apico-lateral angles, and a large medio-apical incision (Figs. 22-23). Intraspecific variation: Similar to that of M. d. dilutus. Comparative notes: For separation from other Western Mediterranean species of Medon, see the comparative notes below M. d. dilutus. From M. d. quadriceps and M. d. pythonissa it is distinguished by the uniformly light coloration, the smaller eyes, and the shorter and narrower elytra, from M. d. pythonissa also by the dif- ferent shape of the aedeagal apex. Map 5: Distributions of Medon dilutus dilutus (Erichson) + M. d. cephalus Koch (filled circles), M. d. quadriceps (Wollaston) (filled squares), and M. d. pythonissa (Saulcy) (open circles), mainly based on examined records. Distribution and bionomics: Medon dilutus cephalus is an Atlanto-Mediterranean element (Maps 4-5), its dis- tribution ranging from southern Spain and Sardinia in the southwest and south to southern England in the north, possibly even to the Swedish island Gotska Sandón (see notes on the distribution of the nominal Volker ASSING: Western Palaearctic Medon 43 subspecies). In Sardinia it is apparently relatively com- mon (see material examined). The records of M. dilutus from Corsica (LUIGIONI 1929; SAINTE-CLAIRE DEVILLE 1906) probably refer to this subspecies, but I have seen only a female from this island. In the British Isles, it is known from only three localities in southern England (ALLEN 1996; OWEN 1998, 2000). This subspecies has been found in similar habitats and under similar circum- stances as M. d. dilutus. Several specimens were sifted from the leaf litter in a mixed stand of Quercus ilex and Pinus sp. in southern Spain (material examined). 3.5.3. Medon dilutus quadriceps (Wollaston, 1864) (Figs. 30-32, Maps 4-5) Lithocharis quadriceps Wollaston, 1864 (WOLLASTON 1864: 586). Medon vitalei Bernhauer, 1936 (BERNHAUER 1936: 306f) syn. n. Medon dilutus eremicus Koch, 1939 (KOCH 1939: 243f) syn. n. Medon algiricus Jeannel & Jarrige, 1949 (JEANNEL & JARRIGE 1949: 361f). Medon marocanum Coiffait, 1970a (COIFFAIT 1970a: 707) syn. n. Medon mateui Coiffait, 1973b (COIFFAIT 1973b: 282) syn. n. Types examined: M. vitalei: Holotype Y: Castanea, 27.X.34, F. Vitale / Vitale: Brnh. Typus unic. Medon / Medon dilutus (Erichson) det. V. Assing 2003 (FMNH). M. dilutus eremicus: Holotype Y: Tauorga nordöstl. Trip., IV.38. G. Frey / Medon eremicus Koch det. C. Koch / Medon dilutus (Erichson) det. V. Assing 2003 (NHMB). M. marocanus: Holotype 4: IXMOART, B. SICAR. (MARR.), COBOS SANCHEZ / MUSEUM PARIS COLL H. COIFFAIT / HOLOTYPE / Medon maroca- num Coiff. H. COIFFAIT det. 1969 / Medon dilutus (Erichson) det. V. Assing 2003 (MNHNP). M. mateui: Holotype 4: MAROC, env. Ifni, H. COIF- FAIT / MUSEUM PARIS COLL H. COIFFAIT / HO- LOTYPE / Medon mateui H. COIFFAIT det. 1973 / Medon dilutus (Erichson) det. V. Assing 2003 (MNHNP). Comments: The original description of Lithocharis quadriceps, which is based on material from Lanzarote and Fuerteventura, leaves no doubt that the types are conspecific with the male non-type specimen seen from Fuerteventura. The original description of M. vitalei is explicitly based on a single female. The holotype was examined and found to be a small specimen of the morph here referred to as M. dilutus quadriceps, hence the synonymy indi- cated above. Similarly, no evidence was found suggest- ing that the examined holotypes of M. dilutus eremidus and M. marocanus should represent a distinct (sub-) species, so that both names are placed in the synonymy of M. d. quadriceps. COIFFAIT (1973b) compares M. mateui with M. maro- canus stating that the former is distinguished from the latter by the dilated male protarsi, the more shallowly convex posterior margin of the male sternite VII, and the apex of the aedeagus having the shape of a semicir- cle and lacking the median incision. An examination of the holotype of M. mateui, however, revealed no such differences in the degree of dilatation of the male pro- tarsi or in the shape of the male sternite VII. Moreover, the apex of the aedeagus clearly has a median incision, though this incision is somewhat smaller than is usually the case in M. dilutus. However, this teratological mal- formation is a common phenomenon in the genus and is present also, for instance, in the holotype of M. aqui- tanicus Coiffait. Aside from the somewhat lower body size and the much smaller head, the holotype of M. ma- teui is, in fact, strikingly similar to the holotype of M. marocanus. Since there is no evidence that it represents a distinct species, M. mateui is here synonymized with the senior name M. dilutus quadriceps. Additional material examined (total: 23 exs.): Tunisia: | ex., Kasserine (=Al Qasrayn), II.1941, leg. Demoflys (cTro); 3 exs., Zarzis (=Jarjis), 11.1951, leg. Demoflys (cTro, cAss); 1 ex., Zarzis, VI.1944, leg. De- moflys (cTro); 1 ex., Gafsa (MHNG); 1 ex., Qibili [“Kebili”, 33°41N, 08°58E] (MHNG); 1 ex., Talah [“Thala”, 35°34N, 08°40E] (cAss); 1 ex., ca. 40 km W Sousse, 35°54N, 10%15E, roadside, under stones, 30.X11.2004, leg. Assing (cAss). Morocco: | ex., Oued Sebou, IV.1961, leg. Comellini (cAss); 1 ex., Anti-Atlas, Agard-Oudad (?), 1200m, 7.- 8.1X.1990, leg. Annot (DEI). West Sahara: | ex., Meseied near Al Aaiun, leg. Franz (NHMW). Portugal: Azores: 4 exs. [identification uncertain], Sao Miguel, leg. Franz (NHMW, cAss). Canary Islands: | ex., Fuerteventura, Cueva del Llano, 4.1V.1992, leg. Oromi (cAss). Italy: Sicilia: 1 ex., locality not specified, leg. Vitale (UCBA); 1 ex., Messina, leg. Vitale (cAss). 44 Bonner zoologische Beitráge 54 (2005) Figs. 17-32. Medon dilutus dilutus (Erichson) (24-29), M. dilutus cephalus Koch (17-23), and M. dilutus quadriceps (Wollaston) (30-32): Habitus (17: Sardinia; 18: Andalucia: 30: Tunisia); forebody (19: Sardinia, 20: holotype of M. dilutus cephalus Koch; 21: Andalucia: 25: Wien; 26: holotype of M. dilutus meridionalis Bordoni; 31: Tunisia: 32: Fuerteventura); male sternite VII (24: Prague); aedeagus in lateral and in ventral view (27, 28: Wien; 29: Prague; 22: Andalucia; 23: Sardinia; ). Scale bars: 17-21, 25- 26, 30-32: 1.0 mm; 24: 0.5 mm; 22-23, 27-29: 0.2 mm. Volker ASSING: Western Palaearctic Medon 45 Diagnosis: This subspecies is distinguished from the nominal subspecies and from M. d. cephalus as follows: Coloration highly variable, but usually similar to that of M. castaneus and M. subcoriaceus: Head, central part of pronotum, and abdominal segments III to anterior half of VII usually dark brown to blackish brown; margins of pronotum near anterior angles broadly, in other places narrowly ferrugineous; elytra and abdominal apex testaceous, rufous, or brown; legs and antennae more or less rufous. Elytra larger, approximately 1.3 times as wide and at su- ture 1.1 times as long as pronotum. Eyes of somewhat variable size, but on average larger, approximately half as long as temples in dorsal view (Figs. 30-32). 3: aedeagal apex as in M. d. cephalus (cf. Figs. 22-23). Intraspecific variation: The extent of intrasubspecific variation is similar to that observed in the other subspe- cies, especially as regards coloration, body size, as well as size and shape of the head. One nanistic specimen seen from Morocco represented a transitional state be- tween the nominal subspecies and M. d. quadriceps in that it had a light coloration and relatively long, but rather narrow elytra. The specimens from the Azores have a very pronounced microsculpture on the head and pronotum; the latter is of uniformly dark coloration. Un- fortunately, only females were available from the Azores, so that their present interpretation must be con- sidered tentative. Comparative notes: Medon d. quadriceps is highly similar to the Ponto-Mediterranean M. d. pythonissa, but distinguished by on average finer puncturation of the head and pronotum, the lower average size of the eyes, the lighter margins of the pronotum, the darker elytra, and the different shape of the aedeagus apex (in M. d. quadriceps similar to that of M. d. cephalus and in M. d. pythonissa similar to that of the nominal subspecies). Distribution and bionomics: The distribution of M. d. quadriceps (Maps 4-5) includes Northwest Africa from Tunisia to southern Morocco and West Sahara, the Ca- nary Islands (Fuerteventura, Lanzarote), and Sicily, pos- sibly also the Azores (material examined; see also HORION (1965), MACHADO & OROMÍ (2000), and NOR- MAND (1935)). Horion (1965) reports it from the Hog- gar range in southern Algeria. Material of this subspe- cies has been collected in various habitats such as caves or a sparrow’s nest (NORMAN 1935), but its real repro- duction habitat must be considered unknown. The dis- tribution suggests that it is adapted especially to hot and dry regions. The material examined was collected in February, April, June, October, and December. 3.6. Medon subcoriaceus (Wollaston, 1864) (Figs. 33-37) Lithocharis subcoriaceus Wollaston, 1864 (WOLLAS- TON 1864: 586). Material examined (total: 1097 exs.; see also material listed in ASSING (1999a, 2000) and ASSING & WUNDERLE (1999)): Canary Islands: Tenerife: 4 exs., Pico del Ingles, 960 m, 4.&13.1V.1992, leg. Assing (cAss); | ex., same lo- cality, 950 m, 28.X1.1996, leg. Schülke & Grünberg (cSch); 2 exs., La Laguna, 750 m, 4.1V.1992, leg. Assing (cAss); 16 exs., El Bailadero, 900 m, 3.1V.1992, leg. Assing (cAss); 22 exs., Anaga, Chamorga, 700 m, 5.1V.1992, leg. Assing (cAss); 1 ex., Anaga, Chinobre, 850 m, 5.1V.1992, leg. Assing (cAss); 1 ex., Anaga, E Chinobre, Bco. d. Corral Viejo, 600 m, 5.X11.1996, leg. Schülke (cSch); 2 exs., E Chinobre, 700 - 800 m, 24.X1.1996, leg. Schülke & Grünberg (cSch); 2 exs., NE Chinobre,. 850 m, 8.V11.1995, leg. Zerche. (DEN); 1 ex., Anaga, Taborno, 1020 m, 5.1V.1992, leg. Assing (cAss); 14 exs., Anaga, leg. Franz (NHMW); 4 exs., Montes de las Mercedes, leg. Franz (NHMW); 1 ex., Las Mercedes, 750 m, 4.1V.1992, leg. Zerche (DEI); 1 ex., Anaga, Estanque, 28°34N, 16°10W, 26.111.2000, leg. Lompe (cAss); 1 ex., Anaga, 28°34N, 16°10W, 25.111.2000, leg. Lompe (cAss); 2 exs., Anaga, Atalaya del Sabinal, 1.5 km SE Lomo de la Bodegas, 500 - 550 m, 11.1.2002, leg. Schülke (cSch); 1 ex., Anaga, 1 km S Lomo de las Bodegas, road to Chamorga, 650 m, 11.1.2002, leg. Schülke (cSch); 16 exs., Anaga, 3 km E El Bailodero, 900 m, 3.1V.1992, leg. Zerche (DEI); 17 exs., El Bailodero, 800 m, 2.VII.1995, leg. Zerche (DEI); 1 ex., Anaga, Cabezo del Tejo, 730 m, 5.V11.1995, leg. Zerche (DEI); 13 exs., Anaga El Pija- ral, 800-850 m, laurisilva, 4.V11.1995, leg. Zerche (DEI); 7 exs., Anaga, Vueltas de Taganana, 780-850 m, Erica forest, 5.V11.1995, leg. Zerche (DEI); 2 exs., same data, but 450 m, laurisilva (DEI); 9 exs., Anaga, Cruz del Carmen, 900 m, Erica litter, 2.V11.1995, leg. Zerche (DEI); 5 exs., Bosque de la Esperanza, W Las Rosas, 1000 m, 26.11.1996, leg. Schülke & Grünberg (cSch):; l ex., Esperanza, 1200 m, 6.VIL1995, leg. Zerche (DEI); 1 ex., Orotava valley, Aguamansa, 1080 m, 25.X11.1995, leg. Stúben & Bahr (DEI); 9 exs., Teno, Erjos, 900 m, 8.-10.1V.1992, leg. Assing (cAss); 3 exs., Erjos, 12.111.1996, leg. Meybohm (cAss); 1 ex., Teno, N Erjos, 700 m, 30.X1.1996, leg. Schülke & Grünberg (cSch); 2 exs., Teno, Monte del Agua, 900-950m, 1:£:10.V11. 1995, leg. -Zerche (DEI); 1. ex., Bajamar, 27.1X.1965, leg. Benick (cAss); 2 exs., Bco. del Infier- no, 500 m, 9.1V.1992, leg. Assing (cAss); 3 exs., Vuel- tas de Taganana, VI.1957, leg. Fernández (NHMW); 4 exs., locality not specified (NHMW). Gran Canaria: 98 exs., NNW Lanzarote, Bco. de la Virgen, 500 m, de- 46 Bonner zoologische Beitráge 54 (2005) graded laurisilva, 20.X11.1997, leg. Assing (cAss); 7 exs., E Osorio, SW El Palmar, 600m, degraded laurisil- va, 20.X11.1997, leg. Assing (cAss); 10 exs., Pinar de Tamadaba, 1200 m, pine forest, 22.X11.1997, leg. Assing (cAss); 3 exs., Pinar de Tamadaba, 1275m, 28°03N, 15°41W, pine forest with Erica arborea, 4.11.1998, leg. Zerche (DEI); 1 ex., S Moya, Los Tilos, 400 m, laurisilva, 23.X11.1997, leg. Assing (cAss); 9 exs., W Moya, El Palmital, 500 m, 23.XII.1997, leg. Assing (cAss); 2 exs., Cruz de Tejeda, 1600 m, 25.X0.1997, leg. Assing (cAss); 1 ex., Tejeda, leg. Gaudin (cSch); 10 exs., Valleseco, leg. Franz (NHMW); 9 exs., laurisilva near Moya, leg. Franz (NHMW); | ex., Isleta near Las Palmas, leg. Franz (NHMW); | ex., Bco. de Mogan, 27°54N, 15°42W, 355 m, Euphorbia obtusi- folia + Echium decaisnei, sifted, 7.11.1998, leg. Zerche (DEI). La Palma: | ex., Los Tilos, leg. Franz (NHMW). El Hierro: | ex., Los Gramales, 21.1V.1985, leg. Oromí (cAss) 9 exs., Valverde, leg. Franz (NHMW ); 7 exs., El Golfo, leg. Franz (NHMW); 4 exs., El Golfo, S El Moncenal, 900 m, Fayal-Brezal, 9.X11.1996, leg. Schülke (cSch); 1 ex., Mirador de la Peña, 650 m, 10.-13.X11.1996, leg. Schiilke & Grünberg (cSch); 2 exs., Sabina, leg. Franz (NHMW); 12 exs. [partly teneral], El Brezal near Tabano, 850/1205 m, fayal-brezal, 27°44N, 18°01W, 17.1.1998, leg. Behne (DEI); | ex., below Sabinosa, 95 m, 27°45N, 18°06W, Eonium sifted, 27.1.1998, leg. Behne (DEI); 31 exs., Lomo Blanco, 6 km W Frontera, 27°45N, 18°03W, 550 m, fayal-brezal, 25.1.1998, leg. Behne (DEI); 5 exs., El Gretime, 8 km W Frontera, 27°44N, 188°04W, 800 m, fayal-brezal, 21.1.1998, leg. Behne (DEI); 4 exs. [1 tene- ral], Mtn. Colorada, 10 km SW Frontera, 27°44N, 18°02W, 1050 m, 21.1.1998, leg. Behne (DEI); 1 ex., Camino de San Salvador near Tabano, 27°44N, 18201 W, 1325 m, fayal-brezal, 17.1.1998, leg. Behne (DEI). La Gomera: | ex., El Cedro, 1000 m, 27.X.1990, leg. Wunderle (cAss); 2 exs., El Cedro, 900 m, 2.X1.1990, leg. Wunderle (cAss, cSch); 35 exs., El Cedro, leg. Franz (NHMW); 2 exs., El Cedro, 29.V1.2002, leg. Klemm & Wisser (cSch); 4 exs., El Cedro, | km S Ermita, 14.1.2002, leg. Schülke (cSch); 20 exs., El Cedro, Ermita, 900 m, 17.&19.VII.1995, leg. Zerche (DEI); 4 exs. [1 teneral], El Cedro, 600 m, 2.1.1998, leg. Stúben & Bahr (DEI); 1 ex., S Las Creces, 21.11.-8.01.2001, leg. Arndt (cSch); 1 ex., N Laguna Grande, laurisilva, 14.1.2002, leg. Schúlke (cSch); 2 exs., Camino Forestal La Meseta, 650 m, laurisilva, 15.V11.1995, leg. Zerche (DEI); 2 ex., barranco above Hermigua, laurisilva, 550 - 580 m, 16.&19.V11.1995, leg. Zerche (DEI); 6 exs., Bco. N La Laguna Grande, 1010-1050 m, laurisilva, 15.&17.VII.1995, leg. Zerche (DEI); 2 exs.; Raso de la Bruma, 1000 m, laurisilva, 18.V11.1995, leg. Zerche (DEI); 1 ex., Mirador de Alo- jera, 26.V.2003, leg. Contreras & Oromi (cOro); 2 exs., Cabecera del Bco. de Juel, 8.V111.2002, leg. Oromí (cOro). Fuerteventura: | ex., Mte. de Betancuria, leg. Franz (NHMW). Diagnosis: 4.0-5.2 mm. Habitus as in Fig. 33. Head brown to blackish; pronotum usually slightly lighter than head, more rarely of similar coloration; elytra light brown to brown; abdomen, except for the lighter apex, of similar coloration as head or pronotum; legs and an- tennae rufous to yellowish brown. Head distinctly oblong, about 1.10 - 1.15 times as long as wide; eyes relatively large, in dorsal view usually slightly more than half the length of temples; punctura- tion very fine, dense, and more or less well-defined; mi- crosculpture usually distinct, more rarely shallow, but always visible especially in lateral areas (Fig. 34). Pronotum 1.05 - 1.10 times as wide as head and ap- proximately as wide as long; puncturation even finer than that of head, often barely noticeable in the pro- nounced microsculpture (Fig. 34). Elytra relatively large, 1.2 - 1.3 as wide and at suture 1.10 - 1.15 times as long as pronotum (Fig. 34); punctu- ration dense, fine, and weakly granulose. Hind wings fully developed. Abdomen narrower than elytra (Fig. 33); puncturation very dense and fine; posterior margin of tergite VII with palisade fringe. ¢: posterior margin of sternite VII deeply concave, with two combs of 5 - 8 palisade setae and on either side of middle with a tuft of long dark setae (Fig. 35); sternite VIII not distinctive; aedeagus of similar general mor- phology as that of M. dilutus, but apical part of different shape (Figs. 36 - 37). Comparative notes: Medon subcoriaceus is readily identified by external characters alone, especially by its oblong head with a dense, fine, and well-defined punc- turation and usually distinct microsculpture, by the ex- tremely finely punctured and distinctly microsculptured pronotum, and the long elytra. In addition it is identified by the presence of tufts of long black setae at the poste- rior margin of the male sternite VIII (one on either side of middle) — a character otherwise exclusive to the fol- lowing species, to M. kabylicus, and to the species of the M. fusculus group — as well as by the distinctive shape of the apex of the aedeagus. Volker ASSING: Western Palaearctic Medon 47 a gy! A Y Aj | A y ¿AN AOS Y NT e Figs. 33-45. Medon subcoriaceus (Wollaston) (33-37) and M. antricola sp. n. (38-45): Habitus (33, 38); forebody (34); male ster- nite VII (35, 42); aedeagus in lateral and in ventral view (36, 37, 44, 45); head and pronotum (39); male protarsus (40); metatar- sus (41); male sternite VII (43). Scale bars: 33, 34, 38, 39: 1.0 mm; 35, 40-43: 0.5 mm; 36, 37, 44, 45: 0.2 mm. 48 Bonner zoologische Beiträge 54 (2005) Distribution and bionomics: Medon subcoriaceus is endemic to the Canary Islands. It occurs in all larger is- lands except Lanzarote (material examined; MACHADO & OROMI 2000) and is common everywhere except Fuerteventura, from where it is here recorded for the first time. The species was found in large numbers espe- cially in the laurisilva, fayal-brezal, and other semi- natural woodland habitats of Tenerife, Gran Canaria, La Gomera, La Palma, and El Hierro, but also in semi-arid biotopes. The examined material was collected through- out the year; teneral adults were repeatedly observed in January. 3.7. Medon antricola sp. n. (Figs. 38-45) Holotype Ö: El Hierro, JN-C/CO-3920, Cueva de Jina- ma, 26-IX-00, GIET leg. / Holotypus 4 Medon antrico- la sp. n. det. V. Assing 2003 (DZUL). Paratypes: 14: El Hierro, JN-C/CO-3779, Cueva de Jinama, 26-09-00, GIET leg.; 19: El Hierro, JN-C/CO-2523, Cueva de Ji- nama, 9-11-2000, GIET leg.; 17: El Hierro, Fi-VG/CO- 2346, Cueva de Fileba, 8.11.2000, GIET leg. (DZUL, MCNT, cOro); 14: El Hierro, Fi-C/CO-2103, Cueva de Fileba, 1.11.2000, GIET leg. (cAss); 14 [teneral]: El Hierro, Cueva de Fileba, 4-XI-01, H.López leg. (cAss); 14 [remains; in ethanol]: El Hierro, Cueva de Jinama, 26.1X.00, JN-C/CO-3779, GIET leg. (DZUL); 1% [re- maines; in ethanol]: NO: FI-C/CO-3868, El Hierro, Cueva de Fileba, Fecha: 15.1X.00, GIET leg. (DZUL ); 19: No: FI-C/CO-5717, El Hierro, Cueva de Fileba, Fe- cha: 28/07/01, leg. S. de la Cruz (cAss). Diagnosis: 5.0-6.8 mm. Habitus as in Fig. 38. Colora- tion of whole body more or less uniformly reddish brown, with the elytra, the legs, and the antennae yel- lowish brown. Head distinctly oblong, 1.10 - 1.15 times as long as wide; widest at a short distance behind eyes, 1.e. at slightly tapering posteriad (Fig. 39); eyes reduced: ru- diments without pigmentation and without distinct om- matidia, in dorsal view about 1/5 - 1/6 the length of temples; integument with very dense, fine, shallow, non-areolate puncturation; interstices with distinct mi- crosculpture. Antennae long and slender; antennomere III approximately 4 times as long as wide and distinctly longer than II; IV shorter than III and more than twice as long as wide; IV - X of gradually decreasing length; X approximately as wide as long (Fig. 39). Pronotum 1.05 - 1.10 times as wide as long and about 0.95 times as wide as head; microsculpture distinct, ren- dering the extremely fine puncturation almost invisible (Fig. 39). Elytra approximately 1.1 times as wide and at suture about as long as pronotum; puncturation very dense, fine, and weakly granulose. Hind wings of reduced length, about' twice as long as elytra. Legs long and slender; metatarsomere I 4 - 5 times as long as wide; I - IV of decreasing length, IV weakly oblong (Fig. 41); protarsus with pronounced sexual dimorphism. Abdomen with very fine and dense puncturation; poste- rior margin of tergite VII with narrow palisade fringe. ¢: protarsomeres strongly transverse and cordiform (Fig. 40); posterior margin of sterniteVII broadly con- cave, laterally with combs of about 10 palisade setae and on either side of middle with tuft of long black setae (Fig. 42); sternite VII not distinctive (Fig. 43); aedeagus as in Figs. 44 - 45. ©: protarsomeres weakly transverse and not cordiform. Etymology: The name (Lat.) is a noun in apposition and means cave inhabitant. Comparative notes and systematics: Medon antricola is readily identified by external characters alone, espe- cially the large size, uniformly light coloration, the re- duced eyes, long antennae and legs, the oblong head, as well as by the puncturation and microsculpture of the head and pronotum. Based on the evident similarities in the shape and chaetotaxy of the male sternite VII, the morphology of the aedeagus, the oblong head, as well as on the puncturation and microsculpture of head and pronotum, M. antricola is doubtlessly the sister species of M. subcoriaceus. It is not closely allied to M. feloi, another cave-dwelling species from the Canary Islands (see section 3.25). Distribution and bionomics: The species is endemic to El Hierro, Canary Islands, where it was found at least in two caves, the Cueva de Jinama and the Cueva de Fileba in the north of the island. Larvae possibly be- longing to this species were also found in caves in the south of El Hierro (P. OROMI, La Laguna, pers. comm. 2003). As can be inferred from the adaptive reductions of the eyes, wings, and pigmentation, as well as from the conspicuously long antennae and legs, M. antricola is a true troglobite. This is also supported by the cir- cumstances of collection: the type specimens were found by handsampling (on bare ground) and with pit- fall traps in the deep and dark parts of the caves, partly near water (P. OROMI, La Laguna, pers. comm. 2003). One specimen taken in November is teneral. 3.8. Medon castaneus (Gravenhorst, 1802) (Figs. 46-52, Map 6) Paederus castaneus Gravenhorst, 1802 (GRAVENHORST 1802: 60). Lathrobium brevicorne Latreille, 1804 (LATREILLE 1804: 342). Paederus quadratus Beck, 1817 (BECK 1817: 25). Volker ASSING: Western Palaearctic Medon 49 Medon ruddii Stephens, 1833 (STEPHENS 1833: 273). Types examined: P. castaneus: Lectotype ©, present designation: 6361 / castanea Er., Paederus Gr. / Synty- pus Lathrobium castaneum Gravenhorst 1802, labelled by Zool. Mus. Berlin, 7.2003 / Hist.-Coll. (Coleoptera) Nr. 6361 Lathrobium castaneum Gravenhorst, 1802, Eu- ropa, Zool. Mus. Berlin / Lectotypus 4 Paederus casta- neus Gravenhorst, desig. V. Assing 2003 / Medon cas- taneus (Gravenhorst) det. V. Assing 2003 (MNHUB). Paralectotype Y: Syntypus Lathrobium castaneum Gravenhorst 1802, labelled by Zool. Mus. Berlin, 7.2003 / Hist.-Coll. (Coleoptera) Nr. 6361 Lathrobium castaneum Gravenhorst, 1802, Europa, Zool. Mus. Ber- lin / Paralectotypus Y Paederus castaneus Gravenhorst, desig. V. Assing 2003 / Medon castaneus (Gravenhorst) det. V. Assing 2003 (MNHUB). Material examined (total: 92 exs.): Portugal: 1 ex., Serra de S. Marmede, Marvao, 39°24N, 07°23W, 665 m, swept from vegetation, 16.111.2002, leg. Meybohm (cAss); 1 ex., Castelo Branco, S Mantel- gas, 1450 m, 16.1V.1960, leg. Besuchet (cSch). Spain: | ex., Castilla-León, Segovia, San Rafael, 14.V.1960, leg. Comellini (cBor). France: | ex., Gironde, Vendays-Montalivet, IX.1950, lectaiempere .(MHNG); 1 ex., Gironde, Pessac (MHNG); 1 ex., Poutou-Charentes, St. Georges-de- Didonne, VIII.1925 (MHNG); 1 ex., Rhone-Alpes, Ar- déche, Saint-Agreve, 900m, 111.1934, leg. Gaudin (U- CBA, cZan); 1 ex., Lyon, 6.1V.1950 (MHNG); 1 ex., Alsace, Strasbourg, X.1953 (MHNG). Locality not i- dentified or ambiguous: | ex., Ft. de St. Germain, 819352=les.. Tempere (MHNG); 1 ex., St. Germain, 24.1.1937, leg. Levasseur (MHNG). England: | ex., Surrey, Pyrford, 26.11.1992, leg. Owen (cOwe). Sweden: 2 exs., Skane, N Landskrona, Alabodarna, vole burrow, 21.V.1971 & 25.V.1975, leg. Gillerfors (cGil). Germany: Nordrhein-Westfalen: | ex., Miinster, Gas- selstiege, 111.1991, leg. Feldmann (cFel). Rheinland- Pfalz: 1 ex., Landau, Rúlzheim, meadow, mole nest, 11.1987, leg. Persohn (cKöh). Schleswig-Hostein: | ex., Krummesse, 22.111.1973 (MHNG). Niedersachsen: 2 exs., Hannover, Herrenháuser Gärten, 24.11.1988, leg. Assing (cAss); 1 ex., Hannover-Ahlem, fallow area, pit- fall trap, 5.V.1992, leg. Sprick (cAss); 1 ex., Harz, Herzberg, pitfall trap, 5.V.1992 (cAss); 2 exs., Hann.- Miinden, left bank of Fulda river, 6.1.1932, leg. Kirch (cSch); 2 exs., Holzminden, leg. Gerhard (NHMW). Hessen: 2 exs., Seeheim-Jugenheim, fallow, mole nest, 11.2003, leg. Hetzel (cFel); 1 ex. [teneral], Karlshafen, 27.1.1934, leg. Folwaczny (MHNG); 2 exs., Erlensee, 10.11.1993, leg. Hóhner (cKóh). Bayern: | ex., Bam- berg, leg. Weise (NHMW). Brandenburg/Berlin: | ex., Frankfurt/O., Markendorf, Helenesee, 22.1V.1984, leg. Gasche (cSch); 4 exs., W Potsdam, Wildpark Golm, mole nest, coll. Neresheimer (DEI); 1 ex., Hónow, mole nest, coll. Neresheimer (DEI); 1 ex., Mittenwalde, mdle nest, coll. Neresheimer (DEI); 10 exs., Strausberg, mole nest, coll. Neresheimer (DEI); | ex., NSG Rietzer See, 52°23N, 12°41E, Helsberg, rabbit burrow, fish bait, 16.V.1979, leg. Uhlig (DEI). Sachsen: | ex., Leipzig, 19.X.1913, leg. Linke (MHNG); 1 ex., Leipzig, Délzig, leg. Linke (NHMW). Austria: Vorarlberg: 2 exs., Feldkirch (NHMW). Ti- rol: 1 ex., Innsbruck, 4.X11.1932, leg. Pechlaner (NHMW); 1 ex., Innsbruck, leg. Strupi (MHNG). O- berösterreich: | ex., Ostermiething, mole nest, leg. Leeder (NHMW). Niederósterreich/Wien: | ex., O- berwaltersdorf, mole nest, leg. Franz (NHMW); 2 exs., Wien, Lobau, mole nest, 21.11.1926, leg. Breit (NHMW); 1 ex., Moosbrunn, mole nest, 1.111.1927, leg. Breit (NHMW); 3 exs., Wien, leg. Schade (NHMW, cAss); 12 exs., Purgstall, mole nests, 14.11.1957, 1.1V.1958, 13,XI1.1958, _4.11.1959, 31.111.1959, 29.18.1959, 4.%.1959 (83 exs.), 5.1959. 4.VD.1960, 20.X.1960, leg. Ressl (NHMW); 1 ex., Oberweiden, 11.11.1968, leg. Gotz (NHMW). Burgenland: | ex., Salmannsdorf (NHMW); 1 ex., Zurndorf, leg. Franz (NHMW). Italy: Trentino-Alto Adige: 1 ex., Bressanone, 24.V.1957, leg. Peez (cBor); 1 ex., Bressanone, 2.111.1953, leg. Peez (cZan); 1 ex., Piemonte, Novara, Invorio, 111.1971, leg. Rosa (cZan). Czech Republic: | ex., Bohemia, Hostivice (MHNG); 1 ex., Prag (NHMW). Poland: | ex., Lubin, 6.11.1916 (NHMW). Slovakia: 3 exs., Banskä Bystrica, X1.1923, leg. Roubal (MHNG, cBor). Locality illegible or not specified: | ex., 16.X.1960 (MHNG); 2 exs. (MHNG). Comments: The original description of Paederus cas- taneus is based on an unspecified number of type specimens, so that the two types found in the historical collection of the MNHUB are attributed syntype status. In order to stabilize the present interpretation of the spe- cies, the male syntype is here designated as the lecto- type. Diagnosis: Large species, 6.0-8.0 mm. Habitus as in Fig. 46. Head dark brown to blackish brown, sometimes with the frons ferrugineous; pronotum dark brown to blackish, with the margins and especially the anterior angles more or less extensively ferrugineous; elytra ru- fous to castaneous; abdomen dark brown to blackish, 50 Bonner zoologische Beitráge 54 (2005) with the apex, paraterga, and the tergal hind margins lighter; legs and antennae rufous. Head at least weakly oblong and of subquadrangular shape, 1.e. with subparallel lateral margins and marked posterior angles; eyes less than half the length of posto- cular region in dorsal view; puncturation moderately coarse, well-defined, and dense, with the interstices much narrower than the punctures, in median dorsal area te N Kae > > ALO = (Up 3 a = / > Ge L 1 SES > Dan sa ad (kar e DR ON l \ i Fy SE 4 ze 2 A ON 4 EN Qe ee y ¡E \ x @ 3 mee a O >) oe Ñ @ == | Cet < y 3 2 en N, <> po. x Y 0 le Mn 3 PR $ 7 / e Ne | y \ ; Y o A ¿0 , Oo y ú wie MER / ® PN an usually sparser and with shining interstices (Fig. 47); shallow microsculpture may be present in posterior and lateral areas. Antenna very long and slender, antennomere IV and V more than twice as long as wide (Fig. 46). Pronotum as wide as long or weakly transverse, slightly to distinctly narrower than head; puncturation denser and less well defined than that of head, often partly conflu- ent; interstices narrow, but more or less shining (Fig. 47). Map 6: Distribution of Medon castaneus (Gravenhorst) based on revised (filled circles) and selected literature records (open circles). Volker ASSING: Western Palaearctic Medon 51 Elytra 1.15-1.20 times as wide and at suture 1.05-1.18 times as long as pronotum (Fig. 46); puncturation dense and granulose, less coarse than that of head and prono- tum. Hind wings fully developed. Legs very long and slender, metatarsus almost as long as metatibia; metatar- somere III at least about 3 times as long as wide. Abdomen with very fine and dense puncturation and with microsculpture; posterior margin of tergite VII with palisade fringe. d: protarsomeres distinctly dilated, protarsomere II cordiform and wider than long; posterior margin of ster- nite VII distinctly concave and with two combs of pali- sade setae, each consisting of usually more than 10 setae (Fig. 48); aedeagus long and slender (Figs. 49-52). ©: protarsomeres weakly dilated, protarsomere II not wider than long and not distinctly cordiform. Comparative notes: Medon castaneus is easily distin- guished from all its Western Palaearctic congeners, ex- cept for M. procerus (see the following section), espe- cially by its large size, the elongate legs and antennae, and by the long and slender aedeagus. Distribution and bionomics: Medon castaneus repre- sents an Atlanto-Mediterranean element (Map 6), its distribution ranging from the western part of the Iberian Peninsula to southern England, southern Scandinavia, Poland, western Russia, Ukraine, Slovakia, and Hun- gary (material examined). For more records see e.g. BARANOWSKI (1977, 1979), BOHÁC (1985), BRUGE et al. (2001), BÚCHE & ESSER (1999), DRUGMAND (1989), ERMISCH & LANGER (1933), FALCOZ (1914), FOWLER (1888), HANSEN (1996), HANSEN et al. (1995), HANSEN (1964), HORION (1965), KOCH (1968), KOHLER (2000), KOHLER & KLAUSNITZER (1998), LUNDBERG (1995), TERLUTTER (1995), and TRONQUET (2001). In Scandi- navia, it is present only in Denmark and Skane. Accord- ing to HORION (1965), there are old records from Slove- nia and Hungary, and also from Ukraine (Wolyns'ka Oblast, Kiev) and Russia as far east as Samara, but these require confirmation. SILFVERBERG (1992) reports the species from Lithuania. Confirmed records from Italy are only known from two localities in the north (mate- rial examined; OSELLA & ZANETTI (1974)); it 1s also in- dicated from Lazio (southern Italy) (CICERONI & ZANETTI 1995; LUIGIONI 1929; PORTA 1926), but this old record has not been confirmed and should be con- sidered doubtful. Medon castaneus is nidicolous and usually associated with the nests and burrows of the European mole (Talpa europaea Linné), which, however, is absent from the western Iberian Peninsula. Thus, M. castaneus may also inhabit the nests of Desmana pyrenaica (Geoffroy Saint Hilaire) or Talpa caeca (Savi), which occur in this re- gion. The observation that the species has also been found in rabbit burrows on two occasions, once on fishbait (material examined; BARANOWSKI (1979)), and once in vole burrows suggests that the host spectrum may also include other mammals with subterranean nests and burrows. / Nests of moles are inhabited by rather many species of Coleoptera and are relatively easy to find and to dig up, which is why there are numerous thorough studies on the beetle fauna of this habitat (e.g. HEINEMANN 1910; HORION 1933; NOWOSAD 1990; OSELLA & ZANETTI 1974; STROUHAL & BEIER 1928). According to these in- vestigations, M. castaneus is rather rarely found in most regions. In the environs of Braunschweig (Germany: Niedersachsen), HEINEMANN (1910) found only 2 speci- mens in 225 nests. In the surroundings of Wien, STROUHAL & BEIER (1928) dug up 118 nests and col- lected 21 beetles. The by far most comprehensive and systematic study of the staphylinid fauna of mole nests was carried out in Poland by NOWOSAD (1990), who re- corded M. castaneus in only 28 nests (48 specimens) of a total of about 5000 nests studied. The literature data on the preferred biotope are diverse or even contradictory. According to most authors (e.g. JOY 1932; HORION 1933, 1965; STROUHAL & BEIER 1928; VOGT 1956), M. castaneus is usually found in damp situations, especially near rivers and streams, whereas LINKE (1927) reports it from dry places; on one occasion he found 14 specimens in a single nest. Adult beetles have been found almost throughout the year, except in August. The low number of summer re- cords, however, may also be due to the probably low number of nests examined during this period. Larvae have been found in July (BOHAC 1985, HORION 1965). A teneral adult was collected in January (material exam- ined). One specimen was swept from vegetation in March (Portugal) and two specimens were caught with pitfall traps in April and at the beginning of May, sug- gesting that dispersal by flight and on the ground proba- bly takes place in early spring. 3.9. Medon procerus (Perez Arcas, 1874), sp. propr. (Figs. 53-57) Lithocharis procera Perez Arcas, 1874 (PEREZ ARCAS 1874: 114 ff). Material examined: Spain: 2 exs., Madrid, Cueva de la Magdalena (NHMW, cAss). Diagnosis: Largest species of the genus in the Western Palaearctic region, 7.0-9.0 mm (Fig. 53). Coloration uniformly ferrugineous. Head very large and of ellipsoid shape, i. e. lateral mar- gins convex and posterior angles almost obsolete; eyes slightly smaller than in M. castaneus; puncturation as in 52 Bonner zoologische Beitráge 54 (2005) M. castaneus, but slightly less coarse (Fig. 54). Anten- nae of similar morphology as in M. castaneus. Pronotum small in relation to head, distinctly tapering posteriad; puncturation less dense and more well de- fined than is usually the case in M. castaneus (Fig. 54). Legs even longer and more slender than in M. casta- ~ neus; length of metatibia 1.7-1.8 mm (maximum length in largest specimens of M. castaneus 1.4 mm). Elytra and abdomen as in M. castaneus (Fig. 53). $: protarsomeres I - IV distinctly dilated; sternite VII of similar morphology and chaetotaxy as in M. castaneus, but more oblong and posterior concavity shallower in relation to total length of sternite (Fig. 55); sternite VIII elongate and slender; aedeagus longer than in M. casta- neus and with longer and more slender ventral process (Figs. 56-57). Figs. 46-57. Medon castaneus (Gravenhorst) (46-52) and M. procerus (Perez Arcas) (53-57): Habitus (46, 53); head and prono- tum (47, 54); male sternite VII (48, 55); aedeagus in lateral and in ventral view (49-57; 49, 50: northern Germany; 51, 52: Portu- gal). Scale bars: 46-48, 53-55: 1.0 mm; 49-52, 56-57: 0.5 mm. Volker ASSING: Western Palaearctic Medon 53 Comparative notes: The species is distinguished from the similar M. castaneus especially by the uniformly ferrugineous coloration, the ellipsoid shape of the head, the longer legs, and the longer and more slender aedeagus. From all other Western Palaearctic congeners, it is readily separated by its much greater body size alone. Comments: The types of this species were not exam- ined, but in view of the distinctive external appearance and the fact that the non-type specimens listed above were collected at the type locality, there is no doubt that the present interpretation is correct. M. procerus was previously regarded as a synonym of M. castaneus (e.g. COIFFAIT 1984). However, the constant differences in external morphology, in the shape of the primary and secondary sexual characters, as well as the different habitat suggest that it represents a distinct species. A similar case of adelphotaxa, one of them widespread and the other an endemic troglobite, is known from the Eastern Mediterranean (ASSING 2004a): Medon fusculus and M. dobrogicus Decu & Georgescu. Distribution and bionomics: This troglobite 1s known only from the Cueva de la Magdalena, a cave near Ma- drid, where it is apparently endemic. According to PEREZ ARCAS (1874) adult beetles were found almost throughout the year. 3.10. Medon pocofer (Peyron, 1857) (Figs. 58-59) Lithocharis pocofera Peyron, 1857 (PEYRON 1857: 718ff). Lithocharis maritimus Aubé, 1863 (AUBE 1863: 36). Material examined (total: 51 exs.; for additional mate- rial see ASSING (2004a)): France: Provence: 2 exs., locality not specified (NHMW); 1 ex., Port Miod near Cassis, “feinkiesiger bis grobschottriger Untergrund”, 10.VIII.1959, leg. Schuster (NHMW); 1 ex., Nice, 111.1954 (cBor); 1 ex., Nice, 25.1X.1918 (MHNG); 10 exs., Nice, 5.11.1954 (MHNG, cAss); 2 exs., La Seyne (NHMW); 3 exs., Saint-Raphaél, leg. Fauvel (NHMW, cAss); 1 ex., Fre- jus, VII.1892, leg. Rey (NHMW); 1 ex., Toulon, leg. Bauduer (NHMW); 1 ex., Toulon (DEI); 1 ex., Var, lo- cality not specified (DEI); 1 ex., Var inundation [W Nice], Alpes Maritimes, IV.1859 (DEI); 1 ex., Antibes (DED. England: 1 ex., Dorset, Lulworth Cove, shingie, 22.111.1990, leg. Owen (cOwe). Italy: Friuli-Venezia Giulia: | ex., Trieste, Sistiana, Adria, leg. Breit (NHMW); 1 ex., Trieste, Grignano, IX.1907, leg. Krekich (MHNG). Liguria: 7 exs., Geno- va, XII.1900, leg. Dodero (NHMW, cAss); 1 ex. [tene- ral], Genova, VII.1899, leg. Dodero (NHMW); 2 exs., Genova, 111.1894, leg. Dodero (NHMW ); 4 exs., Geno- va, 1V.1893, leg. Dodero (DEI, cAss); 2 exs., Genova, 1.1918, leg. Dodero (MHNG); 1 ex.,-Genova, X1.1918, leg. Dodero (NHMW); 4 exs., Genova, leg. Dodero, Hauser (DEL NHMW). Umbria: | ex., locality not specified (NHMW). Diagnosis: See ASSING (2004a). Habitus as in Fig. 58. All tarsomeres III and IV are transverse and more or less cordiform (Fig. 59), a character not emphasized by ASSING (2004a), but distinguishing M. pocofer from all other Western Palaearctic species. Comments: The type material of M. pocofer was looked for, but not found, in the collections of the MNHNP. However, both the characters and the ecologi- cal details indicated in the original description leave no doubt that the present interpretation is correct. Distribution and bionomics: According to the litera- ture, Medon pocofer is distributed on the Atlantic coasts northwards to southern England and in the Western Mediterranean, including Algeria and Tunisia (COIFFAIT 1984; FOWLER 1888; NORMAND 1935). The easternmost record is from Montenegro (ASSING 2004a). In the Western Mediterranean, I have seen material only from southern France (Provence) and Italy. LUIGIONI (1929) reports the species from various Italian regions (Liguria, Venezia Giulia, Lazio, Sardinia, and Sicily) and from Corsica. Based on the material examined, the species is not rare at the coasts of Var, Alpes Maritimes, and Lig- uria. Among Western Palaearctic Medon species M. pocofer is the only representative that is apparently confined to coastal habitats. It is usually found under seaweed and under stones between shingles (COIFFAIT 1984; FOWLER 1888; Joy 1932; NORMAND 1935). FOWLER (1888) ob- serves that the specimens are very quick and immedi- ately hide between shingles when disturbed. One speci- men collected in July was teneral (material examined). 3.11. Medon rufiventris (Nordmann, 1837) (Figs. 60-61, Map 7) Lathrobium rufiventre Nordmann, 1837 (NORDMANN 1837: 147f). Medon anatolicum Coiffait, 1970a (COIFFAIT 1970a: 704ff); synonymy by ASSING (2004a). Medon burdigalensis Coiffait, 1970a (COIFFAIT 1970a: 711f) syn. n. Medon aquitanicum Coiffait, 1970a (COIFFAIT 1970a: 712) syn. n. Medon siculum Coiffait, 1970a (COIFFAIT 1970a: 709ff) syn. n. 54 Bonner zoologische Beitráge 54 (2005) Medon sicilianum Coiffait, 1970c (COIFFAIT 1970c: 712); replacement name for M. siculus Coiffait; syn. n. Types examined: M. burdigalensis: Holotype Ö [ae- deagus missing]: Gironde V.45, Gazinet, H. Coiffait / MUSEUM PARIS COLL H. COIFFAIT / HOLOTYPE / Medon burdigalensis Coiff. H. COIFFAIT det. 1969 / Medon rufiventris (Nordmann) det. V. Assing 2003 (MNHNP). M. aquitanicus: Holotype ©: Gironde, Cussac / Collec- tion H. Coiffait / MUSEUM PARIS COLL H. COIF- FAIT / HOLOTYPE / Medon aquitanicum Coiff. H. COIFFAIT det. 1969 / Medon rufiventris (Nordmann) det. V. Assing 2003 (MNHNP). M. siculus: Holotype Ö: U. Lostia, Sardinia / MUSEUM PARIS COLL H. COIFFAIT / HOLOTYPE / Medon siculum Coiff. H. COIFFAIT det. 1969 / Medon rufi- ventris (Nordmann) det. V. Assing 2003 (MNHNP). M. anatolicus: Holotype Ö: Turquie (Isparta), Bodemli, H. Coiffait, 27.V.54 / HOLOTYPE / Medon anatolicum Coiff. H. COIFFAIT det. 1969 (MNHNP). Paratypes: 14, 12: same data as holotype (MNHNP). Comments: According to COIFFAIT (1970a), M. burdi- galensis is distinguished from M. rufiventris by the finer puncturation of the pronotum and by the different shape of the aedeagus. The puncturation of the pronotum is indeed very fine in the holotype, but this character ıs highly variable in the species and I have seen specimens with a similarly punctured pronotum also from other parts of the range of M. rufiventris, e.g. from Greece. The aedeagus of the holotype is apparently lost, but the drawing provided in the original description is quite in agreement with the aedeagus of M. rufiventris. The api- cal emargination is rather small, but this is probably an artefact resulting from mounting a dry aedeagus. The fact that the illustration of the aedeagus of M. rufiventris in COIFFAIT (1984: 19, figs. 6] and 6J) is taken from the literature suggests that Coiffait had not seen a male M. rufiventris himself. In the original description of M. aquitanicus, COIFFAIT (1970a) states that the species is similar to M. burdi- galensis, but separated by the more parallel shapes of the head and pronotum, by the more numerous black palisade setae at the posterior margin of the male ster- nite VII, the lower distance between the two clusters of palisade setae, and by the different shape of the aedeagus. Head shape is an extremely variable character in M. rufiventris; the majority of specimens have a rela- tively large and somewhat wedge-shaped head, but the head may also be of subparallel shape and/or of reduced size (ASSING 2004a). The apex of the aedeagus (Fig. 61) differs somewhat from the usual condition encountered in M. rufiventris, but teratological aedeagi are a com- mon phenomenon in the genus. Neither the male secondary sexual characters nor the other external characters of the holotypes of M. burdi- galensis and M. aquitanicus (Fig. 60) provide sufficient evidence that these specimens should represent distinct species (not to mention the unlikelihood of endemic Medon species occurring in Gironde), so that both M. burdigalensis and M. aquitanicus is here placed in the synonymy of M. rufiventris. The type locality indicated in the original description of M. siculus (“Sicile”) is clearly incorrect; the label at- tached to the holotype gives “Sardinia”. This error has resulted in the conclusion that M. siculus (later known under the replacement name M. sicilianus) is endemic to Sicily (COIFFAIT 1984; CICERONI & ZANETTI 1995). COIFFAIT (1970a) compares M. siculus with M. brun- neus, probably because of the relatively coarse punctu- ration of the pronotum of the type specimens, but an ex- amination of the holotype revealed that it is conspecific with the senior name M. rufiventris. The puncturation of the pronotum and of the head is indeed coarser than is usually the case in the species, but, like the size and the shape of the head, the puncturation of the forebody is subject to pronounced intraspecific variation in this spe- cies and the condition found in the holotype is still within the range observed in M. rufiventris. An exami- nation of other characters, including the male secondary sexual characters and the morphology of the aedeagus, did not produce any evidence that the types of M. sicu- lus should represent a distinct species. Based on non-type material and the original description (COIFFAIT 1970a), the synonymy of M. anatolicus with M. rufiventris was already established by ASSING (2004a). A recent examination of the types of M. anato- licus confirms this synonymy. Additional material examined (total: 48 exs.; for addi- tional material see ASSING (2004a)): Spain: | ex., Cuenca, Serrania de Cuenca, 40°07N, 02°02W, 17.V.2002, leg. Starke (cAss). A France: 2 exs., Landes, Sos (NHMW); 2 exs., Ile-de- France, Fontainebleau, IV.1904, leg. Mequignon (MHNG); 1 ex., Fontainebleau, leg. Mequignon (MHNG); 1 ex., Fontainebleau, 20.111.1905, leg. Gruar- det (MHNG); 1 ex., Var, St.-Paul-en-Forét, 5.1.1939 (MHNG); 4 exs., Var, Figanieres, 29.1V.1975, leg. Zola (UCBA, cZan); 2 exs., Var, Bagnols-en-Forét, cork tree, IV.1937 (MHNG); 2 exs., Var, Plan de Canjuers, 43°42N, 06°18E, 22.1V.1954 (MHNG); 1 ex., Var, loca- lity not specified, 11.1950 (MHNG). Volker ASSING: Western Palaearctic Medon Un in Map 7: Distributions of Medon rufiventris (Nordmann) (black circles: revised records; grey circles: selected literature records) and of M. africanus (Fauvel) (white circles; examined records only). Luxemburg: 1 ex., locality not specified, leg. Franz (cAss). Italy: 1 ex., Lombardia, Berbenno di Valtellina, Colori- na, 4.1V.1972 (cZan); 1 ex., Piemonte, Brandizzo, mole nest, 28.X11.1973, leg. Osella (cZan); 1 ex., Veneto, Grezzana (VR), 10.1V.1992, leg. Zanetti (cZan); | ex., Elba, 22.-31.V.1993, leg. Rauhut (cAss); 7 exs., Elba, Mt. Capanno, 600 m, under chestnut bark, 30.111.- 13.1V.1921, leg. Moczarski & Scheerpeltz (NHMW, cAss). Germany: | ex., Sachsen-Anhalt, Dessau, leg. Nebel (NHMW). Slovakia: | ex., Hronskä Breznica [48°34N, 19°00E], leg. Roubal (MHNG). Greece: 11 exs., Métsovon, 1200 m, 8.1V.1993, leg. Frisch (MNHUB, cAss); 4 exs., Pelopönnisos, Olympia, Ladzoi, 100 m, 25.11.1992, leg. Frisch (MNHUB, cAss); 1 ex., Pelopónnisos, Taygetos, Katafigion, 1550 m, 10.1X.1995, leg. Zola (cZan). Belarus: 1 ex., Gomel region, Simonitschi, National Park “Pripiatsky”, 20.V11.1998, leg. Sheshurak (cGon). Diagnosis: See ASSING (2004a). Distribution and bionomics: Based on the material ex- amined, M. rufiventris is evidently a Ponto-Mediterran- ean element, its distribution ranging from eastern Ana- tolia, Krasnodar, and Belarus in the southeast and east to the central parts of Poland and Germany and to Lux- emburg in the north, and finally to western France, and northeastern Spain in the west (material examined). There is an isolated record from the Swedish island Oland, the only Scandinavian locality known for the species (LUNDBERG 1995). For additional records see BOHAC (1985), HORION (1965), KOHLER (2000), KOHLER & KLAUSNITZER (1998), LUNDBERG (1995), SCHEERPELTZ (1968), SZUJECKI (1968), and TRONQUET (2001). The species is here recorded from Luxemburg for the first time. It is unknown from the British Isles (LOTT € DUFF 2003), Belgium (BRUGE et al. 2001), and from the Baltic countries (SILFVERBERG 1992). Rather recent German records are from Berlin and Sachsen- 56 Bonner zoologische Beitráge 54 (2005) Anhalt (KORGE 1989; SCHÚLKE 1998). The presence of the species in Sardinia was considered doubtful by CI- CERONI & ZANETTI (1995), but is here confirmed through the holotype of M. siculus Coiffait. In view of the general distribution pattern of the species, as well as of the absence of any confirmed records, the presence of the species in Northwest Africa, as indicated by COIF- FAIT (1984) and HORION (1965), must be considered most unlikely. Previous records from this region are evidently based on confusion with other species, espe- cially the externally similar M. africanus. Medon rufiventris was mostly found associated with dead wood, especially in or under bark of old or dead tree trunks of deciduous trees (various species of oak, beech, chestnut) and pine trees (material examined; see also ASSING (2004a), HORION (1965), KORGE (1989), and VOGT (1968)). One specimen was collected from a mole nest (OSELLA & ZANETTI 1974; specimen exam- ined). The examined beetles (see also ASSING 2004a) were collected during the period from December through July, and in October, the vast majority of them in spring (March through June). One of three specimens taken in October was teneral (ASSING 2004a). 3.12. Medon africanus (Fauvel, 1872) (Figs. 62-69, Map 7) Lithocharis africana Fauvel, 1872 (FAUVEL 1872: 38). Medon bodemeyeri Bernhauer, 1915 (BERNHAUER 1915: 266) syn. n. A Types examined: L. africana: Lectotype 6, present designation: Bone / africanus Fvl. type / Coll. et det. A. Fauvel Medon rufiventris Nordm. R.1.Sc.N.B. 17.479 / TYPE / Medon africanus (Fauvel) det. V. Assing 2003. M. bodemeyeri: Lectotype Y: Ain Draham, Tunis, B. V. Bodemeyer / Bodemeyeri Bernh. Typus / Chicago NHMus, M. Bernhauer Collection / Lectotypus Medon bodemeyeri Y Bernhauer desig. V. Assing 2003 / Me- don africanus (Fauvel) det. V. Assing 2003 (FMNH); 19: same data, but “Cotypus ...” (FMNH); 39 Y: same data, but without type labels (DEI, FMNH); 14: Ain Draham, Tunis, B. V. Bodemeyer / Syntypus / Medon Bodemeyeri Bernh. / O. Leonhard / Coll. DEI Ebers- walde / Paralectotypus Medon bodemeyeri 4 Bernhauer desig. V. Assing 2003 / Medon ripicola (Kraatz) det. V. Assing 2003 (DEI). Comments: FAUVEL (1872) neither specified the num- ber of types nor did he designate a holotype. Therefore, in order to fix the identity of M. africanus, the male syn- type in the Fauvel collection is here designated as the lectotype. The original description of M. bodemeyeri is based on an unspecified number of syntypes (BERNHAUER 1915). Five of them were found in the collections of the FMNH and the DEI; four of them — the Y Y — are conspecific with M. africanus (Fauvel), the male is conspecific with M. ripicola (Kraatz). The original description is in better agreement with the YY, one of which was labelled as “Typus” by Bernhauer. In view of the fact that the type series consists of two species, a lectotype designation is mandatory. The female with Bernhauer’s type label was selected as the lectotype and M. bodemeyeri Bernhauer is placed in the synonymy of M. africanus (Fauvel). Additional material examined (total: 51 exs.): Tunisia: | ex., 2 km E Ain Sobah, 18 km E Tabarka, 100 - 200 m, 1.X.1995, leg. Schulz & Vock (cAss); 3 exs., “Tunis” (NHMW); 1 ex., Ain Draham (= Ayn ad Darahim; 36°47N, 8°42E) (cAss); 1 ex., same locality, 6.1V.1962, leg. Besuchet (MHNG); | ex., same locality, VI.1884 (IRSNB); 2 exs., ca. 2 km S Ain Draham, 36°44N, O8°41E, 670 m, litter of oak forest, 28.X11.2004, leg. Assing (cAss); 2 exs., El Feidja (IRSNB); 11 exs., Aín Soltane, near Ghardimaou, 30.1V.2004, leg. Lackner (cAss). Algeria: | ex., Laverdure [=Mechroha], 7.X.1929, leg. Schatzmayr (FMNH); 6 exs., Bou Berak, Dellys (IRSNB, cSch); 2 exs., Bou Berak (IRSNB, NHMW); 1 ex., Grande Kabylie, Forét d'Akfadou, Adekar, 1300 m, 15.V.1988, leg. Besuchet, Lóbl & Burckhardt (MHNG); 2 exs., Annaba [*Bóne”] (NHMW); 1 ex., Idügh [E- dough] (MHNG); 2 exs., Constantine (MHNG, cAss); 1 ex., Mt. Babor (IRSNB); 1 ex., St. Antoine, leg. Thery (IRSNB); 1 ex., Theniet el Had [“Teniet el H.”], cedar forest (IRSNB); 1 ex., Theniet el Had, 30.V.-5.V1.1954, leg. Fagel (IRSNB); | ex., Ain Babouche [“Babouch”'] (IRSNB); | ex., Philippeville (IRSNB). Locality not identified: 2 exs., Ternano (?) (MHNG). Diagnosis: 3.7-5.0 mm. Habitus as in Figs. 62-63. Head, pronotum, and anterior segments of abdomen dark brown to blackish brown; elytra and abdominal apex rufous; legs and antennae ferrugineous. Head (Figs. 64-65) approximately as wide as long or weakly oblong, of similar shape as in M. cauchoisi; eyes relatively large, clearly more than half the length of postocular region in dorsal view; puncturation dense, relatively coarse, and non-areolate; interstices mostly narrower, in central dorsal area often wider than punc- tures, usually without microsculpture. Pronotum as wide as or slightly wider than head (Figs. 64-65); puncturation similar to that of head or slightly less coarse; microsculpture absent. Elytra about 1.5 times as wide and at suture 1.15-1.20 times as long as pronotum; puncturation fine and dense. Hind wings fully developed. Volker ASSING: Western Palaearctic Medon 57 Abdomen with fine and dense puncturation; posterior margin of tergite VII with palisade fringe. ¢: posterior margin of sternite VII distinctly concave and with two combs of about 10 or more palisade setae (Fig. 66); aedeagus with apex of ventral process of dis- tinctive morphology (Figs. 67-69). Intraspecific variation: The species is extremely vari- able, especially regarding the puncturation (size and density of punctures) of the forebody (Figs. 62-64). Comparative notes: Medon africanus is most similar to M. cauchoisi and M. rufiventris. From the former, it 1s distinguished by the somewhat less coarse and less dense puncturation and the consequently more shining appearance of the head and pronotum, the relatively wider pronotum, the more convex posterior margin and the more numerous palisade setae of the male sternite VI, and by the different morphology of the aedeagus. Medon rufiventris often has a posteriorly dilated head, a finer and sparser puncturation of the head and prono- tum, a male sternite VII with a usually slightly less strongly concave posterior margin and with on average fewer palisade setae, and an apically emarginate aedeagus (ventral view). Distribution and bionomics: The species is currently known only from Tunisia and Algeria (Map 7). For ad- ditional Tunisian and Algerian localities see BORDONI (1988) and NORMAND (1935). Like M. rufiventris, M. africanus apparently lives under bark of deciduous and coniferous trees (NORMAND 1935; PEYERIMHOFF 1919). Two specimens from Tunisia were sifted from the leaf litter of an oak forest. The material with specified dates on the labels was found in April - June, October, and December. 3.13. Medon vicentensis Serrano, 1993 (Figs. 70-72) Medon vicentensis Serrano, 1993 (SERRANO 1993: 4ff). Material examined: Madeira: | ex., Säo Vicente, Gruta dos Cardais, pitfall trap, 5.1.1996, leg. Erber (cErb). Diagnosis: For a more detailed description see SERRANO (1993). 4.4-5.4 mm. Habitus as in Fig. 70. Coloration of whole body uniformly ferrugineous. Head large, approxi- mately as wide as long; eyes reduced to minute rudi- ments, without ommatidia; puncturation distinct, mod- erately coarse and moderately sparse, interstices, except for the sparsely punctured dorso-median area, on aver- age about twice as wide as punctures; surface with pro- nounced microreticulation and almost matt (Fig. 71). Antenna slender, antennomeres III - V more than twice as long as wide; VI about twice as long as wide; VII about 1.5 times as long as wide (Fig. 72). Pronotum slightly narrower than head; approximately as wide as long; puncturation finer and less distinct than that of head, median line impunctate; microreticulatipn pronounced (Fig. 71). Elytra widest near posterior margin, anterior external angles almost obsolete; as wide as or slightly narrower than pronotum; at suture about 0.7 times as long as pronotum; puncturation very fine, ill-defined and mod- erately dense; with much weaker microsculpture and much more shine than head and pronotum (Fig. 71). Hind wings completely reduced. Abdomen with moderately dense, fine, slightly granu- lose puncturation; microsculpture present, but weaker than that of head and pronotum; posterior margin of ter- gite VII without palisade fringe. &: posterior margin of sternite VII of similar shape and chaetotaxy as in M. dilutus; aedeagus as illustrated by SERRANO (1993). Comparative notes: This species is readily distin- guished from its Western Palaearctic congeners by the almost completely reduced eyes, the long and slender antennae, the reduced pigmentation and wings, and by the male sexual characters. The only other troglobiont Medon species with almost completely reduced eyes, i. e. without ommatidia, in the Western Palaearctic are M. feloi from La Palma and M. antricola from El Hierro, Canary Islands. These species are also similar in the dis- tinctly microreticulate and rather shallowly punctate head and pronotum. Medon feloi, however, has shorter antennae, a more convex (cross-section) and more ob- long head, a more slender and posteriorly less distinctly tapering pronotum, longer elytra, not distinctly dilated protarsomeres I - IV, a differently shaped aedeagus, and its male secondary sexual characters are similar to those of M. indigena. Medon antricola, on the other hand, is distinguished by much greater body size, larger eye ru- diments, longer and more slender legs and antennae, a much more oblong and posteriorly tapering head, and completely different male sexual characters. Distribution and bionomics: Medon vicentensis, a true troglobite, is a local endemic of Madeira proper, where it is known only from a cave system near Sao Vicente. 3.14. Medon augur Fauvel, 1906 (Fig. 73) Medon augur Fauvel, 1906 (FAUVEL 1906: 89). Type examined: Holotype Y: Corse / Croatie / augur Fvl.l / Ex-Typis / R. I. Sc.N.B. 17.479, Coll. et det. A. Fauvel (IRSNB). 58 Bonner zoologische Beitráge 54 (2005) Figs. 58-73. Medon pocofer (Peyron) (58-59), M. rufiventris (Nordmann) (holotype of M. aquitanicus Coiffait) (60-61), M. afri- canus (Fauvel) (62-69; 62: lectotype), M. vicentensis Serrano (70-72), and M. augur Fauvel (73, holotype): Habitus (58, 62, 63, 70, 73; 63: Algeria); protarsus (59); forebody (60, 71); head and pronotum (64, 65; 64: Algeria, 65: Tunisia); male sternite VII (66); aedeagus in lateral and in ventral view (61, 67, 68); apical part of aedeagus in ventral view (69); antenna (72). Scale bars: 58, 60, 62-66, 70-73: 1.0 mm; 59, 61, 67-69: 0.2 mm. Volker ASSING: Western Palaearctic Medon 59 Comments: The original description is based on a holo- type from Corsica (“Corse. — Unique”) and an addi- tional specimen from Croatia (“J“en ai vu un second ex- emplaire de Croatia”) (FAUVEL 1906). The label “Croatie” attached to the pin of the holotype suggests that Fauvel apparently exchanged the Croatian specimen with a colleague, but kept the locality label and added it to the pin of the type specimen from Corsica. Types and other material with different locality labels attached to the same pin are not uncommon in the Fauvel collection (see e.g. ASSING 1999b). It seems somewhat unlikely that the holotype is a repre- sentative of a distinct species with a distribution con- fined to Corsica. (The Croatian specimen attributed to this species in the original description probably refers to the similar M. dilutus pythonissa (Saulcy).) The holo- type could be an extremely large specimen of M. rufiventris. However, it cannot be assigned to this nor any of the other widespread species without doubt, so that it is here maintained as a valid species, until males become available from Corsica. Diagnosis: 4.8 mm. Habitus as in Fig. 73. Of similar general appearance as M. dilutus pythonissa. Head, pronotum, and abdomen (except apex and posterior margins of segments) dark brown; elytra, legs, and an- tennae ferrugineous. Head large and convex, as wide as long; eyes relatively small (about as small as in M. dilutus), not distinctly projecting from lateral outline of head and slightly less than one third the length of temples in dorsal view; puncturation coarse (similar to M. dilutus pythonissa and M. rufiventris), along median line without or with very sparse puncturation; microsculpture absent. Pronotum as wide as long, distinctly narrower than head; puncturation slightly less coarse than that of head; median line without punctures; microsculpture absent. Elytra 1.13 times as wide and at suture 0.98 times as long as pronotum. Protarsomeres I - IV dilated (about as wide as in M. rufiventris and M. dilutus pythonissa). Abdomen similar to that of M. dilutus pythonissa, but with slightly less distinct microsculpture; posterior mar- gin of tergite VII with palisade fringe. 4: unknown. Comparative notes: Medon augur is most similar to M. dilutus pythonissa, which is most unlikely to occur in Corsica and from which it is distinguished by the smaller and less prominent eyes and by the slightly shorter and narrower, uniformly ferrugineous elytra. Medon rufiventris is smaller, usually has a more wedge- shaped head, larger eyes and relatively longer and wider elytra. In the paler M. dilutus dilutus, the puncturation of the head and pronotum is finer and the median lines of head and pronotum are usually not free of punctures. Distribution and bionomics: The species is repre- sented only by its holotype from Corsica; the locality 1s not specified. y 3.15. Medon brunneus (Erichson, 1839) (Map 8) Lithocharis brunnea Erichson, 1839 (ERICHSON 1839: 5131. Lathrobium megacephalum Heer, 1839 (HEER 1839: 238). Lithocharis monticola Hampe, 1867 (HAMPE 1867: 372). Medon robustior Roubal, 1920 (ROUBAL 1920: 150). Medon olympicus Scheerpeltz, 1963b (SCHEERPELTZ 1963b: 439ff). Types examined: See ASSING (2004a). Additional material examined (total: 1300 exs.; for additional material see ASSING (2004a)): Sweden: Skane: 2 exs., Skäralid, 4.V.1973 € 6.X11.1975, leg. Gillerfors (cGil); 3 exs., same locality 6.V.1977, leg. Gillerfors (cGil); 2 exs., same locality, 17.1X.1996, leg. Gillerfors (cGil); 1 ex., same locality, 31.V.1997, leg. Gillerfors (cGil); 3 exs., Hall. Väderö, 2.X1.1974, leg. Gillerfors (cGil); 1 ex., Hall. Väderö, 31.V111.1985, leg. Gillerfors (cGil). Spain: Cataluña: | ex., Gerona, Ripoll, 11.11.1978 (cAss); 3 exs., Ripoll — Vic, leg. Franz (NHMW); 1 ex., Gerona, 7 km S Vidreras, 41°43N, 02°50E, 150 m, oak forest, 10.X.1997, leg. Zerche (cAss); 5 exs., 40 km N Barcelona, Sierra de Montseny, S-slope, 700 m, 19.111.1994, leg. Assing (cAss); 1 ex., Sierra de Mon- tseny, 1400 m, beech forest, 19.111.1994, leg. Wunderle (cWun); 1 ex., Sierra de Montseny (cBor); 1 ex., Sierra de Montseny, 1100 m, 20.1V.1999, leg. Tronquet (cTro); | ex., same data, but 1800 m (cTro). England: 1 ex., Kent, Maidstone, 23.1V.1990, leg. Owen (cOwe). France: Languedoc-Roussillon: 8 exs., N Nimes, Pont du Gard, leg. Franz (NHMW, cAss); 5 exs., Gard, W Nimes, Dions, 19.V.1994, leg. Schiilke (cSch); 1 ex., Hérault, St. Jean de Bueges, 17.V1.2000, leg. Tronquet (cTro); 12 exs., Prats de Mollo, leg. Franz, Giraud (MHNG, NHMW, cAss); 1 ex., same locality, 26.V.1937, leg. Gi- raud (MHNG); 1 ex., Pyrénées Orientales, Argeles, Val- lé de la Massane, leg. Franz (cAss); 1 ex., Molitg les Bains, 600 m, 13.V1.1995, leg.Tronquet (cTro); 1 ex., same data, but 28.V.1995 (cTro); 1 ex., Molitg les Bains, 700 m, 7.111.1999, leg.Tronquet (cTro); 1 ex., 60 Bonner zoologische Beitráge 54 (2005) Campóme, 700 m, 16.X1.1996, leg. Tronquet (cTro); 1 ex., Campöme, 800 m, 24.V1.1995, leg. Tronquet (cTro); 2 exs., forét de Boucheville near Rabouillet [42°45N, 2°21E], 750-650 m, 27.11.2001, leg. Tronquet (cTro). Aquitaine: 1 ex., Dordogne, Brantöme (MHNG); 1 ex., Dordogne, Le Bugue, 3.V111.1941 (MHNG); 1 ex., Gironde, Sadirac, 22.1V.1939, leg. ~Tempere (MHNG); 3 exs., Léognan, 26.1V.1936, leg. Tempere (MHNG). Bretagne: 3 exs., Forét de Huel- goat, leg. Franz (NHMW, cAss). Normandie: | ex., Caen (NHMW); 6 exs., Rouen (NHMW). Íle-de- France: | ex., Paris [paralectotype of Lithocharis picea Kraatz]. Bourgogne: | ex., Nievre, Brassy, leg. Mequi- enon (MHNG); 2 exs., Nievre, Dun-les-Places, V11.1903, leg. Mequignon (MHNG). Champagne- Ardenne: 3 exs., Epernay (MHNG); | ex., Marne, Ft. de Troisfontaine (MHNG). Centre: 3 exs., Perrusson, leg. Mequignon (MHNG). Provence: 3 exs., Vaucluse, Plateau de Vaucluse, Gorge de Nesque, 500 m, 28.X11.1995, leg. Assing & Stüben (cAss); 3 exs., Vau- cluse, Mt. Ventoux near Mt. Serein, 1000 m, 27.X11.1995, leg. Assing & Stüben (cAss); 1 ex., Vau- cluse, Montagne du Luberon, Apt, S Auribean, 700 - 800 m, 29.X11.1995, leg. Assing & Stüben (cAss); 2 exs., Carpentras, Vénasque, 300 m, 5.VIII.1997, leg. Stüben (cAss); 4 exs., Tanneron, 350 m, 26.V.1991, leg. Wunderle (cWun); | ex., Tanneron, VI.1988, leg. Wun- derle (cWun); 4 exs., Le Beausset, 30.V11.1921 (MHNG); | ex., Port-Cros, IV.1962 (MHNG); | ex., Ol- lioules, X1.1960 (MHNG); 1 ex., Lantosque (cBor); 3 exs., Sophia Antipolis near Antibes, haystack, 16.X.1991, leg. Schiilke (cSch); 9 exs., Col de Castil- lion, N Monti, 350 m, 5.V.1996, leg. Wolf (cSch); 2 exs., road from Col de Castillion to Col de Braus, 700 m, 8.V.1996, leg. Wolf (cSch); 5 exs., Col de Braus, 1000 - 1150 m, 7.V.1996, leg. Wolf (cSch); 1 ex., SE Sospel, Albarée, 860 m, 9.V.1996, leg. Wolf (cSch); 10 exs., Menton, Annonciade, 13.X1.1983, leg. Zola (cZan); 4 exs., Mandelieu, 6.X11.1992, leg. Zoia (cZan); I ex., St.-Vallier-de-Thiey, leg. Toumayeff (MHNG); 2 exs., same data, but VI.1968 (MHNG); 2 exs., same da- ta, but VIII.1969 (MHNG); 1 ex., same data, but VIII.1976 (MHNG); 2 exs., same locality, leg. Sainte Claire Deville (MHNG); 1 ex., Nice, La Lanterne, X1.1956 (MHNG); 2 exs., Nice, 25.VIII.1921 (MHNG); 1 ex., Nice, leg. Bedel (MHNG); 2 exs., St. Augustin, 111.1958 (MHNG); | ex., St. Augustin, leg. Ochs (MHNG); 1 ex., Beaulieu-sur-Mer, 1.1V.1932 (MHNG); 1 ex., Theoule (MHNG). Rhóne-Alpes: | ex., Bourg-en-Bresse, 450 m, 10.VIII.1997, leg. Stüben (cAss); 1 ex., Voiron, 17.V11.1972 (cBor); 1 ex., Ce- vennes, Col de l’Escrinet, 790 m, 8.-9.V1.1999, leg. Wolf (eSch); 1 ex., W Tullins, road from Venrey to Grenoble, 300 m, 26.V1.1999, leg. Wolf (cSch); 1 ex., Gorges de l’Ardeche, near Grotte de la Madeleine, 10.V1.1999, leg. Wolf (cSch); 1 ex., Saint Jean d’Conz, 15 km W Chambery, C. la Bruyere, 650 m, 26.V1.1999, leg. Wolf (cSch); 1 ex., Ain, Cerdon, 1.1948 (MHNG); 3 exs., Ain, La London [46°10N, 06°01E], 12.VII.1961, 7.1X.1964, leg. Comellini (MHNG); 3 exs., Ain, Longe- ray [46°06N, 05°53E], 15.X11.1961, leg. Comellini (MHNG); 7 exs., Savoie, Pont-du-Fier, 5.X.1960, leg. Comellini (MHNG); 1 ex., Savoie, forét de St. Hugon, V11.19124 (MHNG); 1 ex., Savoie, St.-Alban, 15.1X.1960, leg. Comellini (MHNG); 1 ex., Haute-Savoie, Iles d’Arve [46°12N, 06°08E], 30.V.1959, leg. Comellini (MHNG); 2 exs., Haute-Savoie, Allinges, VII.1960, leg. Comellini (MHNG); 1 ex., Haute-Savoie, Dingy-en- Vuache, 12.11.1961, leg. Comellini (MHNG); 2 exs., Haute-Savoie, Vulbens, 9.V.1960 & 9.V.1961, leg. Co- mellini (MHNG); 3 exs., Haute-Savoie, Rumilly, 10.X.1959, leg. Comellini (MHNG); 1 ex., Haute- Savoie, Arcine, 1.V.1961, leg. Comellini (MHNG); 1 ex., Haute-Savoie, Chamonix, La Fégére (MHNG); 1 ex., Rhone, Vaugneray (MHNG); 3 exs., Dróme, Va- lence (MHNG). Locality illegible: 4 exs. (MHNG). Monaco: 3 exs., Monaco (MHNG). Switzerland: | ex., Bern, Gúmmenen, 19.1V.1986, leg. Feller (cSch); 1 ex., Bern, Gallmitz, 7.1V.1986, leg. Fel- ler (cSch); 4 ex., Vaud, Morges (NHMW). Belgium: | ex., Spa, 6.1V.1983, leg. Assing (cAss). Germany: Nordrhein-Westfalen: 8 exs., Porta West- falica, Wittekindsberg, oak forest, 10.V1.1993, leg. As- sing (cAss); 1 ex., Höxter, Beverungen, 17.V.1990, (cAss); 3 exs., Menden, Hönnetal, Klusenstein, X1.1994 & 29.1V.1996, leg. Feldmann (cFel); 1 ex., Rosbach, leg. Katschak (cFel); 1 ex., Nachrodt, 200 m, XII.1997, leg. Feldmann (cFel); 3 exs., Brühl near Köln, Staats- forst Ville, 2.VIII.1989, leg. Köhler (cKöh, cWun); 1 ex., same data, but 23.VII.1988 (cKöh); 1 ex., Rade- vormwald, 29.X.1985, leg. Wenzel (cKöh). Rheinland- Pfalz: 1 ex., Lahnstein, NSG “Koppelstein”, 18.VI11.1985, leg. Wunderle (cWun); 1 ex., Altenahr, Vischeltal, 17.VII1.1985, leg. Wunderle (cWun); 1 ex., Altenahr, vineyard, 2.V1.1984, leg. Wunderle (cWun); 2 exs., Al- tenahr, Langfigtal, 11.1V.1987, leg. Köhler, Wunderle (cKöh, cWun); 1 ex., Altenahr, Reimertzhoven, 1.1V.1987, leg. Köhler (cKöh); 1 ex., Eifel, Gemünd, Kermeter, Am Steinbach, 23.V1.1992, el. Köhler (cKóh); 3 exs., 3 km S Ahrweiler, 18.1X.1980, leg. Scheuern (cWun). Baden-Württemberg: | ex., Wyh- len, 20.V111.1978, leg. Kiener (MHNG). Schleswig- Holstein: 3 exs., Ratzeburg, 3.1V.1988, leg. Wunderle (cWun). Niedersachsen: 2 exs., Ith, Coppenbrügge, 29.IX.1991, leg. Assing (cAss); 1 ex., same data, 14.X.1984 (cAss); 2 exs., Ith, Lauenstein, 300 m, mixed forest, pitfall trap, 28.VII., 2.IX. & 6.X11.1990 (cAss); 1 ex., same locality, 6.VIII.1988 (cAss). Hessen: 2 exs., Marburg, Caldern, 5.V.1984, leg. Wunderle (cWun); l ex., Rasdorf (MHNG); 1 ex., Darmstadt (NHMW). Volker ASSING: Western Palaearctic Medon 61 Bayern: | ex., Bad Feilnbach, IV.1999, leg. Feldmann (cFel); 4 exs., Chiemgau, Kampenwand, 1000 m, 4.VIII.1995, leg. Wolf (cSch); 2 exs., Königssee, 23.1X.1949, leg. Benick (MHNG). Mecklenburg- Vorpommern: | ex., Rügen, Lohme, 26.V11.1984, leg. Behne (cSch); 1 ex., S Lohme, 1.X.1989, leg. Zerche (DEI). Sachsen-Anhalt: | ex., LKr. Sangerhausen, Questenberg, 26.V1.1998, leg. Schülke (cSch). Bran- denburg/Berlin: 3 exs., Pusack, deciduous forest, 25.1X.1999, leg. Wrase (cSch); 16 exs., Eberswalde, coll. Neresheimer (DEI). Sachsen: 3 exs., Meissen, Za- del, 3.1V.1976 (cSch); 2 exs., Dresden, NSG Rotstein near Sohland, 8.X.1988, leg. Zerche (DEI); 2 exs., Dresden, Zittauer Geb., NSG Lausche, 7.X.1988, leg. Zerche (DEI). Locality not specified: 1 ex. (MHNG). Italy: Trentino-Alto Adige: | ex., Wals near Kaltern, 25.V11.1973 (cAss); 1 ex., Pieve di Ledro, 30.V111.1972, leg. Pace (cBor); 2 exs., Bressanone (BZ), Tiles, 800- 1000m, 30.VII.1973, leg. Sette (cZan); 4 exs., Tiles, VIII.1975, leg. Sette (cZan); 1 ex., Smarano (TN); VII.1972, leg. Zanetti (cZan); 1 ex., same locality, 1000m, 11.V11.1973 (cZan); 2 exs., same data, but 11.VII.1974 (cZan); 2 exs., same data, 21.VIII.1996 (cZan); 1 ex., Trento (NHMW). Piemonte: | ex., Abba- zia, 23.X11.1934, leg. Springer (UCBA). Lombardia: 6 exs., Alpi Orobi, Morbegno, 350 m, deciduous forest, 10.VI11.1994, leg. Assing (cAss); 4 exs., Val Formazza, N Domodossola, 1050 m, 1.VI11.1996, leg. Wolf (cSch); 1 ex., Mantova, Bosco Fontana, 27.X.1980, leg. Zanetti (cZan). Veneto: 1 ex., Fongara, 21.V.1972, leg. Pace (cBor); 1 ex., Valli del Pasubio, 9.V.1972, leg. Pace (cBor); 1 ex., San Giovanni Ilarione, 7.V.1972, leg. Pa- ce (cBor); 1 ex., Belluno, 23.1V.1973, leg. Garagnani (cBor); 2 exs., Feltre (BL), 1.V.1973 (cZan); 7 exs., Pieve d'Alpago (BL), 22.VII.1961 (UCBA, cZan); 1 ex., Valdagno (VI), 21.V.1972, leg. Zanetti (cZan). Friuli-Venezia-Giulia: | ex., Tagliamento near Comi- no, 46°13N, 13°01E, 50 m, 12.1X.1998, leg. Schiilke (cSch); 2 exs., E Cividale del Friuli, Podresca, 750m, 30.V.1986, leg. Torti (cZan); 1 ex., E Trieste, Grópada, 3.1X.1980, leg. Seriani (Zan); 3 exs., Aurisina (TS), ZONENOS6; leg. Torti (cZan); 29 exs., - Trieste env. (NHMW); 14 exs., Trieste, Opacina-Basovizza, V.- V1.1921, leg. Moczarski & Scheerpeltz (NHMW). Emi- lia-Romana: | ex., NW Imola, Mte. Calderano, 570 m, 9.V.1997, leg. Wolf (cSch). Toscana: 8 exs., 30 km SW Firenze, Figline Valdarno, 200 m, 14.V1.1992, leg. As- sing (cAss); 1 ex., Monticiano, R. N. Alto Merse, 8.11.1999, leg. Meybohm (cAss); | ex., Mt. di Calvanı (FI), Mungona, 650 m, 1.V.1991, leg. Wunderle (cWun); 3 exs., Siena Vecchia (SI), 300 m, 9.V.1991, leg. Wunderle (cWun); 1 ex., Mt. Falterona, Castagno di Andrea, 700 m, 7.V.1991, leg. Wunderle (cAss); 2 exs., Ruota, X1.1971, leg. Bordoni (cBor); 1 ex., N Borgo San Lorenzo, Grezzano, X.1971, leg. Bordoni (cBor); 2 exs., S Massa, Lago dell’Accesa, 3.X.1995, leg. Bordo- ni (cBor): 4 exs., P.so Viamaggio (AR), 830m, oak fo- rest, 2.V.1986, leg. Zoia (cZan); 13 exs., Radicondoli (SD, 3.1V.1994 (cZan); 3 exs., Petriolo, 3.X1.1972, leg. Castellini (cTro); 5 exs., Cantagallo, IX.1976, leg. Ca- stellini (cTro). Umbria: 2 exs., Lago di Trasimeño, Monte Solare, 400 m, 24.V.1998, leg. Wolf (cSch); 3 exs., Castiglione del Lago (PG), Pozzuolo, oak forest, 2.V.1986, leg. Zoia (cZan). Abruzzi: 15 exs., Torino di Sangro, 24.-26.V11.1974, leg. Pace (cBor). Campania: 2 exs., Napoli, Ischia, S. Angelo, 27.X.1936, leg. Be- nick (cAss); 5 exs., Napoli, Caserta, X.1924, leg. An- dreini (cBor); 1 ex., Novi Velia, 7.V111.1973, leg. Pace (cBor); 1 ex., Monti Albumi, Salerno, W San Rufo, Pso. di Sentinella, 900 m, 10.X.2000, leg. Wolf (cSch); | ex., Salerno, E Paestum, Capaccio, Mte. Soprano, 350 m, 13.X.2000, leg. Wolf (cSch); 2 exs., NE Salerno, Monte Stella, leg. Liebmann (MHNG). Lazio: 2 exs., Monte Circeo, N-Slope, 300 - 450 m, Quercus ilex forest, 27.X11.1994, leg. Assing (cAss); 2 ex., Monti Lepini ne- ar Carpineto, 27.V1.1973, leg. Pace (cBor); 8 exs. Monti Aurunci NE Formia, 3.V11.1975, leg. Pace (cBor); | ex., Lago di Vico (UT), 10.X1.1974, leg. Rossi (cZan); | ex., Lago di Bracciano, Monti Sabatini, Mte. Guerrano, 500 m, 7.V.1998, leg. Wolf (cSch); 1 ex., Lago di Bracciani, 3 km N Orido Romano, 5.V.1998, leg. Wolf (cSch); 1 ex., 19 km S Rieti, Cerdomare, 19.V.1998, leg. Wolf (cSch); 2 exs., Itri (LT), 320m, 30.1X.1982, leg. Zoia (cZan); 5 exs., Colli Albani, Monte Cavo, 7.X1.1980, leg. Zoia (cZan); 3 exs., Colli Albani, Rocca di Papa, Monte Cavo, 600 - 800 m, 9.V.1998, leg. Wolf (cSch). Puglia: 2 exs., Gargano, W San Marco in Lámis, Mon- tenero, 900 m, oak forest, 30.X11.1994, leg. Assing (cAss); 4 exs., Gargano, 16.V111.1974, leg. Pace (cBor); 1 ex., Foresta Umbra, Foggia, VII.1972, leg. Sciaky (UCBA); 1 ex., Foresta Umbra, 22.V1.1967, leg. Lohse (MHNG); 3 exs., B. Umbra, 26.V.1913, leg. Fiori (UCBA). Basilicata: 1 ex., Maratea, 4001N, 15°44E, 485 m, 12.V.2002, leg. Wunderle (cWun); | ex., 12 km SSW Latrónico, 40°01N, 15°58W, 635 m, 9.V.2002, leg. Wunderle (cWun); 2 exs., Potenza, W San Mauro Forte, Accettura, 600 - 780 m, 22.X.2000, leg. Wolf (cSch). Calabria: | ex., Orsomarso (CS), 1.VII.1986, leg. Angelini (cZan); 4 exs., Orsomarso, Grisolia (CS), 700 m, 17.V1.1997, leg. Angelini (cZan). Austria: Vorarlberg: 3 exs., Gebhardsberg (NHMW). Oberósterreich: 1 ex., Windischgarsten, leg. Franz (NHMW); | ex., Gmunden, leg. Franz (NHMW); | ex., Weyr, leg. Franz (NHMW); 1 ex., Kleinreifling, leg. Franz (NHMW). Tirol: 7 exs., Hall, 1200 m, 7.VUI.1997, leg. Wolf (cSch). Salzburg: | ex., Mand- ling, leg. Franz (NHMW). Niederósterreich/Wien: 2 exs., Frankenfels, leg. Franz (NHMW); 1 ex., Hardegg, leg. Franz (NHMW); 2 exs., Haslau, leg. Scheerpeltz (NHMW); 26 exs., Mödling, leg. Franz, Mandl, Scheer- 62 Bonner zoologische Beitráge 54 (2005) peltz (NHMW); 1 ex., Tullnerbach, leg. Scheerpeltz (NHMW); 2 exs., Vöslau, leg. Franz (NHMW); 1 ex., Krenngraben near Kl. Hollenstein, leg. Franz (NHMW); 1 ex., Donauauen, Wolfsthal, leg. Franz (NHMW); 1 ex., Ebreichsdorf, mole nest, leg. Franz (NHMW); 3 exs., Wien (NHMW); 2 exs., Hütteldorf (NHMW); 1 ex., Neuwaldegg (NHMW); 2 exs., Wien, Weidlingau (NHMW); 2 exs., Rodaun (NHMW); | ex., Wien, Prater, leg. Franz (NHMW); 1 ex., Wienerwald, Hadersdorf, leg. Skalitzky (NHMW); 12 exs., Wienerwald, Reka- winkel, leg. Scheerpeltz, Skalitzky (NHMW); 2 exs., Wien, Bisamberg, leg. Skalitzky, Scheerpeltz (NHMW ); 4 exs., Pressbaum, leg. Scheerepeltz (NHMW); 1 ex., Lunz (MHNG); 3 exs., Scheibbs, 24.V11.1970, leg. Res- se (MHNG); 16 exs., 4 km SE Hof am Liethberge, Franz-Joseph-Warte, 47°56N, 16°37E, 480 m, decidu- ous forest, 15.X.1998, leg. Zerche (DEI); 2 exs., Brühl, leg. Schuster (NHMW); 6 exs., Leithagebirge (NHMW; 1 ex., Schneeberg (NHMW); 2 ex., Rabenstein (NHMW); Tullnerbach, leg. Scheerpeltz (NHMW). Steiermark: 7 exs., Mühlbachgraben near Rein, leg. Franz (NHMW); l ex., Pass im Stein, leg. Franz (NHMW); 2 exs., Peg- gau, leg. Franz (NHMW); | ex., Frauenstein, leg. Franz (NHMW); 1 ex., Tüffer, leg. Franz (NHMW); I ex., Ringkogel, Hartberg, leg. Franz (NHMW); | ex., Hoch- schwab, Bodenbauer, leg. Wimmer (NHMW); I ex., Tobelbad (MHNG); 13 exs., Pleschkogel near Rein, 470 m, 20.V1.1999, leg. Zerche (DEI). Kärnten: | ex., Flat- tach, Mölltal, leg. Franz (NHMW); I ex., Saualpe, E- berstein, 17.V11.1986, leg. Wunderle (cWun); 1 ex., Karnische Alpen, Weidenburger Wasserfall near Wei- denburg, 17.-20.1X.1987, leg. Wunderle (cWun); I ex., Maria Rain (NHMW); 1 ex., Waidisch, leg. Mandl (NHMW); 1 ex., Dobratsch (NHMW); 1 ex., Gailtaler Alpen, Presseggen (NHMW). Burgenland: 1 ex., Rechnitz, leg. Franz (NHMW); 2 exs., Geschrieben- stein, leg. Franz (NHMW); 4 exs., Eisenberg near Burg, leg. Franz (NHMW); 2 exs., Leithagebirge, Zeilerberg, 24.V111.1983, leg. Assing (cAss); 17 exs., Leithagebir- ge, Purbach, leg. Franz (NHMW); 5 exs., Leithagebirge, Wimpassing, leg. Franz (NHMW); | ex., Leithagebirge, Winden, leg. Franz (NHMW); 1 ex., Donnerskirchen, leg. Franz (NHMW); 2 exs., Ruster Hügelzug, leg. Franz (NHMW); 1 ex., Rust, pasture, 11.V11.1973 (cAss); 43 exs., Zurndorf, leg. Franz (NHMW); I ex., Nickelsdorf, leg. Franz (NHMW); I ex., Neusiedlersee, eastern lakeshore, leg. Franz (NHMW); I ex., Groß- Petersdorf, leg. Franz (NHMW); 1 ex., Leithagebirge (MHNG); 20 exs., Neudorf (NHMW). Locality ambi- guous: 4 exs., Gaming (MHNG). Slovakia: | ex., Velka Fatra, Blatnica, 900 m, 11.V11.1992, leg. Assing (cAss); 1 ex., Novy Salas, Presev, 8.V.1998, leg. Smatana (cSch); 3 exs., Pezinok, 400m, 14.V111.1992, leg. Zoia (cZan); 2 exs., Hrhov, 3.V.1967, leg. Lohse (MHNG); 2 exs., Mala Fatra, Rieka W Kralovany, Cha- ta Sútovska dolina, 40°10N, 19°05E, 500 m, 24.V11.2001, leg. Zerche (DEI); 4 exs., E Bánovce, E Ostry Vrach peak, 48°44N, 18°22E, 520 m, beech fo- rest, 3.V111.2001, leg. Zerche (DEI); 3 exs., Slovensky kras, W Borka, 48°38N, 20°47E, 655 m, beech forest, 29.V11.2001, leg. Zerche (DEI); 2 exs., Pieninsky naro- den park, 18.V11.1993, leg. Zerche (DEI); 1 ex., Brati- slava, Svaty jur, 220 m, 26.V.1991, leg. Behne (DEI); 2 exs., Pieniny Geb., Lesnica 500 m, 11.-14.V1.1990, leg. Behne (DEI); 3 exs., Nitra (NHMW); 3 exs., Herlany, 15.V.2000, leg. Hlavaé (cAss). Slovakian or Polish territory: 2 ex., “Beskiden” (NHMW). Czech Republic: 5 exs., Prag env. (NHMW); 2 ex., “Erzgebirge, Hamergrund” (NHMW); 2 exs., “Hof” (NHMW); 1 ex., Oskau (NHMW). Slovenia: 6 exs., Celje, leg. Franz (NHMW); I ex., Li- pica, 27.V1.1981 (cAss); 1 ex., Mozelj Kocevje, 500 m, mixed forest, 11.V.1993, leg. Gasparo (cZan); 7 exs., Senozece, 7.V111.1969, leg. Besuchet (MHNG); 2 exs., Podcetrtek, 26.1V.1930, leg. Kodrik (MHNG); 2 exs. Postojna (NHMW); 5 exs., “Wochein” (NHMW). Hungaria: 2 ex., Sopron (NHMW); 3 exs., Budapest (NHMW; 8 exs., locality not specified or not identified (NHMW). Romania: | ex., Brasov (MHNG); 4 exs., Bihar (NHMW); 8 exs., Baile Herculane (NHMW); 2 exs., “Roterturm Pass” (NHMW); | ex., Sinaia (NHMW); 1 ex., Azuga (NHMW); 2 exs, Tismana (NHMW); 1 ex., Campu Selului (NHMW); | ex., Gaia Arama (NHMW) Bulgaria: 4 exs., Pirin, leg. Weirather (MHNG). Croatia: | ex., Ridnik-Karlovac, 30.1V.1990, leg. Wunderle (cWun); 1 ex., Istria, Opatija, Mt. Ucka, 250 m, 24.V1.1996, leg. Wolf (cSch); 5 exs., same data, but 950 m (cSch); 5 exs., Pula, 13.1V.1974, leg. Zoia (U- CBA); 6 exs., Pula (NHMW); 1 ex., Plitvice, 18.V111.1977, leg. Lohse (MHNG); 1 ex., Plitvice (NHMW); 19 exs., Opatija (NHMW); 8 exs., Noghera (NHMW ); 34 exs., Rab island (NHMW); 9 exs., Mljet island (NHMW ); 16 exs., Velebit (NHMW ); 3 exs., Ra- dostak (NHMW). Bosnia-Herzegovina: | ex., Prozor, 16.V111.1977, leg. Lohse (MHNG); 4 exs., Bjelasnica, 15.V111.1977, leg. Lohse (MHNG); 5 exs., Bjelasnica (MHNG); 5 exs., Nevesinje (NHMW); 4 exs., Jablanica (NHMW ); 2 exs., Metkovic (NHMW); 3 exs., Majevica planina (NHMW); 5 exs., Travnik, 30.V.1934, leg. Natterer (NHMW): 3 ex., Celié (NHMW); 4 exs., Zepce (NHMW); 5 exs., Sarajevo (NHMW). Serbia-Montenegro: | ex., Lovcen (NHMW); 6 exs., Bojana range (NHMW); 18 exs., Rtanj Planina, leg. Volker ASSING: Western Palaearctic Medon 63 Breit (NHMW); 2 exs., Visocka Rzana, Pirot, 750 m, 29.1V.2002, leg. Hlavac (cAss). Macedonia: | ex., KoSevo, leg. Pfeiffer (MHNG); 5 exs., northern shore of lake Prespa, 23.1X.1958, leg. Schuster (NHMW); 3 exs., Monastir (NHMW). Albania: 5 exs., Krujé, leg. Mader (NHMW); 2 exs., Sarisaltik, 1180m, leg. Mader (NHMW). Greece: 6 exs., Pindos, Kalivia, 28.-29.V.1989, leg. Zoia (cZan); 4 exs., Evvoia, Pagóndas, 4.1V.1983, leg. Zoia (cZan); 2 exs., Evvoia, Dirfys, 1.111.1983, leg. Zoia (cZan); 1 ex., Pelopónnisos, Olympia, O. Alfios, 7.1X.1995, leg. Zoia (cZan); 38 exs., Pilion (NHMW); 22 exs., Parnassos Oros (NHMW); 6 exs., Athina (NHMW); 1 ex., Sfendali (NHMW); 12 exs., Nisista, Xeravunel (NHMW); 2 exs., Peloponnisos, Patras, 22.111.1971, leg. Löbl (MHNG); 4 exs., Pelopónnisos, Kalavrita, 800 m, 3.1V.1971, leg. Löbl (MHNG); 1 ex., Pelopönnisos, Enochorion (NHMW); 1 ex., Pelopönni- sos, Olympia, Ladzoi, 100 m, 22.111.1992, leg. Frisch (MNHUB); 1 ex., Pelopómnisos, Taygetos, Kalamata- Sparta, 1300 m, 1.1V.1992, leg. Frisch (MNHUB):.1 ex., Taygetos (NHMW): 6 exs., Drama, Falakró, leg. Weirather (MHNG, cBor): 1 ex., Zakynthos, Skopos, 24.111.1971, leg. Lóbl (cBor); 6 exs., Kefallinia, Sami, 31.11.&2.1V.1971, leg. Löbl (MHNG); 1 ex., Kefallinia, Oros Aenos, 31.111.1971, leg. Mahnert (MHNG); 13 exs., Kefallinia, several collectors (NHMW); 12 exs., Lefkas, Moni near Fryni, 27.1111971, leg. Hauser (MHNG); 44 exs., Lefkas, several collectors (NHMW ); 1 ex., Zante, 1809, leg. Hilf (NHMW); 50 exs., Corfu, leg. Beier, Hummler, Moczarski (MHNG, NHMW); 2 ex., Crete, Amari, leg. Biró (NHMW). Ukraine: 1 ex., Crimea, Evpatoria, 10.-20.VII.1999 (cSch). Turkey: 22 exs., Kirklareli, Demirkóy, 31.VII.1969, leg. Besuchet (MHNG, cAss); 4 exs., Istanbul, “Forét de Belgrade”, 4.V1.1967, leg. Besuchet (MHNG); 1 ex., same data, but 10.VII.1969 (MHNG): 4 exs., Istanbul, Sile, 28.V.1967, leg. Besuchet (MHNG, cAss). Locality not specified: 11 exs. (MHNG). Diagnosis: See ASSING (2004a) and the key in Section 4. Distribution and bionomics: Medon brunneus is a widespread Adriato-Mediterranean element (Map 8). Its distribution ranges from northwestern Turkey, southern Greece (including Crete), and southern Italy in the southeast and south, to southern Sweden and southern England in the north, and to western France and north- eastern Spain in the west (material examined; ASSING 2004a). For additional records see e.g. BOHAC (1985), BRUGE et al. (2001), DRUGMAND (1989), FOWLER (1888), HANSEN (1996), HANSEN et al. (1995), HANSEN (1964), HORION (1965), HUGENTHOBLER (1966), KOCH (1968), KOHLER & KLAUSNITZER (1998), LINKE (1907, 1913), LUNDBERG (1995), SCHILLER (1989), TERLUTTER (1995), and TRONQUET (2001). In Scandinavia, the spe- cies is present only in Denmark and southwestern Swe- den: Skane, Halland, Vástergótland, Bohuslán (HANSEN 1964; LUNDBERG 1995). It is unknown from Norway, Finland, and the Baltic countries (SILFVERBERG 1992). HORION (1965) reports the species from Poland (two lo- calities), Kiev (Ukraine), and from Russia (Wolyns'ka Oblast). The records from the Caucasus region must be considered doubtful until they are confirmed (ASSING 2004a). The species is very common in the southeast of its range: in the Balkans, Italy, and the southeast of Cen- tral Europe; it is much rarer in the northwest. In Italy, it is absent from Sicily and probably also from Sardinia. The records from the latter island (LUIGIONI 1929) are apparently based on misidentifications; | have seen no specimens from there. The previous record from the is- land of Pantelleria (CICERONI & ZANETTI 1995) refers to M. despectus (see Section 3.18). OUTERELO (1980) re- ports M. brunneus from Andalucia, stating that the spe- cies occurs in the whole Iberian Peninsula. These re- cords are most likely to be based on misidentifications (confusion with M. cauchoisi). The occurrence of M. brunneus in the Iberian Peninsula is confined to the very northeast of Spain. DRUGMAND (1989) states that the species is bivoltine, but there is no evidence supporting this. Based on the material examined here, M. brunneus is univoltine. Ac- cording to HORION (1965), the species may be associ- ated with nests of voles and/or mice, but there is still no striking evidence proving this. Previous records from mole nests are probably accidental; NOWOSAD (1990) found the species in only 7 nests out of 5000 nests stud- ied. It is usually — and often in large numbers — found in the leaf litter of forests, especially deciduous forests (oak). 3.16. Medon piceus (Kraatz, 1858) (Figs. 74-78, Map 9) Lithocharis picea Kraatz, 1858 (KRAATZ 1858: cxci f). Medon muscicola Mulsant & Rey, 1878 (MULSANT & REY 1878: 128). Medon murciensis Coiffait, 1970a (COIFFAIT 1970a: 715f) syn. n. 7 Types examined: L. picea: Lectotype ¢ [remounted, aedeagus dissected], here designated: Paris / Coll. Kraatz / Syntypus / Coll. DEI Eberswalde / Medon piceus (Kr.) V.l.Gusarov det. 1996 / Lectotypus ¢@ Lithocharis picea Kraatz desig. V. Assing 2003 / Medon piceus (Kraatz) det. V. Assing 2003 (DEI). Paralecto- types: 14, 19: same data as lectotype (DEI); 14: same data, but “Medon brunneus (Erichson) det. V. Assing 2003” (DED. 64 Bonner zoologische Beiträge 54 (2005) Map 8: Distribution of Medon brunneus (Erichson) (filled circles: revised records; open circles: selected literature records). M. murciensis: Holotype ¢: Sierra de Espuña bei Mur- cia, Hi. m., lg. H. Franz / MUSEUM PARIS COLL H. COIFFAIT / HOLOTYPE / Medon burdigalensis Coiff. H. COIFFAIT det. 1969 / Medon piceus (Kraatz) det. V. Assing 2003 (MNHNP). Comments: One of the syntypes of Lithocharis picea is conspecific with M. brunneus (Erichson), the others are in agreement with the long-standing interpretation of the species. Since the type series is a mixture of two spe- cies, a lectotype designation is mandatory, in order to unambiguously define the name; the male in better con- dition was selected as the lectotype. In the original description of M. murciensis, which is based on a single male, COIFFAIT (1970a) compares the spe- cies with M. ripicola (Kraatz). An examination of the holotype revealed, however, that it is conspecific with M. piceus (Kraatz), so that the junior name M. murcien- sis is here placed in the synonymy of that species. Additional material examined (total: 677 exs.): Portugal: 2 exs., Serra do Geres, Portela de Homem, 41°48N, 08°08W, 750 m, oak forest, 23.111.2002, leg. Lompe (cAss); 5 exs., Serra do Geres, Puerto la d’Home, leg. Franz (NHMW); 2 exs., Serra do Geres, 800 m, oak forest, 16.V.1992, leg. Wunderle (cWun); | ex., Serra de Montezinho, Rio Sabor, 41°54N, 06°47W, 900 m, poplar litter, 21.111.2002, leg. Meybohm (cAss); 1 ex., Serra de Estrela, S Manteigas, 40°22N, 07°33W, 980 m, oak forest, 18.11.2002, leg. Lompe (cAss); 2 exs., Serra da Estrela, S Manteigas, 40°21N, 07°34W, 1070 m, 19.111.2002, leg. Lompe (cAss); 1 ex., Serra da Estrela, N Manteigas, 40°24N, 07°32W, 850 m, 19.11.2002, leg. Meybohm (cAss); 8 exs., Manteigas, Volker ASSING: Western Palaearctic Medon 65 1000 - 1100 m, leg. Franz (NHMW); 10 exs., Serra da Estrela, leg. Franz (NHMW); 1 ex., Guarda, S Gouveia, 1100 m, 17.IV.1960, leg. Besuchet (cAss); 4 exs., Coimbra, leg. Paulino (NHMW); 1 ex., Coimbra (NHMW); 2 exs., Lisboa, Serra do Sintra, 24.V.1992, leg. Wunderle (cAss); 10 exs., Sintra, Lagoa Azul, 6.V1.1966, leg. Besuchet, Comellini (MHNG); 2 exs., Luso, V.1959, leg. Fagel (IRSNB); 41 exs., Algarve, 10 km N S. Bras de Alportel, 400 m, 4.V1.1992, leg. Wun- derle (cWun, cAss, cSch); 22 exs., Algarve, Caldas de Monchique, V.-V1.1960, leg. Fagel (IRSNB); 1 ex., Al- garve, Serra Monchique, E Monchique, 37°19N, 8°31 W, 530 m, 10.1V.2002, leg. Meybohm (cAss). Spain: País Vasco: 2 exs., Sierra de Cantabria, ca. 30 km NW Logroño, NE Pipaón, 42°37’45N, 02°36 18W, 770m, oak forest with dense undergrowth, 14.X.2003, leg. Assing (cAss); 14 exs., Guizpúzcoa, pass between Elgoibar and Azcoitia, leg. Franz (NHMW). Cantá- bria: 2 exs., Santander, Monte de Candino, Liendo, leg. Franz (NHMW); 10 exs., Santander, Monte Aa, Ruente, leg. Franz (NHMW); 4 exs., Santander, Jesus de Mon- tes, W Beranga, leg. Franz (NHMW); 2 exs., Santander, Marron, leg. Franz (NHMW); 1 ex., Santander, Puente Viesgo, leg. Franz (NHMW); 1 ex., Santander, Villa- verde de Pontones, leg. Franz (NHMW); 5 exs., Pechon, Unquera, leg. Franz (NHMW); 1 ex., Reinosa, road to Alto de Campoo, 1400 m, 4.V1.1991, leg. Wunderle (cWun); 1 ex., Alto Campóo, 1600 m, oak forest, 4.V1.1991, leg. Zerche (DEI); 7 exs., same data, but 1400 m (DEI); 1 ex., Espinama, Puerto de Aliva, 28.VII.1972 (cBor); 2 exs., Picos de Europa, 10 km NNW Potes, 43°14’12N, 04°39’11W, 1025 m, N-slope, montane pasture, sifted from grass and moss, 17.V11.2003, leg. Assing (cAss). Navarra: 5 exs., Otxon- do, 16.V11.1968, leg. Besuchet, Comellini (MHNG). La Rioja: 13 exs., Logroño, Monasterio de Valvamera, leg. Franz (NHMW). Asturias: 8 exs., Cabo Vidio, leg. Franz (NHMW ); 8 exs., Posada, leg. Franz (NHMW); 2 exs., S Villaviciosa, leg. Franz (NHMW ); 2 exs., Cova- donga, leg. Franz (NHMW); 1 ex., Nueva, leg. Franz (NHMW); 8 exs., Arriondas, El Fito, V1.1965, leg. Fa- gel (IRSNB). Galicia: 3 exs., Lugo, Alto de Acebo- Cecos, 700 m, 31.V11.1992, leg. Tronquet (cTro); 3 exs., Lugo, Cruzul, Becerreá, leg. Franz (NHMW); 6 exs., Lugo, Valle de Lozera, leg. Franz (NHMW); 3 exs., Lu- go, Villasouto, 2.X.2000, leg. Valcárcel (cSch, cAss); 1 ex., Lugo, Serra do Courel, Rio de Rofión, 23.V.2000, leg. Valcárcel (cSch); 6 exs., Lugo, Monforte de Le- mon, Gullade, 5.1X.2000, leg.. Valcárcel (cSch, cAss); 4 exs., Lugo, Sierra de Ancares, 2 km NE Doiras, 42°47N, 06°58W, 11.V1.2000, leg. Wrase (cSch); 4 exs., Sierra de Barbanza, Noia, leg. Franz (NHMW ); 2 exs., Noia, leg. Franz (NHMW); 3 exs., Nola, Outes, leg. Franz (IRSNB, NHMW); 2 exs., La Coruña, San Saturnino, leg. Franz (NHMW ); | ex., La Coruña, Punta de la Estaca, leg. Franz (NHMW); 1 ex., La Coruña, Puenteseco, leg. Franz (NHMW); | ex., Orense, Verin, Montes del Invernadero, Campo de Becerros, leg. Franz (NHMW); 6 exs., Santiago, Bosque de Cernadas, leg. Franz (NHMW); 2 exs., Santiago, Orilla del Rio Tam- bre, leg. Franz (NHMW); 10 exs., Prov. Pontevellra, Pontevedra, leg. Franz (NHMW); 4 exs., Pontevedra, Isla Cies del Norte, leg. Franz (NHMW); | ex., Ponte- vedra, Testeiro, 800 - 900 m, leg. Franz (NHMW); 2 exs., Pontevedra, Cabo Silieiro, Baiona, leg. Franz (NHMW ); 2 exs., Pontevedra, A Lamosa, Sierra de Fon- teiria, leg. Franz (NHMW); 2 exs., Pontevedra, leg. Franz (NHMW y; 1 ex., Prov. Pontevedra, Rio Umia ne- ar Morana, leg. Franz (NHMW). Castilla-León: 3 exs., Sierra de la Demanda, ca. 40 km E Burgos, NE Valma- la, Rábanos, 42°18N, 03°16W, 1190m, oak forest with grass, 12.X.2003, leg. Assing (cAss); 1 ex., Zamora, Lago de Sanabria, leg.Franz (NHMW); 11 exs., Ponfer- rada, leg. Paganetti (DEI, NHMW); 1 ex., Puerto de Be- jar, 800-1400 m, V.-VI.1957, leg. Fagel (IRSNB). E- stremadura: 1 ex., Prov. Badajoz, Puerto de las Marismas, 8.1V.1959, leg. Besuchet (MHNG). Castilla- La Mancha: 11 exs., SW Albacete, Sierra de Alcaraz, 15 km NNE Riópar, 38°32N, 02°25W, 1350 m, 7.1V.2003, leg. Assing (cAss); 9 exs., same data, but 38°35N, 02°21W, 1125 m, 10.1V.2003, leg. Assing (cAss); 1 ex., same data, but 38°34N, 02°20W, 1120 m, 10.1V.2003 (cAss); 1 ex., Albacete, leg. Comellini (MHNG). Murcia: 5 exs., Sierra de Espuña, leg. Franz (NHMW); 2 exs., Sierra de Espuna, 26.111.1959, leg. Besuchet (MHNG). Andalucía: | ex., Sierra de Segura, Villaverde de Guadalimar, leg. Franz (NHMW); 1 ex., Jaén, Sierra de Segura, Embalse del Tranco, 38°10N, 2°48W, 15.V1.2001, leg. Starke (cFel); 2 exs., Jaén, Sierra de Cazorla, leg. Franz (NHMW ); 3 exs., Sierra de Cazorla, Cazorla, source of Gualaquivir, 6.X.1993, leg. Wunderle (cWun); 15 exs., Sierra de Cazorla, 12.1V.1959, leg. Besuchet (MHNG); 6 exs., SE Ronda (MA), Sierra de Palmitera, 900 m, 24.111.1994, leg. As- sing, Wunderle (cAss, cWun); 20 exs., Sierra de Berme- ja (CA), Jubrique, 350 m, 26.111.1994, leg. Assing, Wunderle (cAss, cWun); | ex., Sierra de Bermeja, 1000 m, 26.111.1994, leg. Assing (cAss); 11 exs., Algeciras (CA), Sierra de Luna, 200 - 350 m, 28.111.1994, leg. As- sing, Wunderle (cAss, cWun); 16 exs., Sierra de Luna, 27.V.1966, leg. Besuchet, Comellini (MHNG); 3 exs., Algeciras (CA), Sierra de Fates, 350 m, 28.111.1994, leg. Assing, Wunderle (cAss, cWun); | ex., Sierra Nevada, Pampaneira, 1000 m, 2.V111.1991, leg.Sprick (cAss); 4 exs., Sierra Nevada, Lanjaron, 600 m, 23.111.1994, leg. Assing, Wunderle (cAss, cWun); 15 exs., Lanjaron, 26.1V.-18.V.1961, leg. Fagel (IRSNB); 1 ex., Sierra Nevada, leg. Franz (NHMW); 3 exs., Sierra Nevada, Guejar Sierra, 1200 m, 28.1X.1993, leg. Wunderle (cWun); 1 ex., Granada, Sierra de Baza, 37°20N, 02°54W, 1700 m, 14.1V.2001, leg. Sprick (cAss); 1 ex., 66 Bonner zoologische Beitráge 54 (2005) Sierra de Córdoba, 9.1V.1959, leg. Besuchet (cBor); 1 ex., Algeciras, Sierra del Niño, 180m, 26.111.1987, leg. Zoia (cZan); 2 exs., Granada, Sierra de la Sagra, 1250 m, 21.V.1966, leg. Besuchet (MHNG); 1 ex., Cordoba, Villaviciosa, 29.V1.1969, leg. Comellini (MHNG); 12 exs., Huelva, Zalamea La Real, 6.V11.1969, leg. Comel- lini (MHNG); 4 exs., Cadiz, 10 km W Los Barrios, 3611N, 05°34W, 210 m, sifted under cork trees, 10.11.1999, leg. Zerche (DEI); 1 ex., ca. 25 km N Almu- ñécar, Sierra de Almijara, 36°53N, 03°42W, ca. 1200 m, mixed pine, oak, and corktree forest, 25.X11.2003, leg. Assıng (cAss). England: | ex., Surrey, Leatherhead, Bookham Com- mon, 24.V.1999, leg. Owen (cOwe). France: Picardie: | ex., Pontarmé, 11.V.1969, leg. Tronquet (cBor). Aquitaine: | ex., Lot-et-Garonne, leg. Pandellé (NHMW); 1 ex., Bordeaux, 11.V.1952, leg. Tempere (MHNG); | ex., Gironde, Cazaux, 29.V.1904 (MHNG); 1 ex., Gironde, Facture, 6.VII.1929, leg. Tempere (MHNG); 1 ex., Gironde, locality illegible, 28.VIII.1931, leg. Tempere (MHNG); 1 ex., Gironde, Guzmet, 3.111.1928, leg. Tempere (MHNG); | ex., Gi- ronde, Taussat-les-Bains, IV.1934, leg. Tempere (MHNG); | ex., Pyrénées Atlantiques, Sare, 400 m, 23.V.1966, leg. Tempere (MHNG); 2 exs., Sos, leg. Fauvel (DEI); 2 exs., Sos (NHMW). Bretagne: 7 exs., Morlaix, leg. Hervé (NHMW); 2 exs., Morlaix (NHMW). Midi-Pyrénées: | ex., Sarrant, 28.V.1936, leg. Giraud (MHNG); 5 exs., Hautes-Pyrénées, SW Ba- eneres-de-Luchon, Lac d’Oo, 1800 m, VII.1919 (NHMW); 5 exs., Saint-Sauveur-les-Bains near Luz- Saint-Sauveur, 17.1X.1965, leg. Tempere (MHNG). Li- mousin: | ex., Haute Vienne (NHMW). Ile-de-France: l ex., Fontainebleau, 16.X.1938, leg. Moingeon (MHNG ); 2 exs., Fontainebleau (MHNG): 1 ex., Versailles (MHNG); 1 ex., Paris (NHMW). Auvergne: | ex., Vi- chy, 12.V11.1934 (UCBA); | ex., Broút-Vernet, ant nest, 21.X11.1883 (MHNG). Centre: 2 exs., Bourges, leg. Boitel (MHNG); 4 exs., F. de Loches, leg. Mequignon (MHNG); 1 ex., Fréteval, leg. Mequignon (MHNG); 1 ex., Vendóme (MHNG); 1 ex., Perusson, leg. Mequi- gnon (MHNG). Rhóne-Alpes: 2 exs., Rhone, Charbon- nieres (MHNG ); 1 ex., Ardeche, Vals-Les-Bains (MHNG). Alsace: | ex., Strasbourg, VII.1955 (MHNG). Locality not identified or amgiguous: | ex., Forét de Moladier All., 23.X.1906 (cSch); 1 ex., St. Germain (MHNG); | ex., Las, leg. Fauvel (MHNG). Netherlands: | ex., Nuenen, mole nest, 10.1.1976, leg. Kanaar (cSch). Germany: Nordrhein-Westfalen: 13 exs., Kreis Stein- furt, Emsdettener Venn, 6.111.1999, leg. Assing, Wund- erle (cAss, cWun); 3 exs., Kreis Steinfurt, N Hopsten, Wiechholz, 6.111.1999, leg. Assing, Wunderle (cAss, cWun); | ex., Neuss, 5.1.1978, leg. Koch (cAss); 1 ex., Vreden, Wallburg, 26.X.1992, leg. Terlutter (cFel); 1 ex., Vreden, Vredener Feld, 3.111.1993, leg. Terlutter (cFel); 3 exs., Troisdorf, Wahner Heide, 12.1V.1987, leg. Wunderle (cWun); 1 ex., Köln, Troisdorf, Wahner Heide, 21.111.1987, leg. Köhler (cKöh); 3 exs., Viersen, Ritzeroder Moor, 19.V.1990, leg. Wunderle (cWun); 2 exs., Brühl, Staatsforst Ville, 20.11.1987, 5.X1.1989, leg. Köhler (cKöh); 1 ex., Köln-Worringen, Worringer Bruch, 5.VI11.1996, leg. Köhler (cKöh); 2 exs., Kölner Bucht, Pulheim-Sinnersdorf, 14.111 € 19.VI.1986 (cKöh); 2 exs., Kölner Bucht, Pulheim-Sinnersdorf, Chorbusch, 10.11.&2.V1.1986, leg. Köhler (cKöh); 1 ex., Mönchengladbach, Gerkerather Wald, 14.11.1986, leg. Köhler (cKöh); 1 ex., Münster-Wölbeck, Tiergarten, 5.VHI.1991, leg. Köhler (cKöh); 4 exs., Elmpt, Elmpter Bruch, 20.X1.1991, leg. Wunderle (cWun); 1 ex., Nie- derkrüchten, Elmpter Bruch, 20.X1.1991, leg. Köhler (cKöh). Rheinland-Pfalz: 3 exs., Lahnstein, NSG Kop- pelstein, 26.VI.1984 & 4.X1.1985, leg. Wunderle (cWun); | ex., Mainz (DEI); 1 ex., Wachenheim, Kam- mertsberg, 25.1V.1995, leg. Köhler (cKóh); 1 ex., Al- tenahr, Reimertzhoven, 11.1V.1987, leg. Köhler (cKöh); | ex., Ahrweiler (NHMW). Schleswig-Holstein: | ex., Schleswig, leg. Varendorff (NHMW). Niedersachsen: l ex., Lüneburger Heide, Niederhaverbeck, heathland, 15.X1.1993 (cAss); 1 ex., Lüneburg, Dahlenburg- Dahlem, 20.-21.V1.1994, leg. Köhler (cKöh); 3 exs., Soltau, Schneverdingen, 16.V.1998, 15.V.1999, 17.X.2000 (cAss); | ex., Winsen/Luhe, Laßrönne, 24.11.1995, leg. Assing (cAss); 76 exs., NW Hannover, Helstorf, heath- land, 1980 - 1990 (cAss); 2 exs., Celle, Scheuen, heath- land, X11.1984 (cAss); 1 ex., N Gifhorn, NSG “Heiliger Hain”, heathland, IX.1985 (cAss); 1 ex., Lehrte, Im- mensen, mixed forest, 30.1V.1986 (cAss); 1 ex., Mep- pen, Sprakeler Heide, heathland, [X.1987 (cAss); 2 exs., Oldenburg, Dötlingen, 22.1X.1970, leg. Lohse (MHNG); 1 ex., Oldenburg, Ostrittrum, 1.V.1971, leg. Lohse (MHNG). Hessen: | ex., Darmstadt, Eberstadt, beech forest, X1.1996, leg. Hetzel (cFel); 1 ex., Darmstadt, 15.V11.1958, leg. Vogt (cBor); 1 ex., Hanau (MHNG). Baden-Wiirttemberg: | ex., Karlsruhe, Baden, 18.1V.1951, leg. Nowotny (MHNG). Berlin/Branden- burg: | ex., Schlaubetal, 9.1.1999, leg. Pütz (cAss); 1 ex., Berlin Karlshorst, Biesenhorster Sand, 9.1V.2001, leg. Nickel (cSch); 2 exs., NSG Rietzer See, 52°23N, 12°40E, pine forest, 11.V.1978, leg. Uhlig (DEI). Thü- ringen: 2 exs., Eisenberg (NHMW); 2 exs., locality not specified (DEI, NAMW). Italy: 1 ex., Trentino-Alto Adige, Laces, 900 m, 10.V.1992, leg. Kapp (cKap). Locality not specified: 5 exs. (MHNG). Diagnosis: Highly variable species. 3.8-4.8 mm. Col- oration somewhat variable; usual coloration: head black, pronotum dark brown to blackish brown, usually no- ticeably lighter than head; elytra brown to dark brown, Volker ASSING: Western Palaearctic Medon 67 usually slightly lighter than pronotum; abdomen dark brown to blackish brown with the apex, the lateral and often also the anterior margins of the segments lighter; legs and antennae reddish brown to ferrugineous, mid- dle antennomeres often slightly darker than the basal and the apical ones. Head of very variable shape, slightly oblong and ap- proximately as wide as pronotum to weakly transverse and distinctly (ca. 1.15 x) wider than pronotum; eyes approximately half the length of postocular region or slightly larger; puncturation very variable, ranging from extremely dense and and fine, rendering the surface of the head almost completely matt, to moderately dense and coarser, with a small median area free of punctures, and with the interstices shining; punctures non-areolate; interstices with or without microsculpture (Figs. 74-75). Pronotum, too, with puncturation of variable size and density; interstices without microsculpture; surface on the whole with more shine than that of head (Figs. 74-75) Elytra of very variable size, 1.15-1.30 times as wide and at suture 1.05-1.17 times as long as pronotum (Fig. 74). Í: protarsomeres I - IV not dilated; posterior margin of sternite VII shallowly concave and laterally with combs of usually 5-9 palisade setae (Fig. 76); sternite VIII not distinctive. Aedeagus with long and slender ventral process, its lateral margins subparallel, shape of apex (ventral view) somewhat variable, truncate to weakly convex, middle area of ventral process in cross-section concave (Figs. 77-78). Comparative notes: Among its Western Palaearctic congeners, M. piceus is especially characterized by the shape of the aedeagus. In addition, it is distinguished by the following character combination: male protarsi not dilated, relatively dark coloration, puncturation of head dense, well defined, and not conspicuously coarse. In the collections examined, M. piceus was most often con- fused with M. brunneus, from which it is separated by its darker average coloration, the distinctly finer punctu- ration of the head, the longer and broader elytra, and by the shape of the aedeagus. Distribution and bionomics: Medon piceus is an At- lanto-Mediterranean element (Map 9), its distribution ranging from the south of the Iberian Peninsula to the south of England and Central Europe (material exam- ined). For a selection of additional records see BOHAC (1985), BRUGE et al. (2001), DRUGMAND (1989), Fow- LER (1888), KOCH (1968), KOHLER (2000), KOHLER & KLAUSNITZER (1998), SCHILLER (1989), TERLUTTER (1995), and VOGEL (1979). Medon piceus is unknown from Scandinavia and the Baltic countries (HANSEN 1996; LUNDBERG 1995; SILFVERBERG 1992), and from southeastern Europe (ASSING 2004a). In the British Isles, it has been reported only from a few localities in southern England. According to LOTT & DUFF (2003), the species was last recorded from England in 1968, but OWEN (2000) indicates a recent record from Bookham. Apparently, the abundance of the species has increased in some areas of Central Europe since the beginning of the 20th century (HORION 1965), and there may have been a recent expansion at the northeastern limit of its range. The first record from the Czech Republic was by BOHAC (1981); only four years later the same author re- ported it from seven additional Czech localities (BOHAC 1985). The species is still unknown from Slovakia. Based on a written statement by Scheerpeltz, HORION (1965) maintains that, in Austria, M. piceus is “aus allen Bundeslandern bekannt ...; zahlr. Belege in coll. Scheer- peltz”; however, the Scheerpeltz collection was exam- ined and not a single Austrian specimen was found in this or in any other collection housed in the NHMW. Until the presence of M. piceus in Austria is confirmed, it should be deleted from the lists of Austrian Staphylinidae. Similarly, the species has been reported from various regions in Italy, including Sicily (CICE- RONI & ZANETTI 1995; LUIGIONI 1929; PORTA 1926; ROTTENBERG 1870), but I have seen only a single Ital- ian specimen from the north (see material examined); KAHLEN (1987) reports the species from the same local- ity (Latsch = Laces). Most, if not all the other Italian re- cords, however, are presumably based on misidentifica- tions, as is suggested both by the revision of the material from various collections and the general distri- bution pattern of the species (Map 9). The record from Rhodos by SCHEERPELTZ (1964) is far outside the range of the species and doubtlessly erroneous. In the Iberian Peninsula, M. piceus is common in the south, the west and the northwest, but it is apparently absent from the northeast, where M. cauchoisi is abundant. Evidently, the species is much more common in the southwest than in the northeast of its range. Medon piceus has been collected from various kinds of nests (moles, ants, etc.) (material examined; see also HORION (1965), RENNER & GRUNDMANN (1984)), but these observations are probably accidental. In Germany, the species is usually found in the leaf litter and humus of bogs, swamps, heathlands, and forests (mostly conif- erous woods) with low-pH soils (material examined: HORION 1965; KOCH 1968; RENNER & GRUNDMANN 1984). In the Iberian Peninsula, according to personal observations, it occurs in various types of woodland (Quercus ilex and other oak species, pine forests, etc.) and in shrub habitats. Adult specimens have been observed throughout the year (material examined). In pitfall trap studies in northwest German heathlands (Assing, unpubl.), teneral beetles were recorded in late summer and autumn (Au- gust through November). On two occasions, mature eggs were observed in the ovaries of females collected 68 Bonner zoologische Beitráge 54 (2005) in May. Based on the material examined here, M. piceus is univoltine, not bivoltine as indicated by DRUGMAND (1989). Dissections of the thorax revealed that the flight muscles of most of the examined specimens from north- ern German heathlands were either rudimentary or ab- sent (Assing, unpubl.) and that consequently only part of the populations is capable of flight. Records of flying specimens are from May and July (KOHLER 2000; Vo- GEL 1979), 3.17. Medon cauchoisi Jarrige, 1949 (Figs. 79-84, Map 10) Medon cauchoisi Jarrige, 1949 (JARRIGE 1949: 66). Medon oculatior Peyerimhoff, 1949 (PEYERIMHOFF 1949: 249f) syn. n. Medon lusitanicum Coiffait, 1970a (COIFFAIT 1970a: 715f) syn. n. Map 9: Distribution of Medon piceus (Kraatz) (filled circles: revised records; open circles: selected literature records). Volker ASSING: Western Palaearctic Medon 69 Medon parviphallus Coiffait, 1973b (COIFFAIT 1973b: 278 ff) syn. n. Medon mazices Coiffait, 1973b (COIFFAIT 1973b: 280ff) syn. n. Medon fongondi Coiffait, 1980 (COIFFAIT 1980: 39f) syn. n. Medon perraulti Coiffait, 1980 (COIFFAIT 1980: 41) syn. n. Medon cribriceps Scheerpeltz 1. 1. Types examined: M. cauchoisi: Holotype @: St M. du Canigou, P.O. VII.48 Dr. Ph. C. / TYPE / Muséum Paris Ex Collection J. JARRIGE 1976 / cauchoisi / Holotypus Medon cauchoisi Jarrige rev. V. Assing 2003 (MNHNP). M. oculatior: Holotype ¢ [aedeagus missing]: O. el Akhdar pr. Tanant, Maroc, 28 juin 1923, Euphorbia serinifera [?] / TYPE / Medon oculatior Peyerimhoff, Type uniq. (MNHNP). M. lusitanicus: Holotype ¢: PORTUGAL ALGARVE, ALGOZ, H. COIFFAIT -168 / MUSEUM PARIS COLL H. COIFFAIT / HOLOTYPE / Medon lusitani- cum Coiff. H. COIFFAIT det. 1969 / Medon cauchoisi Jarrige det. V. Assing 2003 (MNHNP). M. parviphallus: Holotype ©: Maroc, Arbaoua, IV-72, H. C. / MUSEUM PARIS COLL H. COIFFAIT / HO- LOTYPE / Medon parviphallus Coiff. H. COIFFAIT det. 1972 / Medon cauchoisi Jarrige det. V. Assing 2003 (MNHNP). M. mazices: Holotype Y: MAROC 22.4.72, QUEZ- ZANE, H. COIFFAIT / MUSEUM PARIS COLL H. COIFFAIT / HOLOTYPE / Medon mazices H. COIF- FAIT det. 1973 / Medon cauchoisi Jarrige det. V. As- sing 2003 (MNHNP). M. perraulti: Holotype Ö [aedeagus missing]: St. Martin d’Albere / P.O. 26.111.70, G. Perrault / MUSEUM PA- RIS COLLECTION J. JARRIGE / TYPE / perraulti / Medon cauchoisi Jarrige det. V. Assing 2003 (MNHNP). M. fongondi: Holotype © [teneral]: Tizi n’Tchika, Ht Atlas, 9.V.67 / Maroc, H. Fongond / MUSEUM PARIS COLLECTION J. JARRIGE / TYPE / fongondi / Me- don fongondi H. COIFFAIT 1978 / Medon cauchoisi Jarrige det. V. Assing 2003 (MNHNP). Comments: The holotypes of M. cauchoisi, M. ocula- tior, M. lusitanicus, M. parviphallus, M. mazices, and M. perraulti are externally so similar that it seems most remarkable that they were described as different species. The aedeagus of M. parviphallus Coiffait is clearly tera- tologically malformed; for more details on such mal- formations see ASSING (2004a). An examination of the aedeagi and secondary sexual characters, as well as of the external characters of the other holotypes revealed no significant differences suggesting that they should represent distinct species. The teneral holotype of M. fongondi has a distinctly microsculptured head, but oth- erwise no evidence was found that it is specifically dis- tinct from M. cauchoisi, neither in the external nor in the sexual characters. Consequently, M. lusitanicus Coiffait, M. parviphallus Coiffait, M. mazices Coiffait, M. perraulti Coiffait, and M. fongondi Coiffait are all placed in the synonymy of the senior name M. cau- choisi. Since M. oculatior and M. cauchoisi were de- scribed simultaneously and neither of the names has priority by publication date, M. cauchoisi is here desig- nated as the senior name. (The original descriptions of both species were published in 1949; since there is no indication of the precise date, the publication date of both descriptions must be assumed to be December 31.) Additional material examined (total: 405 exs.): Morocco: 4 exs., N-Morocco, Jbel Mousa, leg. Franz (NHMW, cAss); 2 exs., N-Morocco, Xauen, leg. Franz (NHMW, cAss); 1 ex., Rif, Xauen, Puente Fomento, 30.11.1959, leg. Besuchet (cBor); 3 exs., Rif, Dardara, near Xauen, 30.111.1959, leg. Besuchet (MHNG ); 2 exs., Rif, Mt. Lexchab, 1.1V.1959, leg. Besuchet (MHNG); 1 ex., Moyen Atlas, Afourer, leg. Franz (cAss); 2 exs., Je- bel Tazeka, Taza, leg. Franz (NHMW, cAss); 2 exs., M. de Kebdana, 30 km NW Berkane, 35°03N, 02°36W, 350 m, 31.XI11.2001, leg. Bayer (cSch); 1 ex., Rif, 15 km SW Zinat, Tieta-des Beni-Yda-Cherki, 570 m, oak fo- rest, 14.11.2003, leg. Wrase (cSch); 2 exs., Tanger (DEI, IRSNB); 1 ex., Tanger, X11.1894 (MHNG); 3 exs., Oued Sebou near Kenitra, V.1961, leg. Comellini (MHNG); 1 ex., Oued Sebou, IV.1961, leg. Comellini (cAss); 3 exs., 10 km N Larache, Jemis del Sahel, 8.V.1960, leg. Besuchet (MHNG); 1 ex., Jemis del Sa- hel, leg. Franz (NHMW); 4 exs., Ceuta [35°53N, 05°20W], leg. Comellini (MHNG, cAss). Portugal: | ex., Algarve, Tavira, Alcaria do Cume, 37%15N, 07°44W, 440 m, 12.1V.2002, leg. Meybohm (cAss); 1 ex., Algarve, Serra de Monte Figo, 37°06N, 07°48W, 200 m, 4.1V.2001, leg. Meybohm (cAss); 5 exs., Algarve, Serra de Cume, Alcaria, 37°14N, 07°43W, 420-430 m, 31.1112001, leg. Meybohm (cAss); 23 exs., Algarve, 10 km N S. Brás de Alportel, 400 m, 4.V1.1992, leg. Wunderle (cAss); 1 ex., N Al- portel, 37°11N, 07°55W, 340 m, 4.1V.2002, leg. Mey- bohm (cAss); 4 exs., Algarve, N Faro, Freixo-Seco, 37°16N, 08°03W, 440 m, 9.1V.2001, leg. Meybohm (cAss); 2 exs., Algarve, Rocha da Pena, N-slope, 37°15N, 08°05W, 9.1V.2001, leg. Meybohm (cAss); 2 exs., Algarve, N Loule, Querenca, Fonte Benemola, 37°12N, 08°00W, 150 m, 5.1V.2002, leg. Meybohm (cAss); 12 exs., Algarve, Monchique, above road to Al- 70 Bonner zoologische Beitráge 54 (2005) ferce, 37°19N, 8°32W, 590 m, 10.1V.2002, leg. Mey- bohm (cAss); 31 exs., Caldas de Monchique, V- V1.1960, leg. Fagel (IRSNB); 2 exs., Faro, Silves, leg. Franz (NHMW ); 2 exs., Faro, Azinhal, 14.V.1994, leg. Bayer & Winkelmann (cSch). . Spain: Cataluña: | ex., Granollers, La Roca, leg. Franz (cAss); 6 exs., 40 km N Barcelona, Sierra de Montseny, 700 m, S-slope, 19.111.1994, leg. Assing, Wunderle (cAss, Wun); | ex., Sierra de Montseny, Santa Fe, leg. Franz (NHMW ); 5 exs., Sierra de Montseny, leg. Franz (NHMW); 1 ex., Barcelona, Vallirana, leg. Franz (NHMW ); 1 ex., Barcelona (MHNG); 3 exs., Barcelona, Montsenat, 3.1V.1956, leg. Coiffait (MHNG); 2 exs., Barcelona, Puerto Ordal, 22.111.1959, leg. Besuchet (MHNG); 2 exs., Gerona, Ripoll, 11.11.1978 (cAss); 2 exs., Gerona, Puerto de Collsuspina, leg. Franz (NHMW); 2 exs., Gerona, Tapis, 1000 m, 28.11.1997, leg. Tonquet (cTro); 2 exs., Arampruya, Begas, leg. Franz (NHMW); 3 exs., Gerona, Ripoll, 11.111.1978 (MHNG). Madrid: 3 exs., Sierra Guadarrama, Escorial, leg. Franz (NHMW). Aragón: 3 exs., 20 km SE Teruel, La Puebla de Valverde, 40°13N, 00°57W, 1100 m, 14.14.2003, leg. Assing (cAss); 8 exs., 50 km ESE Te- ruel, N Rubielos de Mora, 40°14N, 00°37W, 1255 m, oak forest, 13.1V.2003, leg. Assing (cAss); 1 ex., 30 km S. Teruel, Sierra del Javalambre, NE Javalambre, 40°09N, 01°00W, 1700 m, 14.1V.2003, leg. Assing (cAss); 5 exs., WNW Teruel, Sierra de Albarracin, Ori- huela d. T., 40°32N, 01°40W, 1520 m, 12.1V.2003, leg. Assing (cAss); 1 ex., Huesca, Sierra de Alcubierre, leg. Franz (NHMW). Castilla-La Mancha: 2 exs., Cuenca, 9.1V.1887, leg. Korb (DEI, NHMW); 1 ex., 90 km N Montoro, Sierra del Rey, Puerto de Niefla, 38°32N, 04°22W, 850 m, oak forest, 28.X11.2003, leg. Assing (cAss). Valencia: 2 exs., Serra, leg. Franz (NHMW); 1 ex., Alicante, Sierra de Bernia, 19.111.1978, leg. Me- ybohm (MHNG); 1 ex., Castellón, Cervera del Maestre, 13.V.1966, leg. Besuchet (MHNG). Murcia: 2 exs., Sierra de Espuña, leg. Franz (NHMW ); 4 exs., Sierra de Espuña, 26.111.1959, leg. Besuchet (MHNG). Andalu- cía: 7 exs., Ronda, Sierra de Grazalemma (CA), Puerto las Palomas, 1000 m, 25.111.1994, leg. Assing, Wunder- le (cAss, cWun); | ex., Sierra de Grazalemma, 1100 m, 1.X.1993, leg. Wunderle (cSch); 2 exs., Malaga, leg. Franz (NHMW); | ex., SE Ronda, Sierra de Palmitera (MA), 900 m, 24.111.1994, leg. Assing (cAss); 12 exs., Ronda, Sierra de Ubrique (CA), 1000 m, 25.111.1994, leg. Assing, Wunderle (cAss, cWun); 1 ex., Ubrique, 600 m, 2.X.1993, leg. Wunderle (cWun); | ex., Sierra de Ronda, Mte. Arastepa, leg. Franz (NHMW); 2 exs., Sierra de Bermeja, M. Los Reales, 1450 m, 26.111.1994, leg. Assing (cAss); 10 exs., Sierra de Bermeja, Jubrique, 350 m, 26.111.1994, leg. Assing, Wunderle (cAss, cWun); | ex., Sierra de Bermeja, 1000 m, 26.111.1994, leg. Assing (cAss); 1 ex., Sierra de Bermeja, Puerto de Peñas Blancas; 1200 m, 26.111.1994, leg. Wunderle (cWun); | ex., N Algeciras, 36°15N, 05°26W, moist fo- rest, 21.11.2000, leg. Lompe (cAss); 18 exs., Algeciras, Sierra de Luna (CA), 200 - 350 m, 28.111.1994, leg. As- sing, Wunderle (cAss, cWun); 6 exs., Sierra de Luna, 27.V.1966, leg. Besuchet, Comellini (MHNG); 1 ex., 8 km E Tarifa, Mirador del Estrecho, 36°03N, 05°33W, 310 m, under palm trees and bushes, 10.11.1999, leg. Zerche (DEI); 1 ex., 10 km W Los Barrios, 36°11N, 05°34W, 210 m, leaf litter below cork trees, 10.11.1999, leg. Zerche (DEI); 4 exs., Cadiz, 14 km NE Alcala de los Gazules, 36°32N, 05°39W, 430 m, cork tree forest, 2.11.1999, leg. Zerche (DEI); 5 exs., Algeciras, Sierra de Fates (CA), 350 m, 28.11.1994, leg. Assing, Wunderle (cAss, cWun); 1 ex., Algeciras, leg. Quedenfeldt (NHMW); 1 ex., Algeciras, 1./2.V.1956, leg. Fagel (IRSNB); 2 exs., Algeciras, leg. Fauvel (IRSNB, NHMW); 2 exs., Algeciras, Cerro de Mirador, leg. Franz (NHMW ); 4 exs., Sierra de Aracena, Acebuches, 37°53N, 06°48W, 5.1V.2001, leg. Meybohm (cAss); 2 exs., Cádiz, El Bujeo, 36°04N, 05°32W, 230 m, oak fo- rest, 9.IV.2001, leg. Sprick (cAss); 3 exs., Cadiz, Pto. de Bujeo, 30.V.1966, leg. Besuchet (MHNG); 1 ex., Cadiz, Los Barrios, 22.1V.1960, leg. Besuchet (cAss); 1 ex., Tarifa, Puerto de Ojén, 19.111.2002, leg. Amann (cAss); 1 ex., Tarifa, leg. Franz (NHMW ); 2 exs., Sierra de Cordoba, leg. Franz (NHMW); 1 ex., N Sevilla, Sier- ra Morena, leg. Franz (NHMW); | ex., Sierra de Cazor- la, source of Gualaquivir river, 1200 m, 6.X.1993, leg. Wunderle (cWun); 2 exs., Sierra de Cazorla, leg. Franz (NHMW); 2 exs., Sierra de Cazorla, 12.1V.1959, leg. Besuchet (cBor); 1 ex., W Lanjaron, Alpujarra, 1400 m, 23.111.1994, leg. Wunderle (cWun); 3 exs., Marbella, 1964, leg. Frey (NHMW ); 2 exs., Sierra Nevada, Laro- les, 300m, 22.111.1987, leg. Zoia (cZan); 2 exs., Vejer de la Frontera (CA), 27.111.1987, leg. Zoia (cZan); 4 exs., Malaga, Montes des Malaga, 28.111.1959, leg. Besuchet (MHNG ); 4 exs., Malaga, 4 km NW Yunquera, 36°40N, 04°57W, 725 m, 1.11.1999, leg. Zerche (DED e113 exs;, 15 km NNE Malaga, Montes de Malaga, 36°48N, 04°23W, 750 m, bank of stream with Salix, etc., 24.X11.2003, leg. Assing (cAss); 15 exs., 15 km NNE Málaga, Montes de Málaga, 36°48N, 04°22W, 900 m, mixed pine and corktree forest, 24.X11.2003, leg. Assing (cAss); 6 exs., 15 km NE Málaga, Montes de Málaga, 36°47N, 04°21W, 900 m, N-slope with Quercus suber, 24.X11.2003, leg. Assing (cAss); 1 ex., ca. 20 km S Lo- ja, 5 km NW Zafarraya, 36°59N, 04°10W, 900 m, Quercus ilex forest, 24.X11.2003, leg. Assing (cAss); 23 exs., ca. 30 km NW Córdoba, E Villaviciosa, 38°05N, 04°54W, ca. 300 m, corktree forest, 28.X11.2003, leg. Assing (cAss); 4 exs., ca. 30 km NW Córdoba, E Vil- laviciosa, 38°05N, 04°53W, ca. 500 m, corktree litter at roadside, 28.X11.2003, leg. Assing (cAss); 4 exs., 60 km N Montoro, N Azuel, 38°19N, 04°19W, ca. 600 m, N- slope with scattered corktrees and shrubs, litter and Volker ASSING: Western Palaearctic Medon 7] grass roots sifted, 28.X11.2003, leg. Assing (cAss); 1 ex., locality not specified (NHMW). Gibraltar: 8 exs., The Upper Rock Nature Reserve, 36°09N, 05°21W, 350 m, leaf litter under bushes, 5.11.1999, leg. Zerche (DEI). France: Languedoc-Roussillon: 2 exs., Vernet les Bains, Rte. de Sahorre, 700 m, 17.V1.1999, leg. Wolf (cSch); 1 ex., Molitg les Bains, 500 m, sifted from dry plants, 22.X1.1996, leg. Tronquet (cTro); 1 ex., same data, but 30.111.2001 (cTro); 1 ex., Molitg les Bains, 600 m, 13.V1.1995, leg. Tronquet (cTro); 1 ex., Molitg les Bains, 550 m, sifted moss, 22.11.1997, leg. Tronquet (cTro); 2 exs., Molitg les Bains, 22.11.2002, leg. Tron- quet (cTro); 1 ex., Campóme, 1250 m, 9.IV.1999, leg. Tronquet (cTro); 1 ex., Campóme, 700 m, 13.14.2001, leg. Tronquet (cTro); 4 exs., Collioure, 250 m, 3.X11.1999, leg. Tronquet (cTro); 4 exs., Collioure, 7.1.2000, leg. Tronquet (cTro); 1 ex., Fenouillet, gorges de Saint-Jaume, 11.X1.2002, leg. Tronquet (cTro); 4 exs., Corneilla-de-Conflent, 800 m, Quercus ilex litter, 25.11.2000, leg. Tronquet (cTro). DR. 5 ES e. 00 4 xy = 4 > E ® 2 a “& u Diagnosis: 3.5 - 4.5 mm. Habitus as in Fig. 79. Colora- tion somewhat variable; usual coloration: castaneous to blackish brown, with the head usually slightly darker than pronotum and elytra; legs and antennae testaceous yellowish brown, with the basal antennomeres usually reddish brown to ferrugineous. ] Head of variabie shape, usually at least weakly oblong, rarely about as wide as long; temples behind eyes sub- parallel or weakly converging; eyes moderately large, in dorsal view slightly shorter than half the length of postocular region; puncturation coarse, dense, and well- defined, interstices reduced to narrow ridges or nearly so; occasionally in median dorsal area with small spot with slightly sparser puncturation; interstices usually without microsculpture and shining, sometimes with shallow microsculpture in posterior part of head, or very rarely microsculptured everywhere and matt (Fig. 80). Pronotum approximately as wide as long and usually slightly narrower than head; puncturation variable, as coarse and dense as that of head or somewhat finer and sparser; median line without punctures; microsculpture absent (Fig. 80). Map 10: Distributions of Medon cauchoisi Coiffait (filled circles) and of M. despectus (Fairmaire) (open circles) in the Western Mediterranean, based on examined records only. 72 Bonner zoologische Beitráge 54 (2005) Elytra of variable length and width, usually approxi- mately 1.2 times as wide as and at suture at least slightly longer than pronotum (Fig. 79) 7: protarsomeres I - IV not dilated; posterior margin of sternite VII shallowly concave and laterally with combs of 6-10 (mostly 8) palisade setae (Fig. 81); posterior in- - cision of sternite VIII not very deep. Aedeagus as in Figs. 82-84, central area of ventral process in cross- section concave. Comparative notes: Among most of its Western Medi- terranean congeners, M. cauchoisi is readily identified by the coarse, dense, and well-defined puncturation of the head and the pronotum. Regarding the puncturation of the head, the species is similar to M. brunneus, which, however, differs by its usually larger size, its larger and usually not oblong head, the narrower and shorter elytra, and especially by the morphology of the aedeagus. In M. mirei, the head is relatively larger, of subcircular shape, and less shining, the elytra are much shorter and narrower, the hind wings are reduced, the palisade fringe at the posterior margin of the abdominal tergite VII is reduced, and the aedeagus is of different morphology. Distribution and bionomics: Medon cauchoisi is an Atlanto-Mediterranean element (Map 10), its distribu- tion including Morocco, southern Portugal, the south- ern, central, eastern, and northeastern parts of Spain, and the Pyrénées-Orientales in southwestern France. In- terestingly, it is apparently absent from the northwestern half of the Iberian Peninsula, where M. piceus is com- mon. Similarly, there is only little overlap of the range of M. cauchoisi and that of the closely related M. brun- neus (Map 8) in northeastern Spain (Cataluña) and southwestern France (Pyrénées-Orientales). The distri- butions of M. cauchoisi and M. despectus, another closely allied species, are apparently strictly allo- or parapatric (Map 10). M. cauchoisi has been collected in a wide range of — usually forest — habitats, mostly in the leaf litter of vari- ous species of oak, but also from debris below palm trees and bushes, and from moss. Adult beetles were found during the period from October through June, but the absence of records from the period July to Septem- ber may also be due to low collecting activity in the re- gion where M. cauchoisi occurs. 3.18. Medon despectus (Fairmaire, 1860) (Figs. 85-89, Map 10) Lithocharis despecta Fairmaire, 1860 (FAIRMAIRE 1860: 160). Type examined: Syntype Y: Philippeville / Syntype / despecta Fairm / Coll. A. Fauvel, R.I.Sc.N.B.17.819 / Medon despectus (Fairmaire) det. V. Assing 2003 (IRSNB). Additional material examined (total: 168 exs.): Tunisia: | ex., SW Zaghouan, Jebel Zaghouan, 36°22N, 10°07E, 800 m, N-slope, litter and grass sifted, 29.X11.2004, leg. Assing (cAss); 3 exs., ca. 25 km SW El Fahs, 36°15N, 09°48E, 340 m, reservoir, stream val- ley with poplar etc., moss and litter sifted, 25.X11.2004, leg. Wunderle (cWun); 1 ex., Falnassa, 15 km N Beja, 500 m, 1.X.1995, leg. Schulz & Vock (cAss); 4 exs., “Tunis”, leg. Fauvel (NHMW, cAss); 1 ex., El Feidja (NHMW); 1 ex., Kasserine, X.1937, leg. Demofys (cTro); 31 exs., 25 km W Jendouba, ca. 10 km N Ghar- dimaou, Ain Soltane, 36°29N, 08° 19E, 670 m, litter of oak forest, 27.X11.2004, leg. Assing (cAss); 20 exs., same data, but 600 m, 28.X11.2004, leg. Assing (cAss); 4 exs., NW-Tunisia, Ghardimaou Forest, El Feidja, 700 m, 13.1V.1989, leg. Meregalli (DEI, cAss); 1 ex., Fei- dja, Aín Soltane, 700 m, 13.1V.1989, leg. Meregalli (DEI); 1 ex., Teboursouk (IRSNB). Algeria: | ex., El Biar, [1.1909 (NHMW); 1 ex., Tem- mapes (?), leg. Théry (cAss); 1 ex., Bols de Bainen, [11.1909 (cAss); 3 exs., St. Charles, leg. Théry URSNB, NHMW); 1 ex., Algier (DEI); 2 exs., Algier (MHNG, cAss); 1 ex., Laverdure, 6.X.1929, leg. Schatzmayr (UCBA); 1 ex., Bone (IRSNB); | ex., Bou Berak, near Dellys (IRSNB); 5 exs., Grand Kabylie, Azazga — Ya- kouren, 600 m, 14.V.1988, leg. Besuchet, Lóbl & Burc- khardt (MHNG, cAss); 14 exs., Grande Kabylie, Foret d’Akfadou, 26 km E Yakouren, 1200 m, 16.V.1988, leg. Besuchet, Löbl & Burckhardt (MHNG, cAss); 1 ex., Grande Kabylie, Yakouren, 730 m, 12.V.1988, leg. Be- suchet, Lóbl & Burckhardt (MHNG); 2 exs., Oued Sé- baou, W Dellys, 20.V.1988, leg. Besuchet, Lóbl & Bur- ckhardt (cAss); 4 exs., Grande Kabylie, |’Arbatache sur El Kseur, 300-400 m, 18.V.1988, leg. Besuchet, Lóbl & Burckhardt (MHNG, cAss); 13 exs., Grande Kabylie, Oued Aissi, Ouadhia, 200 m, 12.V.1988, leg. Besuchet, Lóbl & Burckhardt (MHNG, cAss); 13 exs., Grande Kabylie, Djebel Bou-Berak, 350 m, 19.V.1988, leg. Be- suchet, Lóbl & Burckhardt (MHNG, cAss); 2 exs., Djurdjura, sur Boghni, 11.V.1988, leg. Besuchet, Lóbl & Burckhardt (MHNG); 8 exs., Djurdjura, 4 km SW Ti- kjda, 1200 m, 7.V.1988, leg. Besuchet, Lóbl & Burck- hardt (MHNG, cAss); 5 exs., Gorges de la Chiffa, Ruisseau des Singes, 280 - 380 m, 4.V.1988, leg. Besu- chet, Lóbl & Burckhardt (MHNG, cAss); 3 exs., Larba, 180 m, 4.V.1988, leg. Besuchet, Lóbl & Burckhardt (MHNG, cAss). Italy: 14 exs., Pantelleria, Mte. Gibele, 600m, 12.1X.1986, leg. Sette (cZan, cAss); 2 exs., Pantelleria, 11.1913, leg. Dodero (DEI). Volker ASSING: Western Palaearctic Medon 73 Figs. 74-94. Medon piceus (Kraatz) (74-78), M. cauchoisi Jarrige (79-84; 84: holotype), M. despectus (Fairmaire) (85-89), and M. mirei Coiffait, holotype (90-94): forebody (74); head and pronotum (75, 80); habitus (85, 86, 79, 90); male sternite VII (76, 81, 87, 91); aedeagus in lateral and in ventral view (77-78, 82-84, 88-89, 92-94). Scale bars: 74-75, 79-80, 85-86, 90: 1.0 mm; 76-78, 81 -84, 87-89, 91-94: 0.2 mm. 74 Bonner zoologische Beitráge 54 (2005) Locality not specified or ambiguous: | ex. (IRSNB); | ex., with several locality labels (IRSNB). Diagnosis: External characters as in M. cauchoisi, ex- cept for the paler average coloration, the slightly lower average size, and the mostly shorter elytra (Figs. 85-86). Wing-dimorphic species; elytra at suture in macropter- ous specimens 1.05 - 1.10 times as long as pronotum (Fig. 85), in submacropterous specimens approximately as long as pronotum (Fig. 86). ¢: secondary sexual characters as in M. cauchoisi (Fig. 87). Aedeagus smaller and more slender than in M. cau- choisi, apically rounded both in lateral and in ventral view (Figs. 88-89). Comparative notes: Among Western Mediterranean Medon, M. despectus is most similar to M. cauchoisi, from which it is reliably separated only by the different shape of the aedeagus. Medon mirei, which, too, occurs in Northwest Africa, has much smaller eyes, completely reduced hind wings, and it lacks the palisade fringe at the posterior margin of the abdominal tergite VII. From the closely related, highly variable M. semiobscurus (Fauvel) from the Eastern Mediterranean, whose aedeagus has a similar shape, M. despectus is distin- guished by lower average size, shorter antennae with shorter and less oblong antennomeres III - VI, as well as by a smaller and less massive aedeagus. Distribution and bionomics: The known distribution of this species is confined to Tunisia, Algeria, and the Italian island Pantelleria (Map 10), from where it was previously recorded as M. brunneus (CICERONI & ZANETTI 1995). For some additional records see NOR- MAND (1935). The material examined was collected at altitudes of 180-1200 m in autumn (September, Octo- ber), winter (December, February), and in spring (March through May). At least 52 Tunisian specimens were sifted from the leaf litter of an oak forest. 3.19. Medon mirei Coiffait, 1980 (Figs. 90 - 94) Medon mirei Coiffait, 1980 (COIFFAIT 1980: 40f). Medon giachinoi Bordoni, 1988 (BORDONI 1988: 343 ff) syn. n. Types examined: Medon mirei: Holotype ¢@: Ft. de Guerrouch, Alg. P. de Mire / TYPE / Muséum Paris Ex Collection J. Jarrige 1976 / Fagete / Medon mirei H. Coiffait 1979 (MNHNP). Medon giachinoi: Holotype Ö: Algeria — Juel, F.t de Guerrouch, 4/10 — XI — 1984 / M. R. S. N. Spedizione “Algeria “84” Boffa — Casale — Cavazzuti — Gavetti — Giachino — Levi / Holotypus / Medon giachinoi Bordoni det. 1987 / Medon mirei Coiffait det. V. Assing 2003 (MRSNT). Paratypes: 12 exs.: same data as holotype (MRSNT, cBor, cAss). Comments: The locality on the label of the type speci- men of M. mirei is not identical with the type locality indicated in the original description: “gorges de Kedaia, Algérie” (COIFFAIT 1980). The description and the col- lector, however, are in agreement with the above type specimen, and Coiffait’s citation of type labels is not always reliable (see M. siculus). The original description of M. giachinoi is based on 23 specimens collected in the same locality as the holotype of M. mirei. As was expected, the examination of the holotype and 12 paratypes of M. giachinoi revealed that it is conspecific with the holotype of M. mirei; hence the synonymy indicated above. BORDONI (1988) compares M. giachinoi with various congeners, but does not men- tion M. mirei. Diagnosis: Small species, 3.3 mm. Habitus as in Fig. 90. Coloration uniformly ferrugineous, appendages tes- taceous. Head of almost subcircular shape, approximately as wi- de as long; eyes of distinctly reduced size, in dorsal view about one fourth the length of postocular region, and not projecting from lateral outline of head; punctu- ration coarse, dense, and areolate; interstices reduced to narrow ridges and microsculptured; dorsal surface matt. Pronotum distinctly narrower than head, approximately 0.9 times its width, and slightly longer than wide; punc- turation coarse, dense, areolate, and partly confluent; in- terstices and median line with distinct shine, mi- crosculpture absent. Elytra only slightly wider (about 1.1 times) than and at suture only 0.75 times as long as pronotum; punctura- tion moderately dense; interstices distinctly shining. Hind wings reduced. Palisade fringe at posterior margin of abdominal tergite VII reduced. ©: posterior margin of sternite VII of similar shape and chaetotaxy as in M. cauchoisi (Fig. 91); sternite VIII posteriorly with rather broad and not very deep emargi- nation. Aedeagus of similar general morphology (shape and internal structures) as in M. cauchoisi, but in ventral view apically with converging lateral margins and with- out subapical dilatation, and in lateral aspect apically rounded (Figs. 92-94). Comparative notes: Medon mirei is readily distin- guished from its Western Palaearctic congeners by the small eyes, the small body size, the short elytra and re- duced hind wings, and by the male sexual characters. Distribution and bionomics: The species is known only from the type locality in Algeria. The adaptive re- Volker ASSING: Western Palaearctic Medon 75 ductions of eye size, pigmentation, and wings suggest that 1t has a subterranean habitat. 3.20. Medon fusculus (Mannerheim, 1830) (Map 11) Rugilus fusculus Mannerheim, 1830 (MANNERHEIM 1830: 40). Lithocharis fusculus Lacordaire, 1835 (LACORDAIRE 1835: 431). Lithocharis testaceus Lacordaire, 1835 (LACORDAIRE 1835: 432). Lithocharis rufus Mulsant & Rey, 1853 (MULSANT & REY 1853: 66). Lithocharis auranitica Saulcy, 1864 (SAULCY 1864: 40). Lithocharis infuscatus Baudi, 1870 (BAUDI 1870: 392). Medon deficiens Hubenthal, 1911 (HUBENTHAL 1911: 188). Medon abchasicus Bernhauer, 1922 (BERNHAUER 1922: 124). Medon bulgaricus Coiffait, 1970b (COIFFAIT 1970b: 105f). Medon gajaci Coiffait, 1973a (COIFFAIT 1973a: 114ff). Medon paradobrogicus Decu & Georgescu, 1994 (DECU & GEORGESCU 1994: 49f). Types examined: See ASSING (2004a). Additional material examined (total: 216 exs.; for ad- ditional material examined see ASSING (2004a)): France: Aquitaine: 2 exs., Gironde, Cambes, 18.X11.1936, leg. Giraud (MHNG, cAss); 3 exs., Cam- bes, 20.X1.1936 MHNG). Picardie: | ex., Oise, Lai- gneville, IX.1908, leg. Mequignon (MHNG); 11 exs., Laigneville, 24.1V.1924 (MHNG); 1 ex., Laigneville (MHNG); 2 exs., Oise, Noyon, leg. de Brunier (MHNG). Champagne-Ardenne: 2 exs., Les Grandes Loges (MHNG); 2 exs., Les Grandes Loges, VIII.1918 (MHNG). Íle-de-France: 1 ex. Fontainebleau (MHNG). Rhóne-Alpes: | ex., Haute-Savoie, Malagny, 14.X11.1963, leg. Comellini (MHNG); 1 ex., Rhone, Iri- gny, Rhone inundation, X11.1954 (MHNG); 1 ex., Isere, Grenoble, 12.V11.1902 (MHNG); 1 ex., Ain, Poncin, 1.1948 (MHNG). Alsace: 2 exs., Strasbourg, VIII.1953, IV.1954 (MHNG). Lorraine: 2 exs., Metz (MHNG). Locality not specified, ambiguous, or not identified: 2 exs., locality not specified (NHMW); 1 ex., “IN Cen- drieux Bourlet”, 1111960 (MHNG); 1 ex., “Cussac” (MHNG). England: | ex. [aedeagus and male sternite VII tera- tological], East Sussex, Rye, 14.11.1999, leg. Hance (cOwe). Italy: Friuli-Venezia Giulia: | ex., Villotta, 19.11.1973, leg. Zanetti (cBor); 1 ex., W Carnia, bank of Fella river, 45°23N,. 13°07E,-250 m, flood debris, 12.IX.1998, leg. Schúlke (cSch); 2 exs., bank of Isonzo river SE Villesse, 45°51N, 13°27E, 10 m, flood debris, 13.1X.1998, leg. Schiilke (cSch); 2 exs., Villotta (PN), 19.11.1973, leg. Zanetti (UCBA, cZan); 3 exs., Sacile (PN), mole nest, 6.111.1974, leg. Osella & Zanetti (cZan); 5 exs., Gemona del Friuli (UD), mole nest, 6.111.1974, leg. Osella & Za- netti (cZan); 1 ex., Monfalcone, 21.11.1948, leg. Sprin- ger (UCBA); 1 ex., Udine (MHNG). Germany: Niedersachsen: | ex., Hameln, Kónigsfór- de, garden, V.1988, leg. Sprick (cAss). Nordrhein- Westfalen: | ex. [with worker of Lasius brunneus atta- ched to the pin], Kólner Bucht, Pulheim, Stommeler- busch, 4.V.1984, leg. Kóhler (cKóh). Rheinland-Pfalz: 2 exs., Bacharach, bank of Rhine river, flood debris, I- 11.2001, leg. Hetzel (cFel); 3 exs., Bacharach, Steeg, vi- neyard, 56.V1.1996, leg. Köhler (cKöh); 1 ex., same da- ta, but 14.V11.1996 (cKöh); 2 exs., Oppenheim, 14.X.1989, leg. Höhner (cKöh); 1 ex., Schloßböckel- heim, 25.1V.1994, leg. Köhler (cKöh); 3 exs., same da- ta, but 31.V.1996; 1 ex., same data, but 28.VII.1996 (cKöh); 2 exs., Monzingen, 18.-19.V.1991, leg. Köhler (cKóh). Baden-Wiirttemberg: | ex., Kaiserstuhl, Dur- bach, 14.V11.1971 (MHNG). Hessen: | ex., Groß- Gerau, 6.11.1988, leg. Höhner (cKöh). Berlin/Branden- burg: 13 ex., Rüdersdorf, Kalkberge (DEI, cSch); 1 ex., Glambeck (DEI); 9 exs., Berlin, Hohen Neuendorf (DEI); 4 exs., Oderberg (DEI). Austria: Vorarlberg: | ex., Feldkirch (NHMW). Nie- derösterreich/Wien: 27 exs., Wien, several localities (DEI, MHNG, NHMW); 6 exs., Wiener Wald, Rüders- dorf (NHMW); 1 ex., Wien, Donauauen, 29.X1.1952, leg. Malicky (MHNG); 4 exs., Stockerau (NHMW); 1 ex., Albern (NHMW); 1 ex., Metzendorf (NHMW); 1 ex., Rodaun (NHMW); 1 ex., Mödling, Eichkogel (NHMW); 1 ex., Laxenburg (NHMW); 1 ex., Bisam- berg (NHMW); 4 exs., Bisamberg, 27.111.1953, leg. Ma- licky (MHNG); 1 ex., Ulrichskirchen (NHMW); 1 ex., Wechselgebiet (NHMW). Steiermark: | ex., Graz (MHNG); 1 ex., “Siid-Steiermark” (NHMW). Czech Republic: 10 exs., Moravia, Bulhary, 25.V1.1984, leg. Prudek (cSch, cAss). Bulgaria: | ex., Asenovgrad (MHNG). Croatia: 1 ex., Mlini [42°37N, 18°12], 29.IX.-4.X.1958, leg. Benick (MHNG); 3 exs., Brela [43°22N, 16°56E], 16.-26.1X.1958, leg. Benick (cAss). 76 Bonner zoologische Beitráge 54 (2005) Bosnia-Herzegovina: | ex., Montenegro, Budva, leg. Hummler (MHNG); 4 exs., Montengro, Herzeg-Novi (MHNG). Serbia-Montenegro: | ex., Zvonacka Banja, 28.1V.2002, leg. Hlavác (cAss). Greece: 8 exs., Voiotia, Levadia, wet moss, 4.1X.1964, leg. Puthz (MHNG); 2 exs., Makedhónia, Nomós Kavá- la, 10 km W Eleftheröpoli, Pangéo, 900 m, beechforest, 10.1V.1994, leg. Schawaller (SMNS); 2 exs., Makedho- nia, Nomós Kavála, 10 km N Paleá Kavála, 600 m, macchia & chestnut forest, 20.1V.1994, leg. Schawaller (SMNS, cAss); 1 ex., Makedhónia, Nomós Kavala, Ne- stos delta S Khrisöpoli, floodplain forest, 26.1V.1994, leg. Schawaller (SMNS); 2 exs., Thrakia, Nomós Xán- thi, Nestos near Toxótes, 25.1V.1994, leg. Schawaller (SMNS, cAss); 3 exs., Pelopónnisos, Kalavrita, 800 m, 3.1V.1971, leg. Löbl (MHNG); 1 ex., Pelopönnisos, Ta- ygetos, Meligalás, Dhervénion, 600 m, 26.111.1992, leg. Frisch (MNHUB); 1 ex., Corfu, leg. Hummler (MHNG). Turkey: Tokat: 6 exs., Tokat — Almus, 1200 m, 21.V.1967, leg. Besuchet (MHNG). Adana: 2 exs., Kozan, 600 m, 5.V.1967, leg. Besuchet (MHNG); | ex., Kozan, 300 m, 5.V.1967, leg. Besuchet (MHNG). Konya: | ex., Cam- lik env., Toroslar mts., 1345-1380 m, 6.-8.V1.2003, leg. Smatana (cSch). Adiyaman: | ex., Gólbasi, 900 m, 10.V.1967, leg. Besuchet (MHNG). Ukraine: | ex., Odessa (NHMW). Locality not specified or illegible: 10 exs. (MHNG). Diagnosis: See ASSING (2004a). Distribution and bionomics: Medon fusculus is a widespread Ponto-Mediterranean element (Map 11), its distribution ranging from the Middle East, eastern Ana- tolia, West Caucasus, and Ukraine in the southeast and east to the southwest of France, southern England, and southern Norway, southern Sweden, and Lithuania in the west, northwest, and north (material examined; ASSING 2004a). For a selection of additional records see BOHAC (1985), BRUGE et al. (2001), DRUGMAND (1989), FOWLER (1888), HORION (1965), HUGENTHOBLER (1966), KOCH (1968), KOHLER (2000), KOHLER & KLAUSNITZER (1998), LINKE (1907), SCHOLZE & JUNG (1993), SILFVERBERG (1992), SZUJECKI (1968), TER- LUTTER (1995), and UHLIG & VOGEL (1981). On the whole, M. fusculus is much more common in the south- east of its range (Turkey, Greece) than in the northwest, where it is rather rare. In France, it is absent from the Pyrénées-Orientales (TRONQUET 2001). In the British Isles, it is local and confined to the southern parts of England (FOWLER 1888). According to HANSEN (1996), it is absent from Denmark. In Sweden, it is known only from Sódermanland (LUNDBERG 1995), but this record is somewhat ‘doubful (M. SÓRENSSON, Lund, pers. comm. 2003). OUTERELO (1980) reports a record from southern Spain (Andalucia), adding that the species is known from Spain and Portugal. As can be inferred from the general distribution and the abundant Medon material examined from the Iberian Peninsula, however, M. fusculus is absent from the Iberian Peninsula and these records are most likely to be based on misidentifi- cations (probably confusion with M. ripicola). Medon Jusculus has been reported from various regions in Italy, including Sardinia and Sicily (CICERONI & ZANETTI 1995; HORION 1965; LUIGIONI 1929; PORTA 1926), but the results of the present revision suggest that all the re- cords, except for those from the northeast (Friuli- Venezia Giulia), are probably erroneous. Medon fusculus has been collected from various kinds of substrates, especially leaf litter, nests of mammals (mole, fox, vole, rabbit), compost, hollow trees, and flood debris in forests, gardens, gravel pits, river banks, etc. (ASSING 2004a; BOHAC 1985; FRANZ 1938; HORION 1933, 1965; WAGNER 2002; ZERCHE 1980). Many re- cords are from mole nests; OSELLA & ZANETTI (1974) repeatedly found the species in this habitat, partly in larger numbers. On the other hand, STROUHAL & BEIER (1918) found only six specimens in 118 nests examined. According to NOWOSAD (2000) only very few speci- mens were collected in a total of 7000 nests studied in Poland; NOWOSAD (1990) had not found more than two specimens in 5000 mole nests dug up all over Poland. MEYBOHM (1997) reports two flight records (car-net); on one occasion 15 flying specimens were collected. Adult beetles have been collected throughout the year. DRUGMAND (1989) states that the species is bivoltine, but there is no evidence supporting this. Based on the material examined here, M. fusculus is univoltine. Ac- cording to BOHAC (1985), teneral specimens were ob- served in March. In the male seen from England, the male sternite VII and the aedeagus were teratologically malformed. 3.21. Medon sardous Dodero, 1922 (Figs. 95-99) Medon sardous Dodero, 1922 (DODERO 1922: 67f). Material examined (total: 18 exs.): Italy, Sardegna: 6 exs., NNW Villaputzu, Salto di Quirra, 400 m, 9.X.1989, leg. Wunderle (cWun, cAss); 3 exs., grotta S. Puddu near Villaputzu, 17.V111.1973 (cBor); 6 exs., Flumini (NHMW, cAss); 1 ex., Lanusei (NU), 12.V.1980, leg. Torchia (cZan); 1 ex., NW Lanu- sei (NU), 650 m, 21.1V.1992, leg. Schawaller (SMNS ); l ex., N Domusdemaria, 38°58N, 8°52E, 23.X.1981, leg. Malicky (cAss). Volker ASSING: Western Palaearctic Medon 77 rs M Map 11: Distribution of Medon fusculus (Mannerheim) (filled circles: revised records; open circles: selected literature records). Diagnosis: In external appearance similar to M. fuscu- lus. 4.5-5.5 mm. Coloration reddish brown to dark brown, with the head usually slightly darker; legs and antennae yellowish brown to ferrugineous. Head about as wide as long or weakly transverse and 1.10-1.15 times as wide as pronotum (Fig. 95); eyes relatively large, in dorsal view more than half the length of temples; puncturation dense, coarse, well-defined, and areolate; interstices with noticeable shine. Pronotum approximately as wide as long or weakly transverse (Fig. 95); puncturation on the whole similar to that of head, but usually shallower, slightly less dense, and less well-defined; usually with, more rarely without impunctate median line. Elytra relatively large, 1.20-1.25 times as wide and at suture about 1.1 times as long as pronotum (Fig. 95); puncturation much finer and less dense than that of head and pronotum; interstices shining. Hind wings appar- ently fully developed. Protarsomeres I - IV without sex- ual dimorphism, transverse in both sexes; tarsi relatively short, metatarsomeres III and IV less than twice as long as broad. Abdomen finely and densely punctate; posterior margin of tergite VII with palisade fringe. $: posterior margin of sternite VII of similar morphol- ogy and chaetotaxy as in other species of the M. fuscu- lus group, deeply and broadly concave (not distinctly trapezoid), with two lateral combs of palisade setae and on either side of middle with a distinct tuft composed of very long dark setae (Fig. 96); sternite VIII similar to that of M. fusculus and related species (Fig. 97); aedeagus of distinctive shape (Figs. 98-99). Systematics and comparative notes: Based on external similarities (relatively large size, puncturation of head and pronotum), the male secondary sexual characters, especially the characteristic shape and chaetotaxy of sternite VII, and on the morphology of the aedeagus, M. sardous doubtlessly belongs to the M. fusculus group. From all the species of this group, none of which occurs in Sardinia (but see notes on M. ripicola below), it is distinguished especially by the distinctive shape of the aedeagus. From M. fusculus, the only geographically close congener of the fusculus group, it is additionally distinguished by the more well-defined puncturation and more shining interstices of the head and pronotum, by 78 Bonner zoologische Beitráge 54 (2005) the shorter tarsi (metatarsomeres HI and IV in M. fuscu- lus more than twice as long as broad), transverse protar- someres | - IV, the concave (not trapezoid) posterior ex- cavation of the male sternite VII, and by the much more distinct and longer tufts of setae at the posterior margin of the male sternite VII. Distribution and bionomics: Medon sardous is en- demic to Sardinia and — together with M. fusculus, M. ripicola, and M. kabylicus — one of the westernmost rep- resentatives of the M. fusculus group. Additional Sar- dinian localities are summarized by BORDONI (1981), all of them are in the southern half of the island. The spe- cies was sifted from leaf litter and found in caves. 3.22. Medon ripicola (Kraatz, 1854) (Fig. 100, Map 12) Lithocharis ripicola Kraatz, 1854 (KRAATZ 1854: 127). Lithocharis kellneri Kraatz, 1875 (KRAATZ 1875: 123). Medon pectiniventris Donisthorpe, 1932 (DONISTHORPE 1932: 252). Types examined: See ASSING (2004a). Material examined (total: 379 exs.; for additional ma- terial see ASSING (2004a)): Tunisia: 2 exs., Ain Draham (= Ayn ad Darahim; 36°47N, 8°42E), leg. Bodemeyer [1 paralectotype of M. bodemeyeri Bernhauer] (DEI, MHNG); 2 exs., Ain Dra- ham, 1X.1944, leg. Demoflys (cTro); 2 exs., Ain Dra- ham, VIII.1945, leg. Demoflys (cTro). Algeria: 2 exs., Grande Kabylie, Oued Sébaou, W Del- lys, 20.V.1988, leg. Besuchet, Lóbl & Burckhardt (MHNG, cAss). Morocco: 3 exs., Salé, leg. Thery (NHMW); 2 exs., Haute Atlas, Oukaimeden, leg. Franz (cAss); 1 ex., Oued Sebou, IV.1961, leg. Comellini (MHNG). Portugal: 2 exs., Coimbra (NHMW); 1 ex., Algarve, Praia de Faro, 37°02N, 07°59W, 5 m, laguna, 26.111.2002, leg. Meybohm (cAss); 1 ex., Algarve, Serra de Monchique, Portela Viuva, 20.11.1999, leg. Mey- bohm (cAss); 1 ex., Lagoa da Casa, 28.11.1986, leg. Winkelmann-Klóck (cSch). Madeira: 2 exs., above Seixal, Rib. do Seixal, 550m, bank of stream, 31.11.1996, leg. Lompe (cAss); 1 ex., locality not speci- fied, leg. Franz (cAss). Azores: 6 exs., Sáo Miguel, La- goa do Congro, 27.V111.2003 (cAss). Spain: Cataluña: | ex., Seo de Urgel, leg. Franz (cAss); 5 exs., Seo de Urgel, Arfa, V.-VI.1962, leg. Fa- gel (IRSNB); 4 exs., Seo de Urgel, V.-VI.1962, leg. Fa- gel (IRSNB); 4 exs., Gerona, Figueres, 10.VII.1965, leg. Comellini (MHNG, cAss). Galicia: | ex., Lugo, Linares, 1000 m, 27.V.1996, leg. Starke (cAss); 1 ex., Orense, Manzaneda, VII.1956, leg. Gonzales (MHNG). Asturias: | ex., Playa de Concho de Artedo, 3 km W Cudillero, 43°34N, 06°11W, 14.V1.2000, leg. Wrase (cSch); 2 exs., W Puerto de Pajares, Puerto de la Cubilla — Sotiello, 500 - 1400 m, car-net, 8.V1.1991, leg. Wun- derle (cWun); 4 exs., Covadonga, V1.1965, leg. Fagel (IRSNB); 2 exs., Cudillero, Conche de Ariedo, 43°34N, 6°11W, 14.V1.2000, leg. Starke (cFel). Aragón: | ex., Sierra de Albarracín, V.1953, leg. Comellini (cBor). Andalucía: 2 exs., Sierra Nevada (GR), Lanjaron, 400 - 600 m, 23.111.1994, leg. Assing, Wunderle (cAss, cWun); 8 exs., Lanjaron, 26.1V.-18.V.1961, leg. Fagel (IRSNB). British Isles: Wales: 1 ex., Powys, Brecon, 26.V1.1994, leg. Owen (cOwe). Sweden: | ex., Halland, Dagsás, 4.1X.1986, leg. Giller- fors (cGil). France: Bourgogne: 2 exs., Cóte d'Or, Montigny sur [illegible] (NHMW). Nord-Pas-de-Calais: 2 exs., Lille (DEI, NHMW). Picardie: 3 exs., Laigneville (Oise), 1.1V.1930, leg. Mequignon (MHNG ); | ex., Laigneville, leg. Mequignon (MHNG); 3 exs., Laigneville, 12.1V.1926 (MHNG); 1 ex., Bailleval (Oise), 24.1V.1927 (MHNG). Ile-de-France: 2 exs., Sucy-en-Brie, leg. Mequignon (MHNG); 1 ex., Sucy-en-Brie, leg. Mequignon (MHNG). Aquitaine: | ex., Arette, 29.111.1960, leg. Jeanne (cBor); 1 ex., Castets, 30.V1.1938, leg. Gaudin (UCBA ); | ex., Gironde, Cambes (MHNG); 3 exs., Cambes, 11.01.1939, leg. Giraud (MHNG); 1 ex., Gironde, loca- lity illegible (MHNG); 4 exs., Bordeaux, bank of Ga- ronne, 16.11.1930, leg. Tempere (MHNG); 1 ex., Gironde, Taussat-les-Bains, 10.1X.1934, leg. Tempere (MHNG); | ex., Gironde, Cubzac-les-Ponts, 23.111.1935, leg. Tem- pere (MHNG); 1 ex., Gironde, Langoiran, 15.1V.1946, leg. Tempere (MHNG). Midi-Pyrénées: 2 exs., Hautes- Pyrénées, locality not specified, leg. Pandellé (NHMW, UCBA); 1 ex., locality illegible, 25.V.1943, leg. Giraud (MHNG). Languedoc-Roussillon: 1 ex., Molitg les Bains, 550 m, 2.1X.2001, leg. Tronquet (cTro); 1 ex., Molitg les Bains, 600 m, oak leaf litter, 10.1V.2002, leg. Tronquet (cTro).7 exs., Carcassonne (NHMW); 1 ex., Cevennes, Villefort, Bresis, 9.111.2002, leg. Af mann (cAss). Auvergne: | ex., Cantal, Thiézac, 28.V111.1947, leg. Tempere (MHNG). Rhone-Alpes: 1 ex., Savoie, Belmont-Tramonet (NHMW); 3 exs., Haute-Savoie, Vougy, 6.1V.1965, leg. Comellini (MHNG, cAss); 1 ex., Ain, Divonne-les-Bains (MHNG); | ex., Ardéche, Mau- ves, 18.111.1906 (cSch); 3 ex., Haute-Savoie, Malagny, 14.X11.1961, leg. Comellini (cAss). Provence: | ex., Var, Roquebrune sur Argens, leg. Demoflys (cTro); l ex., Var, Gapeau inundation [near La Garde], X.1958 (MHNG); 1 ex., Var, Hyeres, X.1958 (MHNG); 1 ex., Var, Vésubie inundation [near Lantosque], V1.1957 (MHNG); 1 ex., same locality, VI.1958 (MHNG); 1 ex., same locality, X1.1947 (MHNG); 1 ex., same locality, Volker ASSING: Western Palaearctic Medon 79 IV.1956 (MHNG); 2 exs., Argens (Le Muy), 11.1937 (MHNG); 1 ex., Var, Dardennes, 11.1956 (MHNG); | ex., Var, E Vidauban, Aille inundation, 1.1955 (MHNG); 2 exs., Var, Loup inundation (W Cagnes), leg. Ochs (MHNG); 8 exs., Loup inundation, 111.1951 (MHNG); 1 ex., Nice, 1.1960 (MHNG); 3 exs., Alpes Maritimes, St.-Vallier-de-Thiey, V.1975, leg. Tou- mayeff (MHNG); | ex. [teneral], same locality, X.1975, leg. Toumayeff (MHNG); 1 ex., Eyguieres, leg. Perrot (MHNG). Alsace: | ex., Strasbourg, Rhine inundation, IX.1954 (MHNG). Corse: 1 ex., S Galeria, Col Palma- rella, 5.V1.1987, leg. Tronquet (cTro); 2 exs., E Ajaccio, Gravone inundation, X.1960 (MHNG). Locality illegible or ambiguous: | ex., Vilue (?), leg. Falcoz (MHNG); 2 exs., Touxadons, Canarive (?), X11.1960 (MHNG); 1 ex., Cussac, VI.1913 (MHNG); 2 exs., IX.1903, leg. Mequignon (MHNG). Germany: Nordrhein-Westfalen: 3 exs., Südlohn, chalk quarry, 21.1V.2001, leg. Scharf (cFel); 1 ex., Hürth, Tagebau Ville, 26.111.1989, leg. Köhler (cKöh); 3 exs., Wesseling, Berzdorf, Kiesgrube, 24.1V.1991, leg. Köhler (cKöh); 4 exs., Vadrup, Emsufer, Hof Schulte- Bisping, 25.V.1997, leg. Köhler (cKöh); 1 ex., Trois- dorf, Kiesgrube Eschaar, 27.1V.1991, leg. Köhler (cKöh). Hamburg: 6 exs., locality not specified (NHMW). Rheinland-Pfalz: 2 exs., Eifel, Cochem, Bengel, 5.V1.1992, leg. Köhler (cKöh); 1 ex., Sinzig, 26.1V.1991, leg. Köhler (cKöh); 2 exs., Schaidt Bien- wald, NWR Stuttpferch, 30.V.1996, leg. Köhler (cKöh); l ex., Wiltingen, car-net, 7.V1.1996, leg. Köhler (cKöh). Bayern: | ex., München, V.1930, leg. Pfaundler (MHNG). Berlin/Brandenburg: | ex., Eisenhütten- stadt, 9.1X.1980, leg. Pütz (cSch); 1 ex., Eisenhütten- stadt, 2.V11.1984, leg. Pütz (cSch); 13 exs., Sperenberg (DEI). Thüringen: 1 ex., locality not specified (NHMW). Sachsen: | ex., Klitten, 30.V.1978, leg. Vo- gel (cWun); 1 ex., Oberlausitz, Daubitz, flood debris, 23.1V.1994, leg. Schülke (cSch); 1 ex., Oberlausitz, Kreis Bautzen, Guttau, edge of pond, 26.V.1985, leg. Schülke (cSch); 2 exs., Guttau, car-net, 25.V.1985, leg. Schülke & Heinig (cSch); 2 exs., Guttau, 22.V.1983, leg. Schülke (cSch). Switzerland: | ex., Morges (NHMW); 3 exs., shore of Lac de Neuchätel, Vaumarcus, 26.V11.1992, leg. Zanettı (cZan); 1 ex., Bressigny (MHNG). Austria: Vorarlberg: 5 exs., locality not specified, leg. Müller (NHMW); 1 ex., Lustenau/Lauterach, 400 m, flood debris, 13.V.1991, leg. Möseneder & Brand- stetter (cSch). Niederösterreich/Wien: 2 exs., Wien (NHMW); 1 ex., Stockerau (NHMW); 1 ex., Klos- ternau (?), leg. Scheerpeltz (NHMW); 1 ex., Lobau, leg. Mandl (NHMW); 1 ex., Ysper, 16.1V.1992, leg. Peschel (cSch). Steiermark: | ex., Graz, leg. Strupi (NHMW). Italy: Trentino-Alto Adige: | ex., Trento, Storo, 28.V.1977 (cAss); 5 exs., Adige (“Etsch-Au”), leg. Breit (NHMW). Lombardia: 2 exs., 10 km E Mantova, Barbassolo, 1X.1976, leg. Carpacchia (cBor); 9 exs., Val Camonica, Cogno, leg. Krüger (NHMW, cAss); | ex., Solferino (MN), 24.X.1971, leg. Cormacchio (cZan); | ek., Sondrio, mole nest, 7.X1.1971, leg. Zanetti (cZan); | ex., Bergamo, Buco d. Corno, 9.V111.1976, leg. Rossi (cZan); l ex., Ostiglia, Palude del Busatello, 12.111.1981, leg. Zanetti (cZan); 2 exs., Torbiere d Iseo (BS), 28.111.1982, leg. Zanetti (cZan); 1 ex., Pozzolengo (BS), 15.1V.1984, leg. Zanetti (cZan); 1 ex., Mantova, Ponti sul Mincio, 27.1.1980, leg. Zanetti (cZan); 1 ex., Monte Marenzo (BG), 200m, 23.11.1997, leg. Tagliapietra & Zanetti (cZan); 5 exs., Pavia, bank of Ticino river, 3.X1.1958, leg. Rosa (UCBA); 2 exs., Cogno near Bienno, leg. Krüger (MHNG). Piemonte: 2 exs., Vercelli, 5.8 12.1V.1964, leg. Olmi (cBor); 6 exs., Leini (TO), mole nest, 8.11.1966, leg. Osella (cBor); 5 exs., Leini, in Salix litter with Lasius fuliginosus, 25.1V.1964 (cZan); 14 exs., Leini, 20.-21.11.1965, leg. Osella (UCBA, cZan); 6 exs., Locana (TO), inundation, X.1960 (MHNG). Liguria: 4 exs., Altare, VIII.1968 & VII- IX.1974, leg. Bordoni (cBor, cTro); 1 ex., Arenzano (GE), 12.X1.1977, leg. Parodi (UCBA); 1 ex., same lo- cality, 20.X.1969, leg. Briganti (UCBA); | ex., same lo- cality, 15.11.1960, leg. Gardini (UCBA); 5 exs., NW Al- benga, Leca, 15.1V.1973, leg. Briganti, Zoia (UCBA); 4 exs., Genova, 1920, leg. Mancini (UCBA). Veneto: 2 exs., NW Monselice, Colli Euganei (NHMW); 7 exs., NW Villafranca di Verona, Custoza, 22.1.1972, leg. Za- netti (cZan); 1 ex., Nogara (VR), Ponte Molino, V.1974, leg. Zanetti (cZan); | ex., Montecchia di C. (VR), mole nest, 22.11.1976, leg. Zanetti (cZan); 1 ex., Gazzo (VR), Palude del Busatello, caricetum, 12.111.1981, leg. Zanet- ti (cZan); 3 exs., Pellegrina (VR), mole nest, 3.11.1980 (cZan); 1 ex., Pellegrina, mole nest, 26.11.1999, leg. Za- netti (cZan); 1 ex., S Vicenza, Fimon, Lago di Fimon, 31.X.1982, leg. Zanetti (cZan); 1 ex., San Martino Buon Albergo (VR), Musella, Ferrazzetta, mole nest, 28.11.1991, leg. Zanetti (cZan); 1 ex., Cavaion Veronese (VR), Palude Canova, 13.11.1994, leg. Zanetti (cZan); 1 ex., Piombino Dese (PD), 20.11.1994, leg. Zanetti (cZan); 1 ex., Feltre (BL), Anzu, 240 m, mole nest, 11.1.1998, leg. Zanetti (cZan); 1 ex., Lovadina near Spresiano, VI.1939, leg. Burlini (MHNG). Friuli- Venezia Giulia: 13 ex., Tagliamento, Fella estuary, 46°22N, 13°07E, 250 m, 12.-13.1X.1998, leg. Schiilke (cSch, cAss); 6 exs., bank of Tagliamento river near Codroigo, 45°57N, 12°55E, flood debris, 12.1X.1998, leg. Schülke (cSch, cAss); 5 exs. Venezia, Tagliamento, Foce fiume, 19.1V.1973 & 20.1V.1990, leg. Zanetti (cZan); 1 ex., bank of Isonzo river, SE Villesse, 4551N, 13°27E, flood debris, 13.1X.1998, leg. Schiilke (cSch); 1 ex., Frisanco (PN), 1.V.1990, leg. Zanetti (cZan). Toscana: 2 exs., “Toscana” (NHMW); l ex., 80 Bonner zoologische Beitráge 54 (2005) Toscana, S Altopascio,Vallino di Macchia (PI), 16.1X.1995, leg. Bordoni (cBor); 1 ex., Mte. Argentario, macchia, 29.11.1921, leg. Moczarski & Scheerpeltz (NHMW). Campania: 2 ex., Capriati al Volturno, 21.V11.1975, leg. Rossi (cZan). Basilicata: | ex., Lago- negro, Rivello Fiume Noce, 40°03N, 15°46E, 250 m, 10.V.2002, leg. Wunderle (cWun); 1 ex., Episcopia (PZ), bank of Sinni, 500m, 22.VII.1993, leg. Zanetti (cZan). Sicilia: | ex., M. Iblei, Noto (SR), F. Manghisi, 25.V11.1994 (UCBA); 1 ex., M. Iblei, Noto, 25.VII.1993 (UCBA); 2 exs., Peloritani, F.ra di Floresta, Santa Ve- nera del Bosco, Frascianida (ME), 450 m, 23.V1.1995, leg. Adorno (UCBA); | ex., M. Iblei, Isola Ardito (near Palazzolo) (SR), Valle d'Ánapo, 5.111.1997 (cAss). Lo- cality ambiguous: 2 exs., Fiume, leg. Meyer (NHMW). Czech Republic: | ex., Bohemia, Doudleby nad Orlicí, 1944, leg. Roubal (MHNG). Polish or Czech territory: 1 ex., Cieszyn (“Teschen”), leg. Wanka (MHNG). Croatia: 2 exs., Istria, Rabac Luka, 15.-17.V.1990, leg. Wrase (cSch). Locality not indicated or illegible: 9 exs. (MHNG). Diagnosis: See ASSING (2004a); habitus as in Fig. 100. Distribution and bionomics: Medon ripicola is an ex- pansive Atlanto-Mediterranean element (Map 12). Based on the revised material and reliable literature re- cords, its distribution extends from the Azores, Madeira, Northwest Africa, and the Iberian Peninsula in the west to southern England and southern Scandinavia in the north and northwest, and to Poland, Romania, Croatia, Bulgaria, and Montenegro in the east and southeast (ma- terial examined; ASSING 2004a). It has also been re- corded from Lithuania (SILFVERBERG 1992). In Scandi- navia, it is known only from Halland (material examined), Malmó, and Oland in southern Sweden (BARANOWSKI 1982; LUNDBERG 1995), as well as from Denmark (HANSEN 1996; HANSEN et al. 1994; V. HAN- SEN 1964). For a selection of additional records see AL- LEN (1969), BERNHAUER (1940), BOHAC (1985), BRUGE et al. (2001), DRUGMAND (1989), FOWLER (1888), GIL- LERFORS (1986), HORION (1965), HUGENTHOBLER (1966), KOCH (1968), KÖHLER & KLAUSNITZER (1998), LUNDBLAD (1958), NORMAND (1935), SCHILLER (1989), TERLUTTER (1995), VOGEL (1979), and WITTWER (1988). According to CICERONI & ZANETTI (1995), the species has been recorded from Sardinia, where the closely related endemic M. sardous occurs, but I have seen no material from there. In the north of its range, M. ripicola was repeatedly found in coastal habitats (ALLEN 1969; BARANOWSKI 1982; HANSEN 1964; HORION 1965). Frequently, it was collected from flood debris (spring and winter) and in other habitats near or at the banks of rivers and streams (material examined; see also HORION (1965), KOCH (1968), and LINKE (1913)). In northern Italy, it is very common and abundant in mole nests, especially near rivers and streams; in this habitat it excludes M. perni- ger, which inhabits mole nests in drier places (OSELLA & ZANETTI 1974). In a very comprehensive study of mole nests in Poland, however, NOWOSAD (1990) did not find a single specimen of M. ripicola. On many oc- casions, the species was sifted from the leaf litter of various kinds of — mostly deciduous — forests and even of macchia. As can be inferred from the examined mate- rial with precise altitude specification, M. ripicola oc- curs at lower and intermediate elevations; none of these specimens was collected above 1000 m. Adult beetles have been observed throughout the year (material examined), though most specimens were found in spring. Flying specimens were caught by car- net in May and June. One examined specimen collected in October was teneral. According to DRUGMAND (1989), the species is bivoltine, but there is no evidence supporting this. Based on the material examined here, it is univoltine. Medon ripicola is the host of the endoparasitic tylenchid nematode Parasitylenchus me- donis Wachek (WACHEK 1955). 3.23. Medon kabylicus sp. n. (Figs. 101-105, Map 12) Types: Holotype ©: Gde Kabylie: Yakouren, forét Be- ni-Ghobri, 800m, V.1953, G. Fagel / G. Fagel det., 1954, Medon ripicola / Holotypus Ö Medon kabylicus sp. n. det. V. Assing 2003 (IRSNB). Paratypes: 230, 39 Y: same data as holotype (IRSNB, cAss); 19, 39 9: Gde Kabylie: Yakouren, ravine au-dessus Tala Teg- zirine, 860m, V.1953, G. Fagel (IRSNB); 599: Gde Kabylie: Yakouren, forét Beni-Ghobri, Bois Sacré, 750m, V.1953, G. Fagel (IRSNB, cAss); 14: Gde Kabylie: forét d’Akfadou, Tala Kitan, 1100m, 18V.1953, G. Fagel (cAss). Description: 3.9-5.5 mm. Facies as in Fig. 101. Aside from the slightly wider protarsomeres I-IV (both sexes) and the male sexual characters, externally indistinguish- able from Medon ripicola. Usual coloration: head blackish brown to black, with the mouth parts lighter; pronotum and elytra rufous to yel- lowish, the elytra usually slightly lighter than the prono- tum; abdomen reddish brown; legs and antennae rufous. Head of subquadrate shape, approximately as wide as long; puncturation very dense, coarse, and areolate; sur- face almost without shine; interstices reduced to narrow ridges; eyes large, approximately 2/3 the length of post- genae in dorsal view. Volker ASSING: Western Palaearctic Medon 8] Map 12: Distributions of Medon ripicola (Kraatz) (filled circles: revised records; open circles: selected literature records) and of M. kabylicus sp. n. (filled squares); the record of M. ripicola from the Azores is omitted. Pronotum approximately as wide as long and at least slightly narrower than head; posteriorly weakly taper- ing; puncturation as in M. ripicola highly variable, rang- ing from fine and indistinct to relative coarse and well- defined. Elytra distinctly wider than and at suture at least about 1.1 times as long as pronotum; puncturation very dense, slightly granulose, and not very well defined. Hind wings apparently fully developed. Protarsomeres I-IV slightly dilated in both sexes; protarsomere II distinctly transverse. Abdomen with very fine and very dense puncturation; posterior margin of tergite VII with palisade fringe. ¢: posterior margin of sternite VII with broad trapezoid excavation (not shaped like a very wide V as in M. ripi- cola), with lateral combs of approximately 6 palisade setae and on either side of middle with tuft of several stout black setae (Fig. 102); sternite VIII similar to that of M. ripicola (Fig. 103); aedeagus very distinctive (Figs. 104 - 105), in lateral view apically wider and straight (in M. ripicola slightly bent dorsad) and in ven- tral apically shallowly incised, not deeply bifid as in M. ripicola. Etymology: The name (adj.) is derived from the name of the Algerian mountain range, where the types were collected. Comparative notes and systematics: Medon kabylicus and M. ripicola are doubtlessly adelphotaxa, as can be 82 Bonner zoologische Beiträge 54 (2005) inferred from the synapomorphic highly derived mor- phology of the aedeagus. Both species can be distin- guished only by the width of the protarsi (slightly di- lated in M. kabylicus), by the morphology and chaetotaxy of the male sternite VII, and by the shape of the distinctive aedeagus. Distribution and bionomics: Medon kabylicus is cur- rently known only from Grande Kabylie, Algeria, where it apparently replaces M. ripicola. As far as is known today, both species seem to have an allo- or parapatric distribution (Map 12). Apart from the altitude (750- 1100 m) and the fact that the types were collected in forests, no further bionomic data are available. 104 105 Figs. 95-105. Medon sardous Dodero (95-99), M. ripicola (Kraatz) (100), and M. kabylicus sp. n. (101-105): forebody (95); male sternite VII (96, 102); male sternite VIII (97, 103); aedeagus in lateral and in ventral view (98-99, 104-105); habitus (100, 101). Scale bars: 95, 100, 101: 1.0 mm; 96-99, 102-105: 0.2 mm. Volker ASSING: Western Palaearctic Medon 83 3.24. Medon indigena (Wollaston, 1857) (Figs. 106-110) Lithocharis indigena Wollaston, 1857 (WOLLASTON 1857: 93). Material examined (total: 133 exs.): Madeira: Porto Santo: 11 exs., Pico do Castelo, 400m, mixed laurel and pine wood, 1.1V.1993, leg. Assing, Wunderle (cAss, cWun): | ex., Pico do Castelo, 300- 430m, 11.1X.1998, leg. Schuh (cAss); 105 exs., Pico Juliana, 400m, mixed laurel and pine wood, 1.1V.1993, leg. Assing, Wunderle (cAss, cWun); 12 exs., same lo- calityay I“IV.1996, leg. Lompe, Zerche (DEL, cAss, cSch); 4 exs., Pico Facho, 500m, pine wood with scat- tered laurel trees, 1.1V.1996, leg. Assing (cAss). Diagnosis: 4.2-5.0 mm. Habitus as in Fig. 106. Colora- tion variable, castaneous with blackish abdomen to blackish with slightly lighter elytra; legs light brown; antennae of variable coloration, light brown to dark brown, often with the central antennomeres slightly darker than the basal and apical ones. Head weakly oblong, approximately 1.1 times as long as wide (Fig. 107); eyes small, less than half the length of postocular region in dorsal view and not projecting from lateral outline of head; puncturation distinct, relatively sparse, non-areolate, and rather fine; interstices on aver- age wider than punctures; microsculpture usually absent from central dorsal area, visible only in posterior and lateral areas. Pronotum approximately as wide as head or slightly nar- rower, weakly oblong; puncturation non-areolate, finer and denser than that of head, interstices narrower than punctures; microsculpture shallow, present at least in lateral areas of pronotum (Fig. 107). Elytra small, only slightly wider than pronotum and at suture only approximately 0.8 times as long as prono- tum; puncturation densely coriaceous; surface almost without shine (Fig. 107). Hind wings reduced. Abdomen with very fine and dense puncturation, with distinct microsculpture, and with subdued shine; poste- rior margin of tergite VII without palisade fringe. Pro- tarsomeres dilated in both sexes, with weak sexual di- morphism. ¢: protarsomeres | - IV slightly more dilated than in 9; posterior margin of sternite VII very weakly concave, in the middle with fringe of weakly modified, long black setae (Fig. 108); sternite VIII posteriorly with deep emargination; aedeagus very distinctive (Figs. 109-110). Comparative notes: This highly distinctive species is readily distinguished from its Western Palaearctic con- geners by the sparse, fine, and non-areolate puncturation of the head, the reduced wings, the coriaceous elytra, and especially by the male sexual characters: the ab- sence of palisade setae or distinctly modified setae at the almost truncate posterior margin. of sternite VII, as well as the morphology of the aedeagus. / Comments: The types were not examined, but in view of the distinctive external morphology, sexual charac- ters, and the restricted distribution, there are no doubts regarding the identity and interpretation of the species. Based on the male primary and secondary sexual char- acters, M. indigena is most closely related to M. feloi Assing from the Canary Islands (see below). A closer relationship to the Madeiran endemic M. vicentensis Serrano, in contrast, is unlikely, based on the differ- ences in the shape and chaetotaxy of the male sternite VU. Distribution and bionomics: Medon indigena is en- demic to the Madeiran archipelago and has been re- corded from both Madeira proper and Porto Santo. While there are no recent records from the former, the species has repeatedly been found in Porto Santo in the past decade. Numerous specimens were sifted from leaf litter and rotting wood on the northern slopes of Pico Juliana, Pico do Castelo, and Pico Facho. 3.25. Medon feloi Assing, 1998 (Fig. 111) Medon feloi Assing, 1998 (ASSING 1998: 143f). Type examined: See ASSING (1998). Diagnosis: For a more detailed description see ASSING (1998). 4.1 mm. Coloration of whole body uniformly ferrugine- ous. Head large (Fig. 111), weakly oblong; eyes reduced to minute rudiments, without ommatidia; puncturation fine and relatively sparse, interstices much wider than punctures; surface with pronounced microreticulation and almost matt. Pronotum distinctly narrower than head; weakly, but no- ticeably oblong; puncturation very fine and sparse, barely noticeable; microreticulation pronounced (Fig. 111); Elytra widest near posterior margin, anterior external angles almost obsolete; only slightly wider and at suture somewhat shorter than pronotum; puncturation more distinct and microsculpture much weaker than on head and pronotum; dorsal surface with distinct shine. Hind wings completely reduced. Abdomen with moderately dense and very fine punctu- ration; posterior margin of tergite VII without palisade fringe. 84 Bonner zoologische Beitráge 54 (2005) ©: posterior margin of sternite VII of similar shape and chaetotaxy as in M. indigena; aedeagus as illustrated by ASSING (1998). Comparative notes: This highly distinctive cave- dwelling species is readily distinguished from other Western Palaearctic congeners by the completely re- duced eyes, the reduced pigmentation and wings, and by the male sexual characters. The latter are highly similar to those of M. indigena, its presumable sister species. The only other species with (almost) completely re- duced eyes known from the Western Palaearctic region are M. vicentensis Serrano from Madeira and M. antri- cola from El Hierro. Distribution and bionomics: Medon feloi is a true troglobite and endemic to La Palma, Canary Islands. 3.26. Medon subterraneus Coiffait, 1970 Medon subterraneum Coiffait, 1970a (COIFFAIT 1970a: 7036). Comments: The holotype of this species, whose de- seription is based on a single male from Ibiza without indication of the type depository, was looked for, but not found in the Coiffait collection and in the Zoologi- cal Museum in Barcelona. COIFFAIT (1970c) corrected the locality data, but failed to specify the whereabouts of the holotype. Until this specimen becomes available, an interpretation of the species must rely on the original description; additional material has not become known. According to COIFFAIT (1970a), M. subterraneus is characterized especially by adaptive reductions resulting from a subterranean habitat and by the male sexual characters. The eyes are only one fourth the length of the postocular region, the hind wings are of reduced length, the abdominal tergite VII lacks a palisade fringe, and the posterior margin of the male sternite VII is shal- lowly concave and has a row of sparse, weakly modified marginal setae; the aedeagus is of similar general mor- phology as that of M. dilutus, M. pocofer, and related species. 4. KEY TO THE MEDON SPECIES OF THE WESTERN MEDITERRANEAN In the key below, M. subterraneus (described from Spain: Ibiza) and M. spelaeus (southeastern Spain) are not accounted for, since no material has become available for study. Medon augur from Corsica 1s tentatively included in the key, although its male sexual characters are un- known. Based on external characters, this doubtful spe- cies is close to M. rufiventris and M. dilutus, so that even if it should prove to be a valid species, it can be expected to have similar male secondary sexual characters. Eyes strongly to almost completely reduced, at most 1/5 the length of postocular region in dorsal view, without pigmentation or ommatidia. Atlantic Islands. Eyes at least 1/4 the length of postocular region, composed of at least some ommatidia. ................... > Large species of at least 5 mm. Eye less strongly re- duced, approximately 1/5 the length of temples in dor- sal view; head distinctly oblong and posteriorly ta- pering. Legs and antennae very long and slender. ©: posterior margin of sternite VII as in Fig. 42, with a tuft of long black setae on either side of middle; aedeagus as in Figs. 44-45. Endemic to El Hierro, Canary Islands; troglobite. .......... M. antricola sp. n. Eyes reduced to minute rudiments. Smaller species. Sexual characters different. ....coinn e 3 Head about as wide as long (Fig. 71). Antennae long and slender; antennomere VII distinctly oblong (Fig. 72). Pronotum approximately as wide as long, dis- tinctly tapering caudad, and with shallow, but rather well-defined and distinct puncturation (Fig. 71). Pro- tarsomeres I - IV dilated in both sexes. Elytra shorter. Posterior margin of Ö sternite VII distinctly concave and with two combs of palisade setae; aedeagus: SERRANO (1993). Madeira: Sao Vicente. o A M. vicentensis Serrano Head noticeably oblong (Fig. 111). Antennae shorter; antennomere VII not oblong. Pronotum distinctly oblong, weakly tapering caudad, and with extremely fine, barely noticeable puncturation (Fig. 111). Pro- tarsomeres not distinctly dilated. Elytra longer. Pos- terior margin of Í sternite VII almost truncate, with two clusters of long black setae in the middle, but without palisade setae; aedeagus: ASSING (1998). Canary Islands: La Palma. ............... M. feloi Assing Posterior margin of @ sternite VII with more or less pronounced clusters of long black setae, but without combs of palisade setae, „nn. er 5 Posterior margin of Ö sternite VII with two combs Of palisade selae. „nen 9 Head shining, with sparse, well-defined, non- areolate puncturation (Fig. 107). Elytra at suture dis- tinctly shorter than pronotum (Fig. 107); hind wings reduced. Abdominal tergite VII without palisade fringe. ©: posterior margin of sternite VII truncate to weakly concave, with a cluster of rather sparse black setae on either side of middle (Fig. 108); aedeagus broadly truncate apically (Figs. 109-110). Endemic to Madeira proper and Porto Santo. ....................... a ere he M. indigena (Wollaston) Volker ASSING: Western Palaearctic Medon 85 Head matt or nearly so due to very dense punctura- tion and/or pronounced microsculpture. Elytra at su- ture longer than pronotum; hind wings not reduced. Abdominal tergite VII with palisade fringe. ©: pos- terior margin of sternite VII at least weakly bisinu- ate, with much more numerous black setae, and, in addition to lateral clusters, also with central cluster of black setae; aedeagus with bifid or with narrowly elongated apex (ventral view). The M. apicalis A ew sacs aussen ts niiisideobisscnrasuncorscateesssasccssnenspe 6 Head with shallower and less coarse puncturation (Fig. 13). Puncturation of pronotum very fine, barely noticeable. Pronotum almost completely without shine due to pronounced microsculpture (Fig. 13). &: posterior margin of sternite VII with rather few and very slender black setae (Fig. 14); aedeagus with very long and slender ventral process of distinctive shape (Figs. 15-16). Canary Islands: Gran Canaria. . M. oromii sp. n. Head with coarser puncturation. Puncturation of pronotum fine and often shallow, but dense and dis- tinct. Pronotum with at least subdued shine. @: pos- terior margin of sternite VII in the middle with more numerous and stouter black setae (e.g. Fig. 3); with much shorter and apically bifid ventral process. .... 7 Head on average with slightly more distinct punctu- ration (Fig. 1). $: posterior margin of sternite VII in the middle with distinct convex projection (1.e. dis- tinctly bisinuate) and, especially in the middle, with darker, stouter, and longer modified setae (Figs. 2- 3); aedeagus with ventral process of distinctive mor- phology: deeply bifid, apices widely separated, and subapically dentate (Figs. 5-7). Italy, southern Swit- zerland, southeastern France (Map 3). .....................- RES Le nstieaanknsacslauhlssbedenu M. perniger Coiffait Puncturation of head usually ill-defined and conflu- ent. Í: posterior margin of sternite VII in the middle weakly convex (i.e. more weakly bisinuate) or con- cave, with shorter and more slender modified setae. d: posterior margin of sternite VII in the middle weakly convex; aedeagus compact and apically bifid (ASSING 2004a, figs. 1-3). Widespread in the West- ern Palaearctic region, from the Atlantic Islands to Croatia and Bosnia-Herzegovina in the southeast IMA 22) IN M. apicalis (Kraatz) Í: posterior margin of sternite VII in the middle weakly concave (Fig. 10); aedeagus with very long and apically acute ventral process (Figs. 11-12). N- Algeria, NW-Tunisia (Map 2) en ENS eres DA M. sericellus (Fairmaire) 9: Posterior margin of % sternite VII with very deep and broad, trapezoid or concave emargination, with two combs of palisade setae, and, except for one species (M. ripicola), additionally with numerous long black setae on either side of the middle, usually grouped in more or less distinct tufts. Protarsi At most weakly dilated and without sexual dimorphism. Aedeagus either with apically truncate or with dis- tinctly elongated ventral process of highly distinc- tive shape. Elytra distinctly wider and at suture longer than pronotum. The M. fusculus species TOW) Fare ORO 10 Posterior margin of Ö sternite VII with shallower excavation and — except for one species from the Canary Islands — without (tufts of) long black setae on either side of the middle; sternite VII usually with smaller emargination posteriorly. ................. 13 10. Posterior margin of Ö sternite VII with deep excava- tion of distinctly trapezoid shape and on either side of middle with long thin setae not arranged in dis- tinct tufts. Aedeagus apically truncate (ASSING 2004a, figs. 66-71). Widespread species; distribution ranging from Great Britain to the Middle East, but absent from the southwestern Mediteranean (Map Dal) A ÓN M. fusculus (Mannerheim) Posterior excavation of Í sternite VII either shal- lower or not distinctly trapezoid, on either side of middle either without or with tufts of stout black se- tae. Aedeagus with apically distinctly elongated ven- GA process nenne ee 11 .Larger and on average darker species (Fig. 95), forebody brown to dark brown; head only slightly darker than pronotum. Posterior margin of Ö sternite VII as in Fig. 96. Aedeagus as in Figs. 98-99. En- demic 10 Sardinia, 4... M. sardous Dodero Smaller and more distinctly bicoloured species, pronotum usually rufous and distinctly contrasting with the blackish or blackish brown head. © sexual Characters PCr Nts: nern 12 . Protarsomeres I-IV not dilated, protarsomere II ap- proximately as wide as long; posterior margin of sternite VII in the middle obtusely angled, on either side of middle without tufts of stout black setae; aedeagus distinctive (ASSING 2004a, figs. 118-120), apically in ventral view narrowly and deeply incised. Widespread from NW-Africa, the Canary Islands and the Azores to southern Scandinavia, Lithuania, Poland, Bulgaria, and Montenegro (Map 12). ........... M. ripicola (Kraatz) Protarsomeres I-IV slightly dilated, protarsomere II much wider than long; posterior margin of sternite VII with trapezoid excavation, in the middle trun- 14. Bonner zoologische Beitráge 54 (2005) cate, on either side of middle with a tuft of stout black setae (Fig. 102); aedeagus distinctive (Figs. 104-105), apically in ventral view without narrow deep incision. Algeria (Map 12). M. kabylicus sp. n. $: protarsi I-IV unmodified, in one species weakly dilated; aedeagus apically rounded, truncate, or with obtuse or rounded apico-lateral angles. ................ 14 “: protarsi I-IV more or less strongly dilated; aedeagus apically either incised or with the apical and the lateral margins meeting at an acute angle; ventral process not strongly elongated. ................ 18 Eyes very small, about one fourth the length of tem- ples in dorsal view (Fig. 90). ©: aedeagus as in Figs. 92-94. Known only from one locality in northern POI Nay. A A M. mirei Coiffait Eyes larger, more than one third the length of tem- ples ın dorsal view. Aedeagus different. More wide- SPEER DES dnd 15 .Head with finer puncturation (Figs. 74-75). Head and pronotum usually blackish. ©: aedeagus as in Figs. 77-78. From the south of the Iberian Peninsula to southern England and Central Europe. Absent from NW-Africa, the Balkans, and apparently also from Austria and from central and southern Italy (Maps mea M. piceus (Kraatz) Head with very coarse puncturation. Head and pronotum usually (but not always!) rufous to dark brown. Ö: aedeagus of different shape. ................ 16 . Of larger average size and often with relatively lar- ger head (though these characters are subject to con- siderable intraspecific variation). Elytra usually shorter, at suture approximately as long as prono- tum. ¢: aedeagus larger (ASSING 2004a, figs. 40- 41). From the Bosporus, Greece, and southern mainland Italy in the south and southeast to southern Scandinavia, southern England, western France in the north and west (Map 8). Distribution in the Ibe- rian Peninsula limited to the northeast (Cataluña). ... M. brunneus (Erichson) On average smaller and with relatively smaller head. Elytra at suture usually at least slightly longer than pronotum, only in brachypterous morph of M. despectus as long as pronotum. @: aedeagus smaller and of different shape. NW-Africa, Iberian Penin- sila, 31 SW-REINEE. nenn 17 “: ventral process of aedeagus of trapezoid shape in ventral view (Figs. 82-84). Distribution: Morocco to SW-Fralce nern M. cauchoisi Jarrige <: ventral process of aedeagus apically rounded in ventral view (Figs. 88-89). Confirmed records only from Tunisia and Algeria. M. despectus (Fairmaire) 18. Y Head and'pronotum with conspicuously coarse and well-defined puncturation; punctures of head areo- late and very dense, interstices reduced to narrow ridges. Abdomen with rather sparse puncturation. ©: aedeagus as figured by ASSING (2004a). Predomi- nantly Eastern Mediterranean species, ranging from eastern Anatolia to southeastern Austria, the Czech Republic, and Slovakia. . M. ferrugineus (Erichson) Head and pronotum with finer puncturation; punc- tures of head not areolate. Abdomen with very dense and fine puncturation. ¢: aedeagus of different SHAPE: 2.:2....u ee 19 . Very large species, body length usually >6.0 mm. Legs and antennae very slender; metatarsi almost as long as metatibiae. Sexual dimorphism of protarsi distinct, protarsomeres I - IV more strongly dilated in male than in female. @: aedeagus very long and A oo 20 Smaller species, body length in normal preparation usually <6.0 mm. Legs and antennae not conspicu- ously long and slender; metatarsi much shorter than metabiae. Í: aedeagus much shorter and less slen- O O odio 21 .Largest species in the Western Palaearctic region, 7.0-9.0 mm (Fig. 53). Coloration uniformly ferrugi- neous. Head very large and of ellipsoid shape, lateral margins convex and posterior angles almost obsolete (Fig. 54). Length of metatibia 1.7 - 1.8 mm. ¢: ster- nite VII more oblong and posterior concavity shal- lower in relation to length of sternite (Fig. 55); aedeagus longer and with longer and more slender ventral process (Figs. 56-57). Known only from caves near Madrid. ......... M. procerus (Perez Arcas) Usually smaller, 6.0-8.0 mm (Fig. 46). Head, most of pronotum, and anterior segments of abdomen blackish. Head of subquadrangular shape and with more distinct posterior angles (Fig. 47). Length of metatibia at most 1.4 mm. @: sternite VII less ob- long and posterior concavity relatively deeper (Fig. 48); aedeagus shorter and with less slender ventral process (Fig. 49-52). Widespread from the south of the Iberian Peninsula to southern England, southern Scandinavia, and Central Europe (Map 6). ............... Ek dis wae ah Racha datas ee M. castaneus (Gravenhorst) . Protarsomeres I - IV strongly dilated in both sexes (Fig. 59); meso- and metatarsomeres III and IV at least weakly transverse and almost cordiform. Head relatively large, weakly transverse, and somewhat flattened (Fig. 58). Í: sternite VII and aedeagus as figured by ASSING (2004a). Confined to the coasts (under shingles and sea-weed) of western Europe, eastwards to Montenegro. ........ M. pocofer (Peyron) Volker ASSING: Western Palaearctic Medon RT — Protarsomeres I - IV less strongly dilated; meso- and metatarsomeres HI and IV oblong and not cordi- form. Head not flattened. ¢: aedeagus of different shape. Species not usually occurring in coastal habi- ada 22.Head distinctly oblong (Figs. 33-34); head and pronotum with pronounced microsculpture and very fine puncturation (Fig. 34). ©: posterior margin of sternite VII with distinct tufts of long dark setae on either side of middle (Fig. 35); aedeagus as in Figs. 86-37. Endemic to Canary Islands. ..............-...........: A SEES Soon nenasese- M. subcoriaceus (Wollaston) — Head at most weakly oblong; head and pronotum usually with very shallow microsculpture and with coarser puncturation; exceptionally the microsculp- ture may be pronounced. ©: posterior margin of sternite VII without distinct tufts of long dark setae on either side of middle; aedeagus of different shape. A A 23 23. Puncturation of head coarser, in dorso-median area sparser than in lateral areas. On average smaller spe- ASIN A 24 — Puncturation of head finer and evenly distributed, in dorso-median area as dense as in lateral areas. On average larger species, 4.0 - 6.0 mm. Aedeagus as in LAS ee sere see A O 26 24. Large species of about 5.0 mm. Eyes smaller, weakly prominent and less than one third the length of postocular region in dorsal view (Fig. 73). Corsica. . ooo edad M. augur Fauvel — Smaller species of 3.5-4.5 mm. Eyes larger, dis- tinctly projecting from lateral outline of head and about half the length of postocular region in dorsal A Ramee meee ee NE IE TEFRRRENE 25 25. Head on average more transverse and often slightly dilated posteriorly. 4: aedeagus with apical incision (ASSING 2004a, figs. 57-58). Widespread, from Cau- casus region and Anatolia to Spain; confirmed re- cords from NW Africa absent (Map 7). .................... SSCS SUSU a M. rufiventris (Nordmann) — Head on average less transverse and usually not di- lated posteriorly (Figs. 62-65). 4: aedeagus without apical incision (Figs. 67-69). Known only from Tu- nisia and Algeria (Map 7). .... M. africanus (Fauvel) 26. Coloration of body more or less uniformly rufous to ferrugineous. Eyes slightly smaller. Elytra at suture approximately as long as pronotum (Figs. 17-21, 25- 26). Unknown from North Africa and the Atlantic NS Feat hs Spore eee a ceo e eesarsaner 2 — Head, usually at least the central part of the prono- tum, and abdominal segments III - VII blackish. Eyes slightly larger and more prominent. Elytra at suture distinctly longer than pronotum (Figs. 30-32). Azores, Canary Islands, NW-Africa, Sicily (Maps 4, 5). ee een M. dilutus quadriceps (Wollaston) 27.8: apex of aedeagus (ventral view) with subparallel lateral margins, pronounced apico-lateral angles, and with small median incision (Figs. 27-29). Southern mainland Italy and Central Europe. Unconfirmed re- cords also from southern Scandinavia (Maps 4, 5). .. ee ee Eee M. dilutus dilutus (Erichson) — : apex of aedeagus (ventral view) with convex lat- eral margins, indistinct apico-lateral angles, and with large median incision (Figs. 22-23). Southwestern Europe, Sardinia, southern England (Maps 4, 5). ...... REES aiii Sacos M. dilutus cephalus Koch 5. THE MEDON SPECIES OF THE EASTERN MEDITERRANEAN AND ADJACENT REGIONS: ADDENDA Below, the material that has become available since the submission of the first part of the revision is listed. The records are commented on only when they are outside the known range of the species (see ASSING 2004a, 2004b). Medon maronitus (Saulcy, 1864) Additional material examined: Macedonia: 6 exs., Vardar plain, leg. Schatzmayr (NHMW, cAss). Israel: 12 exs., Galilee, Safad, 500 m, 30.V.1973, leg. Lóbl (MHNG, cAss); 5 exs., Galilee, Mt. Meron, 900 m, 27.V.1973, leg. Lóbl (MHNG, cAss). Azerbaijan: 2 exs., Talysh mts., road Lenkoran- .Lerik km 14-18, 30.1V.-9.V.2001, leg. Lackner (cAss); 1 ex., Talysh mts., Masalli — Isti Su, 10.-13.V.2001, leg. Lackner (cAss). Medon bucharicus Bernhauer, 1902 (Figs. 112-116) Medon bucharicus Bernhauer, 1902 (BERNHAUER 1902: 244). Types examined: Lectotype Í, present designation: ¢ / Buchara Staudinger / bucharicus Brh. Centralasien Type / Dr. M. Bernhauer donavit 29.1V.1942 / ex coll. Scheer- peltz / Typus Medon bucharicus Dr. Bernhauer / Lecto- typus Medon bucharicus Bernhauer desig. V. Assing 2003 (NHMW). Paralectotypes: 14: same data as lecto- type (NHMW); 14: same data, but “Q / ... / Cotypus Medon bucharicus ...” (NHMW). Additional material examined: Tajikistan: 2 exs., Pamir Alai, Hissar Mts., Adshuk- cleft near Warsob, 1200 m, 1.-3.VII.1990, Schiilke & Wrase (cSch); 1 ex., Hissar Mts., Warsob Valley, Siddi, 2000 m, 29.VI.1990, leg. Schülke & Wrase (cAss); 88 Bonner zoologische Beitráge 54 (2005) l ex., Seravshan Mts., locality illegible, 1. VI. 1984, leg. Michailov (cSch). Kirghizia: 7 exs., Ferganskiy Alatau, Yarodar, 1400-1500 m, 16.-19.V.1993, leg. Schawaller (SMNS, cAss). Comments: The original description is based on an un- specified number of syntypes. In order to secure the pre- sent interpretation of the species, one of the syntypes in the collections of the NHMW is here designated as lec- totype. Medon bucharicus was previously attributed to Hy- pomedon e.g. by COIFFAIT (1984), who, on p. 15 of the same work, however, mentions the same species as Me- don in a footnote. As can be inferred especially from the morphology of the male sexual characters, the species doubtlessly belongs to the Medon apicalis group. Diagnosis: 3.6-4.4 mm. Habitus as in Fig. 112. Head and pronotum castaneous to dark brown, head some- times slightly darker than pronotum; elytra rufous to brown; abdomen brown to dark brown with lighter apex; legs and antennae rufous to brown. Coloration somewhat variable; usual coloration: casta- neous to blackish brown, with the head usually slightly darker than pronotum and elytra; legs and antennae tes- taceous yellowish brown, with the basal antennomeres usually reddish brown to ferrugineous. Head 1.10 — 1.15 times as long as wide (Fig. 113); eyes of variable size, slightly more than half the length of postocular region in dorsal view, or much smaller, ca. one third the length of postocular region; integument with distinct microreticulation and with moderately dense, relatively shallow, but relatively large punctures. Antennae slender, preapical antennomeres only indis- tinctly transverse. Pronotum weakly oblong, approximately as wide as head; microsculpture similar to that of head, but punctu- ration much finer, sparser, shallower, and ill-defined (Fig. 113). Elytra at suture slightly longer (large-eyed specimens) (Fig. 112) or shorter (small-eyed specimens) than pronotum; puncturation dense and somewhat granulose; microsculpture indistinct, surface with much more shine than head and pronotum. Legs relatively slender; metatarsi rather long, all tar- someres much longer than wide, second tarsomere about 2.5 - 3.0 times as long as wide. Abdomen with fine and moderately dense puncturation and with pronounced microsculpture; posterior margin of tergite VII with palisade fringe. ¢: posterior margin of sternite VII broadly and shal- lowly concave, and with cluster of modified black setae (Fig. 114); sternite VIII posteriorly broadly and deeply excavate; aedeagus of similar morphology as in M. ma- ronitus, but larger, broader (ventral view), and in lateral view apically more distinctly dentate (Figs. 115-116). Comparative notes: Medon bucharicus is somewhat similar to M. maronitus, but distinguished by larger body size, more uniform coloration of head and prono- tum (in M. maronitus, the head is usually much darker than the pronotum), less dense puncturation of the head, clearly longer antennae, a much more strongly mi- crosculpured and less densely punctured pronotum, rela- tively shorter elytra, longer tarsi (in M. maronitus the second metatarsomere is less than twice as long as wide), a different shape and chaetotaxy of the male ster- nite VII, a much deeper posterior emargination of the male sternite VIII, and a larger and broader aedeagus. From M. perniger and M. apicalis, M. bucharicus is readily distinguished by less dense and less coarse punc- turation of the head, by more pronounced microsculp- ture of the pronotum, by relatively smaller elytra, shorter metatarsi, and by different male sexual charac- ters. Intraspecific variation: Apparently, there are two morphs: the type specimens from Uzbekistan have rela- tively large eyes (more than half the length of postocu- lar region in dorsal view) and longer elytra (Figs. 112- 113), whereas the material seen from Tajikistan has small and weakly prominent eyes (about one third of the length of postocular region) and shorter elytra. How- ever, the male primary and secondary sexual characters are essentially the same, so that the observed differences are attributed to intra- rather then interspecific variation. Distribution and bionomics: Medon bucharicus has become known only from Tajikistan, Khirghizia, and Uzbekistan in Middle Asia. The specimens examined were collected at altitudes of 1200-2000 m during the period from May through July. Medon cerrutii Coiffait, 1976 Additional material examined: Crete: 6 exs., Zakros, 28.11.1973, leg. Meybohm & Fül- scher (MHNG); 1 ex., Zakros, 20.11.1975, leg. Mey- bohm & Fiilscher (MHNG). Medon subquadratus Assing, 2004 Additional material examined: Turkey: Adana: 2 exs., Kozan, 300 m, 5.V.1967, leg. Besuchet (MHNG, cAss). Volker ASSING: Western Palaearctic Medon 89 112 eae A NN Dipl Nodes 108 110 25 116 Figs. 106-116. Medon indigena (Wollaston) (106-110), M. feloi Assing, holotype (111), and M. bucharicus Bernhauer (112-116; 112, 113: lectotype): habitus (106, 112); forebody (107, 111); male sternite VII (108, 114); aedeagus in lateral and in ventral view (109-110, 115-116); head and pronotum (113). Scale bars: 106-107, 111-113: 1.0 mm; 108-110, 114-116: 0.2 mm. Medon guignoti Coiffait, 1987 Medon semiobscurus (Fauvel, 1875) Additional material examined: Medon ruber Sahlberg, 1908 (SAHLBERG 1908: 33f) syn. n. Greece: 14 exs., Serrai, Hagiai Ioannis, wet moss, 17.V111.1964, leg. Puthz (MHNG, cAss). Medon sahlbergi Scheerpeltz, 1933 (SCHEERPELTZ 1933: 1255) [replacement name for Medon ruber Sahl- berg] syn. n. 90 Bonner zoologische Beitráge 54 (2005) Types examined: Medon ruber: Lectotype Ö, here des- ignated: Galilea / Vall. Kison / J. Sahlb. / Spec. typ. / 2410 / Mus. Zool. H:fors Spec. typ. No. 875 Medon ru- ber J. Sb. / Lectotypus Medon ruber Sahlberg desig. V. Assing 2003 / Medon semiobscurus (Fauvel) det. V. Assing 2003 (ZMH). Paralectotype +: same data as lec- totype (ZMH). Additional material examined: Turkey: 14 exs., Mugla, Marmaris, V.1969, leg. Fagel (IRSNB); 4 exs., Antalya, Sugözü near Alanya, 15.11.2000, leg. Esser (cEss); 1 ex., Antalya, Akseki, 14.111.2000, leg. Esser (cEss); 1 ex., Antalya, Taskesigi, 17.111.2000, leg. Esser (cEss); 1 ex., Antalya, Side, 13.111.2000, leg. Esser (cEss); 2 exs., Antalya, Killik, cave entrance, 780 m, 2.-4.V1.2003, leg. Lohaj (cSch): 11 exs., Kilik, 800 m, 3.V1.2003, leg. Smatana (cSch, cAss). Syria: | ex., Caifa (?), leg. Simon (NHMW) Israel: | ex., Carmel ridge, Dalya/Galed, 19./30.111.1995, leg. Sama (cZan); 3 exs. Mt. Carmel, Little Switzerland, 28.V.1973, leg. Lóbl (MHNG). Iran: | (NHMW). ex. [macropterous], Central Iran, Shapoor Comments: The original description of M. ruber Sahl- berg — a preoccupied name later replaced by M. sahl- bergi Scheerpeltz, 1933 — is based on an unspecified number of syntypes. In order to secure the present inter- pretation of the name, the male syntype in the collec- tions of the ZMH is here designated as the lectotype. The two type specimens belong to the short-winged, microsculptured and light-coloured morph of M. semiobscurus (see ASSING 2004a), so that M. ruber and its replacement name M. sahlbergi are synonymized with the senior name M. semiobscurus (Fauvel). Distribution: Medon semiobscurus is here recorded from Syria and Iran for the first time. Medon dilutus pythonissa (Saulcy, 1864) Medon pythonissa (Saulcy, 1864); ASSING (2004a). Additional material examined: Greece: Thrakia: 3 exs.. Nomós Xanthi, Nestos near Toxótes, 25.1V.1994, leg. Schawaller (SMNS, cAss). Turkey: 4 exs., Mersin, Tönük, 380m, 8.VII.1987, leg. Gardini, Rizzo, Zoila (cZan); 5 exs., Balikesir, Ayvalik, 15.V11.1969, leg. Besuchet (MHNG); 1 ex., Istanbul (NHMW); 2 exs., Antalya, Sugózú near Alanya, 15.11.2000, leg. Esser (cEss); 2 exs., Antalya, Güclüköy near Akseki, 14.11.2000, leg. Esser (cEss); 3 exs., Anta- lya, Kilik, 800 m, 3.V1.2003, leg. Smatana (cSch, cAss). Ukraine: | ex., Crimea, Jalta (NHMW ); 2 exs., Odessa, Lusanwskij forest, rotting leaves, 22.X. 2003, leg. Gon- tarenko (cAss). Jordan: | ex., Amman, 800 m, 25.111.1958, leg. Klap- perich (MHNG). Caucasus region: | ex., “Kaukas” (NHMW). This taxon was previously referred to as M. pythonissa by ASSING (2004a), but is here regarded as a subspecies of M. dilutus. For further details see Section 3.5. Medon ferrugineus (Erichson, 1837) Additional material examined: Austria: 2 exs., Niederósterreich, Mistelbach, oak forest, 11.V.2002, leg. Zanetti (cZan); 3 exs., Bad Deutsch Altenburg, leg. Blühweiss (MHNG, cAss). Locality not identified: 3 exs., “Hackelsberg” (MHNG). Slovakia: 1 ex., Stúrovo, VI.1963, leg. Rous (cBor); 3 exs., Kovácova, leg. Roubal (MHNG); 2 exs., Nitra, Zobor, 27.1X.1980, leg. Moravec (cKGh); 2 exs., Bratis- lava, Svaty jur, 220 m, 26.VI.1991, leg. Behne (DEI, cAss); | ex., Plast’ovce, 14.1V.2000, leg. Hlavác (cAss). Romania: | ex., Comana Vlasca, leg. Montadon (MHNG ); 2 exs., Cheile Turzli, 20.1V.1962, leg. Comel- lini (MHNG. cAss). Bulgaria: | ex., Stara planina, VI.1968, leg. Rous (Bor). Macedonia: | ex., Keretschkoi, leg. Schatzmayr (MHNG); 1 ex., Vardar plain, leg. Schatzmayr (MHNG). Greece: | ex., Thessaloniki, leg. Schatzmayr (MHNG). Medon beroni Coiffait, 1969 Additional material examined: Greece, Crete: | ex. [aedeagus teratological], W-Crete, Rethimnon, Dramia, bank of Mouselas river, [V.1997, leg. Feldmann (cFel). Medon lindbergi Scheerpeltz, 1958 Additional material examined: Israel: 49 exs. [partly teneral], Mt. Carmel, Little Swit- zerland, 28.V.1973, leg. Lóbl (MHNG, cAss); 9 exs., Galilee, below Safad, 500 m, 30.V.£14.V1.1973, leg. Lóbl (MHNG, cAss): 1 ex., Galilee, Ginosar, 20.- 21.V.1973, leg. Löbl (MHNG); 3 exs., 3 km E Ginosar, 24.V.1973, leg. Löbl (MHNG). Volker ASSING: Western Palaearctic Medon 9] Medon fusculoides Coiffait, 1969 Additional material examined: Iran: | ex., Fars, Sivand, 30°07N, 52°58E, 10.V1.1974, leg. Senglet (MHNG); 19 [identification uncertain], Kohkiluyeh, Charam, 30°44N, 50°44E, 23.V.1974. leg. A. Senglet (MHNG). Medon lanugo Assing, 2004 Additional material examined: Turkey: Antalya: | ex., ca. 30 km N Akseki, Bademlh, Bademli gecidi, 2.-3.V.2001, leg. Lohaj (cAss). Amasya: 3 exs., Amasya-Ezinepazari, 22.V.1967, leg. Besuchet (MHNG). The species was previously unknown from Amasya province (ASSING 2004a). Medon sparsiventris Eppelsheim, 1889 Additional material examined: Azerbaijan: 2 exs., Masally, Istisu W Masally, 300 m, 18.-19.V1.1996, leg. Schawaller (SMNS). 6. THE MEDON SPECIES OF THE EASTERN MEDITERRANEAN AND ADJACENT REGIONS: CORRIGENDA The recently published revision of the Medon species of the Eastern Mediterranean and adjacent region (ASSING 2004a) is unfortunately based on an early draft of the manuscript and not a revised version submitted to the editorial staff approximately one year prior to publica- tion (18 June, 2003), in which the omission of a paper by COIFFAIT (1987) on two homonymies was corrected. In consequence, the revision contains a number of errors that require emendation. The following corrections (un- derlined) are proposed: p. 33, abstract: 11. 5 f.: The previously unknown male sexual characters of M. guignoti Coiffait, 1987 are described. Il. 17 ff.: ... = M. besucheti Bor- doni, 1980, syn. n.; M. lydicus Bordoni, 1980 = M. mimulus Fagel, 1970, syn. n.. = M. rhodicus Franz, 1987, syn. n. , = M. choparti Coiffait, 1987, syn. n.: M. fusculoides Coiffait, 1969 ... PIS paragraph, ll. 1 ff: Six species groups are present in the studied region: The M. apicalis group (3 species), the M. petrochilosi_ group (7 species), de same paragraph, Il. 17 ff.: M. guignoti, M. petro- chilosi, M. seleucus, M. cerrutii, and M. caricus, which he referred to four different groups, are evidently closely related and here all included in the M. petrochilosi group, ... p. 41: p. 43, p. 44, p. 63: p. 69, Ase paragraph, 11.2 ff.: ... the Mediterranean re- gion, the M. petrochilosi and the M. fusculus group are confined to the Eastern Mediterranean. The known distribution of the M._perrochilosi group ... or paragraph, ll. 8 ff.: ..., e. g. the majority of species of the M. petrochilosi group (M. guignoti, M. cerrutii, M. seleucus, M. impar, M. subquadratus). , 7" paragraph, IL I f.: The distribution patterns within the M. apicalis and the M. petrochilosi group, ... , 3 paragraph, Il. 5 £.: M. cerrutii is the sole rep- resentative of the M. petrochilosi group in Crete 3.6. Medon guignoti Coiffait, 1987 (Figs. 20-23, Map 4) Medon coriaceum Coitfait, 1969: 714f. (preoc- cupied). Medon guignoti Coiffait, 1987: 497 (replacement name). , legend to Figs. 20-23: Medon guignoti Coiffait: Aedeagus ... 2, section 3.6, comparative notes, Il. I f.: From M. cerrutii, M. petrochilosi, M. caricus, and related species, M. guignoti 1s distinguished ... legend to Map 4: Distributions of Medon petro- chilosi Coiffait (filled circles) and M. guignoti Coiffait (squares) in the Balkans, except for the type locality of M. guignoti based on revised re- cords. Distribution and bionomics, 11. 5 ff.: ... but due to the external similarity of the species of the petro- chilosi group, their identification is uncertain. ..., the species was found together with M. lydicus, M. fusculus, and M. semiobscurus. , Comparative notes, Il. I ff: M. seleucus is read- ily separated from M. cerrutii, M. caricus, M. guignoti, M. petrochilosi, and M. impar by the ... , Distribution and bionomics, Il. 18 f.: ... the spe- cies was found together with M. lydicus, M. cari- cus, and M. semiobscurus. legend to Map 12: Distributions of Medon beroni Coiffait (squares), M. subfusculus Fagel (open circles), and M. lydicus Bordoni (filled circles) ... legend to Figs. 94-97: Medon lydicus Bordoni: Aedeagus ... 92 Bonner zoologische Beiträge 54 (2005) : 3.24. Medon lydicus Bordoni, 1980 (Figs. 94- 97, Map 12) Medon lydicum Bordoni, 1980a: 114ff. Medon umbilicatum Coiffait, 1970: 102ff.; syn. n. (preoccupied). Medon mimulus Fagel, 1970: 157f. ; syn. n. (preoccupied) Medon rhodicum Franz, 1987: 76ff.; syn. n. Medon choparti Coiffait, 1987: 497; syn. n. (re- placement name for M. umbilicatus). p. 70, Comparative notes: M. /ydicus is distinguished from other species of the fusculus group only by the ... p. 70, Comments: delete last three paragraphs and re- place with: A comparison of the type material indicated above revealed that M. umbilicatus and M. mi- mulus are conspecific. The descriptions of both species were published in 1970; that of M. um- bilicatus was published on June 30 (HERMAN 2001), whereas the date provided on the front cover of the issue containing the description of M. mimulus is September 15 (A. SMETANA, Ot- tawa, pers. comm. 2002). Both names are preoc- cupied, so that neither of them is eligible as the valid name of the species. Medon umbilicatus Coiffait is a primary junior homonym of M. um- bilicatus Cameron, 1924 and M. mimulus Fagel is a secondary junior homonym of Lithocharis mimulus Casey, 1886, which is now in Medon. Two species are represented in the type series of M._lydicus. The paratype belongs to M. semiob- scurus (Fauvel); the holotype 1s conspecific with M. umbilicatus and the other names listed above. Since _M. lydicus is the oldest unpreoccupied name, it is to be regarded as the valid name of the species. Medon choparti Coiffait is a re- placement name for M. umbilicatus Coiffait and consequently a junior synonym of M. Iydicus Bordoni. An examination of the types of M. rhodicus Franz revealed that they are not specifically dis- tinct from M. umbilicatus, so that M. rhodicus is here synonymized with the senior name M. um- bilicatus. p. 77, Key, couplet 3, ll. 8 f.: ... modified setae. The M. petrochilosi group. p. 78, couplet 9,1. 10: ..... M. guignoti Coiftait p: 30, couplet 30,18: M. lydicus Bordon p. 81, after Coiffait (1984) insert: COIFFAIT H. (1987): Nouvelles corrections homonymiques (Col. Sta- phylinidae). Nouvelle Revue d’Entomologie 3 (4) (1986): 497-498. Acknowledgements. | am most grateful to all the col- leagues indicated in the material section for the loan and/or gift of material under their care. In addition, my sincere thanks are due to the following colleagues for the commu- nication of Medon records from England and Scandinavia: T. Clayhills (Parainen), E. Helve (Espoo), D. Lott (Birstall), I. Rutanen (Hyvinkáá), and M. Sórensson (Lund). Benedikt Feldmann and Michael Schiilke kindly proofread the manuscript. REFERENCES ALLEN, A. A. 1969. Notes on various little-known, doubt- ful, or misidentified British Staphylinidae (Col.). The Entomologist’s Monthly Magazine 105: 193-196. ALLEN, A. A. 1969. A note on Medon dilutus (Er.) (Col.: Staphylinidae) in Britain. Entomologist’s Record and ournal of Variation 108: 108. ASSING, V. 1998. Zur Kenntnis der Staphylinidenfauna der Atlantischen Inseln: neue Arten, Synonyme und Fund- daten (Col., Staphylinidae). Entomologische Nachrich- ten und Berichte 42: 139-146. ASSING, V. 1999a. On the Staphylinidae of La Gomera (Canary Islands), with descriptions of four new species (Coleoptera). Beitrage zur Entomologie, Berlin 49: 343-356. ASSING, V. 1999b. A revision of the genus Piochardia Heyden, 1870 (Insecta: Coleoptera: Staphylinidae: Aleocharinae). Annalen des Naturhistorischen Muse- ums Wien 101B: 277-301. ASSING, V. 2000. On the Staphylinidae of El Hierro, with additional new records from the Canary Islands (Col- eoptera). Entomologische Zeitschrift 110: 114-118. ASSING, V. 2004a. A revision of the Medon species of the Eastern Mediterranean and adjacent regions (Insecta: Coleoptera: Staphylinidae: Paederinae). Bonner zoolo- gische Beiträge 52: 33-82. ASSING, V. 2004b. A revision of Western Palaearctic Me- don STEPHENS (Coleoptera: Staphylinidae, Paederi- nae). Supplement I. — Linzer biologische Beitráge 36(2): 655-662. ASSING, V. & WUNDERLE, P. 1999. On the Staphylinidae of La Palma, Canary Islands, with descriptions of new species and remarks on some Canarian Stenus Latreille (Coleoptera). Entomologische Zeitschrift 109: 433- 443. BARANOWSKI, R. 1977. Intressanta skalbaggsfynd 1 (Col- eoptera). Entomologisk Tidskrift 98: 11-28. BARANOWSKI, R. 1979. Skalbaggar som lever 1 kaningan- gar (Coleoptera). Entomologisk Tidskrift 100: 57-60. BARANOWSKI, R. 1982. Intressanta skalbaggsfynd 6 (Col- eoptera). Entomologisk Tidskrift 103: 130-136. BERNHAUER, M. 1902. Neue Staphyliniden der paláarkti- schen Fauna. Deutsche Entomologische Zeitschrift 1901: 241-251. BERNHAUER, M. 1915. Beiträge zur Kenntnis der paláark- tischen Staphyliniden-Fauna. IV. Münchener Koleo- pterologische Zeitschrift 4: 262-270. — Volker ASSING: Western Palaearctic Medon 93 BERNHAUER, M. 1936. Neuheiten der palaearktischen Staphylinidenfauna. H. Pubblicazioni del Museo En- tomologico “Pietro Rossi” Duino 1: 303-325. BERNHAUER, M. 1940. Staphyliniden von den Azoren und Madeira. Commentationes Biologicae 8 (2): 1-9. BERNHAUER, M. & SCHUBERT, K. 1912. Staphylinidae HI. In: SCHENKLING, S.: Coleopterorum Catalogus 5 (40): 191-288. Berlin: Junk. BOHÁC, J. 1981. Faunistic records from Czechoslovakia. Acta Entomologica Bohemoslovaca 78: 230. BOHÁC, J. 1985. Review of the subfamily Paederinae (Col- eoptera, Staphylinidae) of Czechoslovakia. Part II. Ac- ta Entomologica Bohemoslovaca 82: 431-467. BORDONI, A. 1980. Studi sui Paederinae — III - I Medon Steph. paleartici con descrizione di nuove specie medi- terranee (Col. Staphylinidae). Bollettino del Laborato- rio di Entomologia Agraria “Filippo Silvestri” di Por- tici 37: 73-125. BORDONI, A. 1981. Coleotteri stafilinidi raccolti in grotta da Sergio Puddu nella Sardegna centro meridionale, 61° contributo alla conoscenza degli Staphylinidae. Bollettino della Societa Sarda di Scienze Naturali 21: 137-147. BORDONI, A. 1988. Note su alcuni Stafilinidi raccolti du- rante la spedizione “Algeria 1984” del Museo Regio- nale di Scienze Naturali di Torino (Coleoptera). Bol- lettino del Museo Regionale di Scienze Natural, Torino 6: 341-347. BRUGE, H. DRUGMAND, D. & HAGHEBAERT, G. 2001. Co- leoptera Staphylinidae de Belgique et du Grand-Duché de Luxembourg. Catalogue commenté et éléments de biogéographie. Bulletin de la Société royal belge d’Entomologie/Koninklijke Belgische Vereniging voor Entomologie 137: 139-172. BUCHE, B. & ESSER, J. 1999. Faunistisch bemerkenswerte Käferfunde aus Mecklenburg-Vorpommern. Entomo- logische Nachrichten und Berichte 43: 129-135. CICERONI, A. & ZANETTI, A. 1995. Paederinae e Staphyli- ninae (generi 079-147). Pp. 19-33 in: CICERONI, A., PUTHZ, V. & ZANETTI, A. (eds.) Coleoptera, Polyphaga II (Staphylinidae). Checklist delle specie della fauna italiana, Calderini Bologna, Fasc. 48: 1-65. COIFFAIT, H. 1970a. Formes nouvelles ou mal connues des genres Medon et Hypomedon. Annales de Spéléologie 24 (1969): 701-727. COIFFAIT, H. 1970b. Staphylinides nouveaux ou mal connus de la région paléarctique occidentale. Bulletin de la Société d'Histoire Naturelle de Toulouse 106: 99-111. COIFFAIT, H. 1970c. Corrigenda. Annales de Spéléologie 25: 909. COIFFAIT, H. 1973a. Staphylinides nouveaux ou mal connus d'Europe, d’Afrique du nord et du Moyen- Orient. Les sous-genres du genre Bledius. Nouvelle Revue d Entomologie 3: 107-123. COIFFAIT, H. 1973b. Staphylinides nouveaux ou mal connus du Maroc. Bulletin de la Société des Sciences Naturelles et Physiques du Maroc 53: 269-292. COIFFAIT, H. 1978. Staphylinides nouveaux ou mal connus de la France et de la région méditerranéenne. Nouvelle Revue d’Entomologie 8: 267-279. COIFFAIT, H. 1980. Staphylinides nouveaux. Nouvelle Re- vue d’Entomologie 10: 37-49. COIFFAIT, H. 1984. Coléopteres Staphylinidae de la région paléarctique occidentale. V. Sous famille Paederinae Tribu Paederini 2,-Sous famille Euaesthetinae. Nou- velle Revue d’Entomologie Supplement 13 (4): 1-424. COIFFAIT H. 1987. Nouvelles corrections homonymiques (Col. Staphylinidae). Nouvelle Revue d’Entomologie 3 (4) (1986): 497-498. DODERO, A. 1922. Appunti coleotterologici. Bolletino del- la Societa Entomologica Italiana 54: 67-76. DRUGMAND, D. 1989. Distribution et phénologie des Pae- derinae de Belgique (Coleoptera, Staphylinidae). Stu- diedocumenten van het Koninklijk Belgisch Instituut voor Natuurwetenschappen 55: 1-52. ERICHSON, G. F. 1839. Die Kafer der Mark Brandenburg. 1. Bd., 2. Abt., Berlin: 385-740. ERMISCH, K. & LANGER, W. 1933. Ueber die Käfergäste im Winterlager des Maulwurfs, zugleich ein Beitrag zur Käferfauna des Vogtlandes. Koleopterologische Rund- schau 19: 16-24, FAIRMAIRE, L. 1860. [New species]. In: FAIRMAIRE, L. & COQUEREL, C.: Essai sur les coléopteres de Barbarie. Annales de la Société Entomologique de France (3) 8: 145-176. FALCOZ, L. 1914. Contribution a l'étude de la faune des microcavernes, faune des terriers et des nids. Annales de la Société Linnéenne de Lyon, Nouvelle Série, 61: 59 - 243. FAUVEL, A. 1872. Lithocharis africana Fauvel. In: REICHE, L. Catalogue des coléopteres de l’Algerie et contrées voisines avec descriptions d’especes nouvelles. Mé- moires de la Société Linnéenne de Normandie 15 (1869): 1-44. FAUVEL, A. 1886. Les Staphylinides du Nord de l'Afrique. Notices Entomologiques 9: 1-92. FAUVEL, A. 1906. M. augur n. sp. In: SAINTE-CLAIRE DE- VILLE, J. Catalogue critique des Coléopteres de la Corse. Revue d’Entomologie. Supplement 25: 1-112. FOWLER, C. 1888. The Coleoptera of the British Islands. Vol. II. Staphylinidae. London: 444 pp. FRANZ, H. 1938. Eine Káferfauna aus Fuchsbauen des nórdlichen Burgenlandes. Koleopterologische Rund- schau 24: 30-32. GILLERFORS, G. 1986. Contribution to the coleopterous fauna of the Azores. Boletim do Museu Municipal do Funchal 38: 16-27. GILLERFORS, G. 1988. Second contribution to the coleop- terous fauna of the Azores. Bocagiana, Museu Mu- nicipal do Funchal 118: 1-6. GRAVENHORST, J. L. C. 1802. Coleoptera microptera Brun- svicensia. Brunsuigae, 206 pp. GUSAROV, V. I. 1989. Fauna 1 ekologua stafilinid (Coleop- tera, Staphylinidae) Kryma. Podsemeistva Metopsiinae, Proteinae, Omaliinae, Piestinae, Tachyporinae, Oxy- porinae, Steninae, Paederinae, Xantholininae, Staphylininae [In Russian]. Vestnik Leningradskogo Universiteta. Biologii 1989: 3-17. GUSAROV, V. I. 1992. Novye 1 maloizvestnye paleark- ticheskie stafilinidy (Coleoptera, Staphylinidae). Soobshchenie 4. Vestnik Leningradskogo Universiteta, Biologia Ser.3 (3): 11-25. HANSEN, M. 1996. Katalog over Danmarks biller. Ento- mologiske Meddelelser 64 (1&2): 1-231. HANSEN, M., MAHLER, V., PRITZL, G. & RUNGE, J. B. 1994. 13. tilleg til 'Fortegnelse over Danmarks biller' (Col- eoptera). Entomologiske Meddelelser 62: 65-89. 94 Bonner zoologische Beitráge 54 (2005) HANSEN, M., LILJEHULT, H., MAHLER, V. & PEDERSEN, J. 1995. 14. tilleeg til “Fortegnelse over Danmarks biller” (Coleoptera). Entomologiske Meddelelser 63: 21-50. HANSEN, V. 1964. Fortegnelse over Danmarks biller (Coleoptera). Entomologiske Meddelelser 33: 1-507. HEINEMANN, R. 1910. Käfer in Maulwurfsnestern. Ento- mologische Blatter 6: 121-126, 157-164. Horton, A. 1933. Die Koleopteren-Fauna der Maulwurfs- nester der Rheinprovinz. Verhandlungen des Naturhis- torischen Vereins der preußischen Rheinlande und Westfalens 89: 144-163. HORION, A. 1965. Faunistik der Mitteleuropáischen Kafer, Bd. X: Staphylinidae, 2. Teil, Paederinae bis Staphylininae. Überlingen, Bodensee. 335 pp. HUGENTOBLER, H. 1966. Beitrag zur Kenntnis der Käfer der Nordostschweiz. — Naturwiss. Ges. St. Gallen. 248 PP- JANAK, J. 1993. Interessante Funde der Kurzflügler aus Mähren und aus der Slowakei (Coleoptera: Staphylinidae). Klapalekiana 29: 1-17. JARRIGE, J. 1949. Contribution a l’etude des Staphylinides circummediterraneens. Annales de la Société Entomo- logique de France 116 (1947): 59-72. Joy, N. H. 1932. A Practical Handbook of British Beetles. Vol. I. London. 622 pp. KAHLEN, M. 1987. Nachtrag zur Káferfauna Tirols. Ergán- zung zu den bisher erschienenen faunistischen Arbei- ten über die Käfer Nordtirols (1950, 1971 und 1976) und Südtirols (1977). Beilageband 3 zu den Veröffent- lichungen des Museum Ferdinandeum 67: 288 pp. KocH, C. 1938. Apunti sugli stafilinidi italiani IV. Bollet- tino della Societa Entomologica Italiana 70: 135-141. Koch, C. 1939. Die Käfer der libyschen Ausbeute des Herrn Georg Frey. Mitteilungen der Múnchener Ento- mologischen Gesellschaft 29: 216-293. Koch, K. 1968. Käferfauna der Rheinprovinz. Decheniana (Bonn), Beihefte 13: 1-382. Koch, K. 1974. Erster Nachtrag zur Káferfauna der Rhein- provinz. Decheniana (Bonn) 126: 191-265. KÖHLER, F. 2000. Erster Nachtrag zum “Verzeichnis der Käfer Deutschlands”. Entomologische Nachrichten und Berichte (Dresden) 44: 60-84. KÖHLER, F. & KLAUSNITZER, B. 1998. Verzeichnis der Kä- fer Deutschlands. Entomologische Nachrichten und Berichte (Dresden) Beiheft 4: 1-185. KOLBE, W. 1918. Beiträge zur schlesischen Käferfauna. Entomologische Mitteilungen 7: 200-211. KORGE, H. 1989. Beiträge zur Kenntnis der Tierwelt von Berlin (West). Teil I: Kurzflügelkäfer (Coleoptera, Staphylinidae). Berliner Naturschutzblätter 33: 28-40. KRAATZ, G. 1857. Naturgeschichte der Insecten Deutsch- lands. Abt. 1. Coleoptera. Zweiter Band. Lief. 3-4. Berlin: 377-768. KRAATZ, G. 1858. Description de quelques nouvelles espe- ces de Coléopteres. Bulletin de la Société Entomologi- que de France 6: CLXXXVIHI-CXCH. LATTIN, G. de 1967. Grundriss der Zoogeographie. Gustav Fischer Verlag, Stuttgart: 602 pp. LINKE, M. 1907. Verzeichnis der in der Umgebung von Leipzig beobachteten Staphyliniden. Sitzungsberichte der Naturforschenden Gesellschaft zu Leipzig 1906- 1907: 1-54. LINKE, M. 1913. Erster Beitrag zur Kenntnis der Staphyliniden des Königreichs Sachsen. Entomologi- sche Blätter 9: 76-81. LINKE, M. 1927. Zweiter Beitrag zur Kenntnis der Staphyliniden des Freistaates Sachsen und einiger Grenzgebiete. Coleopterologisches Centralblatt 1: 355- 364. Lott, D. A. & DUFF, A. G. 2003. Staphylinidae. In: Check- list of Beetles of the British Isles. Revised 28th Janu- ary 2003. www.coleopterist.org.uk. LUIGIONI, P. 1929. I coleotteri d'Italia. Catalogo sinonimi- co-topografico-bibliografico. Memorie Pont. Accademia delle Scienze — I nuovi Lincei. Serie II, 13: 1-1160. LUNDBERG, S. 1978. Bidrag till kännedom om svenska skalbaggar. 17 (Coleoptera). Entomologisk Tidskrift 99: 31-34. LUNDBERG, S. 1995. Catalogus Coleopterorum Sueciae. — Naturhistoriska Riksmuseet, Entomologiska Fórenin- gen: without pagination. LUNDBLAD, O. 1958. Die Arthropodenfauna von Madeira nach den Ergebnissen der Reise von Prof. Dr. O. Lundblad Juli-August 1935. XXXV. Die Káferfauna der Insel Madeira. Arkiv fór Zoologi 11 (30): 461-524. MACHADO, A. & OROMI, P. 2000. Elenco de los coleópte- ros de las Islas Canarias. Instituto de Estudios Cana- rios, La Laguna. 306 pp. MEYBOHM, H. 1997. Meldungen zur Káferfauna von Schleswig-Holstein, Hamburg und Nord-Niedersachsen. 3. Teil. Bombus, Faunistische Mitteilungen aus Nord- westdeutschland 3: 102-104. NORMAND, H. 1935. Contribution au Catalogue des Co- léoptéres de Tunisie (4. fascicule). Bulletin de la So- ciété d’Histoire Naturelle de l’Afrique du Nord 25 (1934): 356-390. . NOWOSAD, A. 1990. Staphylinidae (Coleoptera) gniazd kreta — Talpa europaea L. w Polsce. Uniwersytet im. Adama Mickiewicza w Poznaniu, Seria Zoologia Nr. 15: 1-254. NowosAD, A. 2000. Wstepne wyniki badan nad chrzaszczami kusakowatymi Staphylinidae (Micrope- plinae — Tachyporinae) gniazd ptaków 1 ssaków. Pp. 29-40 in: Konferenzmaterialien Erstes Staphylinidae- Symposium Rogów, 10-12 November 1999, Rolle der Kurzfliigler (Coleoptera, Staphylinidae) in der Funkti- on und im Schutz von Waldókosystemen. Poznañ. OSELLA, G. & ZANETTI, A. 1974. La coleotterofauna del nidi di Talpa europaea L. nell Ttalia settentrionale a nord del fiume Po. Bollettino di Zoologia Agraria e di Bachicoltura, Ser. II, 12: 43-200. OUTERELO, R. 1980. Los Staphylinoidea de la Sierra de Cazorla. In: M. VIEDMA (ed.): Fauna de Cazorla. In- vertebrados. Monografias. Instituto Nacional para la Conservacion de la Naturaleza 23: 53-71. OWEN, J. 1998. Medon dilutus (Erichson) (Col.: Staphylinidae) in Richmond Park, Surrey. Entomolo- gist’s Record and Journal of Variation 110: 269-270. OWEN, J. 2000. Coleoptera occurring underground at the roots of old trees. Entomologist's Gazette 51: 239-256. PEREZ ARCAS, L. 1874. Especies nuevas 6 críticas de la fauna española. Anales de la Sociedad española de Historia natural 3: 111-155. PEYERIMHOFF, P. de 1919. Notes sur la biologie de quel- ques Coléopteres phytophages du Nord-Africain (troi- siéme série) avec les descriptions de cinq especes nou- velles et de sept sous-especes ou variétés. Annales de la Société Entomologique de France 88: 169-258. Volker ASSING: Western Palaearctic Medon 95 PEYERIMHOFF, P. de 1949. Etudes et descriptions de co- léopteres marocaines. II. Bulletin de la Société des Sciences Naturelles du Maroc 25-27: 248-308. Porta, A. 1926. Fauna Coleopterorum Italica, 2. Staphyli- noidea. Piacenza: 1-405. RENNER, K. & GRUNDMANN, B. 1984. Neuheiten und Sel- tenheiten der westfälischen Käferfauna III. Entomolo- gische Blatter 80: 71-84. ROTTENBERG, A. v. 1870. Beiträge zur Coleopteren-Fauna von Sicilien. Berliner Entomologische Zeitschrift 14: 11-40. SAHLBERG, J. 1908. Coleoptera mediterranea et rosso- asiatica nova et minus cognita, maxima ex parte intine- ribus annis 1895-1896, 1898-1899 et 1903-1904 col- lecta. Ofversigt af Finska Vetenskaps-Societetens Fór- handlingar 50 (7): 1-94. SAINTE-CLAIRE DEVILLE, J. 1906. Catalogue critique des Coléopteres de la Corse. Revue d’Entomologie. Sup- plement 8, 25: 1-112. SCHEERPELTZ, O. 1933. Staphylinidae VII. In: Schenkling, S. (ed.): Coleopterorum Catalogus 6 (129): 989-1500. Berlin: Junk. SCHEERPELTZ, O. 1958. Staphyliniden aus den Picentinischen Bergen im südlichen Mittel-Italien. Memorie del Museo Civico di Storia Naturale Verona 6: 143-173. SCHEERPELTZ, O. 1964. Ergebnisse der von Dr. O. Paget und Dr. E. Kritscher auf Rhodos durchgeführten Ex- kursionen. Teil XI. Coleoptera, Fam. Staphylinidae. Annalen des Naturhistorischen Museums Wien 67: 667-672. SCHEERPELTZ, O. 1968. Catalogus Faunae Austriae. Teil XV fa: Coleoptera-Staphylinidae. — Österreichische Akademie der Wissenschaften, Wien: 279 pp. SCHILLER, W. 1989. Die Staphylinidae (Kurzflúgler) des Belchengebietes im Súdschwarzwald. In: Der Belchen — Geschichtlich-naturkundliche Monographie des schönsten Schwarzwaldberges. Natur- und Land- schaftsschutzgebiete Baden-Württemberg 13: 1031- 1081. SCHOLZE, P. & JUNG, M. 1993. Beiträge zur Faunistik der Kurzflügler (Coleoptera, Staphylinidae) im Nordharz und Vorharz. I. Unterfamilien Micropeplinae bis Tachyporinae. Entomologische Nachrichten und Be- richte 37: 225-234. SCHÜLKE, M. 1998. Medon rufiventris (Nordmann) in Sachsen-Anhalt (Staph.). Kleine Mitteilung Nr. 2182. Entomologische Blätter 94: 98. SCRIBA, W. 1870. Lithocharis spelaea Scriba n. sp. — In: HEYDEN, L. v.: Entomologische Reise nach dem stidli- chen Spanien, der Sierra Guadarrama und Sierra Morena, Portugal und den Cantabrischen Gebirgen. Berliner Entomologische Zeitschrioft 14 (Beiheft): 58- 176. SERRANO, A. R. M. 1993. Medon vicentensis n. sp., a new species of eyeless rovebeetle (Coleoptera: Staphylinidae: Paederinae) from a cave in the island of Madeira. Bo- cagiana, Museu Municipal do Funchal No. 165: 1-7. SILFVERBERG, H. 1992. Enumeratio Coleopterorum Fenno- scandiae, Daniae et Baltiae. — Helsingin Hyönteis- vaihtoyhdistys, Helsingfors Entomologiska Bytes- förening, Helsinki: 94 pp. SMETANA, A. 1970. Staphylinidae (Coleoptera) from the Azores and Madeira. Boletim do Museu Municipal do Funchal. 25 (110): 53-67. STROUHAL, H. & BEIER, M. 1928. Beitrag zur Coleopteren- fauna der Maulwurfsnester in der nächsten Umgebung Wiens. Zeitschrift für Morphologie und Ökologie der Tiere 12: 191-239. SZUJECKI, A. 1968. Wstep do znajomosci kusakowatych (Staphylinidae, Col.) Niziny Mazowieckiej na podsta- wie zbioru W. Maczynskiego. Polskie Pismo Entomo- logiczne 38: 693-752. TERLUTTER, H. 1995. Coleoptera Westfalica: Familia Staphylinidae, Subfamiliae Oxytelinae, Oxyporinae, Steninae, Euaesthetinae, Paederinae. Abhandlungen aus dem Westfälischen Museum für Naturkunde 57: 3- 84. TRONQUET, M. 2001. Catalogue des Coleopteres des Pyre- nees-Orientales. Vol. 1. Staphylinidae. Revue de l’Association Roussillonnaise d’Entomologie 10, Sup- plement: 1-88. UHLIG, M. & VOGEL, J. 1981. Zur Staphylinidenfauna der Umgebung von Waren/Miiritz (Mecklenburg). Mitte1- lungen des zoologischen Museums Berlin 57: 75-168. VOGEL, J. 1979. Faunistisch bedeutsame und fiir die DDR neue Staphylinidae (Coleoptera) aus der Oberlausitz. Abhandlungen und Berichte des Naturkundemuseums Görlitz 52, 8 (1978): 1-13. VOGEL, J. 1982. Faunistisch bedeutsame und ftir die DDR neue Staphylinidae (Coleoptera) aus der Oberlausitz, Teil H. Abhandlungen und Berichte des Naturkunde- museums Görlitz 55, 6: 1-26. VOGT, H. 1956. Käfer in Maulwurfsnestern. Naturschutz- stelle Darmstadt, Institut zur Erforschung, Pflege und Gestaltung der Landschaft, Schriftenreihe IH, 3: 119- 134. VOGT, H. 1968. Bemerkenswerte Káfergesellschaften II. Anbrúchige Buche mit Lasius brunneus Latr. Nach- richtenblatt der Bayerischen Entomologen 17: 50-55. WACHEK, F. 1955. Die entoparasitischen Tylenchiden. Pa- rasitologische Schriftenreihe Heft 3: 1-119. WAGNER, T. 2002. Zur Káferfauna (Coleoptera) einer Kies- grube im Neuwieder Becken. Mitteilungen der Ar- beitsgemeinschaft Rheinischer Koleopterologen (Bonn) 12: 45-63. WITTWER, A. 1988. Recherche faunistique sur les Staphy- linidae (Col.) de la rive sud du lac Neuchatel. Mit- teilungen der Schweizerischen Entomologischen Ge- sellschaft 61: 233-239. WOLLASTON, T.V. 1857. Catalogue of the Coleopterous In- sects of Madeira in the Collection of the British Mu- seum. London. 234 pp. WOLLASTON, T.V. 1864. Catalogue of the Coleopterous In- sects of the Canaries in the Collection of the British Museum. London. x11 + 648 pp. ZERCHE, L. 1980. Faunistisch interessante Staphylinidae aus der DDR (Coleoptera). Entomologische Nachrich- ten 24: 145-165. Author’s address: Dr. Volker ASSING, Gabelsberger- straße 2, D-30163 Hannover, Germany; E-mail: vassing.hann@t-online.de Received: 19.11.2003 Accepted: 15.04.2004 Revised: 07.01.2005 Corresponding editor: M. Schmitt Bonner zoologische Beitráge Band 54 (2005) | Heft 1 | Seiten 97 Bonn, April 2006 SCHNEIDER, H. (2005): Bioakustik der Froschlurche — Einheimische und verwandte Arten. Mit Audio CD (beides zusammen erschienen als Supplement der Zeitschrift fiir Feldherpetologie 6). Laurenti Verlag, Bielefeld. 135 Seiten, 146 Abbildungen. ISBN 3-933066-23-9. Preis: 28,- € Mit diesem kleinen Buch und der dazugehórigen CD legt der deutsche Nestor bioakustischer Forschung an Froschlurchen eine Zusammenfassung seiner langjáhri- gen Forschungsarbeiten vor. Ausführlich dargestellt werden das Rufverhalten und die Rufe von Männchen und Weibchen von 25 Taxa aus verschiedenen Teilen Europas, aus dem Vorderen Orient und Kasachstan in Text und Abbildungen. Vorangestellt sind zwei kurze Einführungen, eine zu den Verhaltenszusammen-hängen, in denen Froschlurche Rufe äußern, und in die Termino- logie dieser Rufe, die andere zur Bioakustik unter dem Titel „Akustische Merkmale der Paarungsrufe und Ana- lyse der Rufe“. Die meisten Rufformen sind im Buch in paralleler Darstellung derselben Rufe im Oszillogramm (Zeitsignal) und Lautspektrogramm abgebildet. Der Text umfasst 1m wesentlichen die Beschreibungen der akustischen Strukturmerkmale der Rufe und geht für viele Taxa ausführlich auf deren Abhängigkeit von Luft- bzw. Wassertemperatur, bei der die Tiere rufen, sowie von ihrer Körpergröße ein. Die meisten Ruffor- men sind auf der CD mit Angabe der jeweiligen Tempe- ratur, bei der diese aufgezeichnet wurden, mit Hörbei- spielen dokumentiert. Veröffentlichungen dieser Art wünscht man sich auch für weitere Tiergruppen, die zur Lautbildung befähigt sind, vor allen Dingen wie hier mit begleitender Audio- Dokumentation und so detaillierter Darstellung der ver- schiedenen Lautformen in Text und Analyse mit den dazugehórigen Abbildungen. So willkommen und niitz- lich diese Neuerscheinung ohne Zweifel ist, sie hat lei- der auch einige Aspekte, die man kritisch erwáhnen muss. Schnell fallt auf, dass eine nennenswerte Anzahl im Text zitierter Publikationen im Literaturverzeichnis fehlt (z.B. BLAB 1982; HEINZMANN 1970; HEUSSER 1969; LÖRCHER 1969; LÖRCHER & SCHNEIDER 1973; MÜLLER 1984; NASCETTI et al. 1996, 1999; NEUBAUR 1949; RADWAN & SCHNEIDER 1988). Die online- Version der Weltliste ,,Amphibian Species of the World“ (FROST 2004) ist zwar zitiert, im Text des Bu- ches wird aber bei Formen mit ‚wechselvoller’ Taxo- nomie, zu der auch der Buchautor aktiv auf der Grund- lage bioakustischer Befunde beigetragen hat, wie z.B. im Fall von Rana balcanica, darauf und den aktuellen Stand der Diskussion nicht Bezug genommen. Angaben über ihr Verbreitungsgebiet erfolgen nur für wenige Taxa. Nahezu alle abgebildeten Lautspektrogramme wurden mit relativ großem Zeitfenster und damit hoher Fre- quenz- und entsprechend geringer Zeitauflösung analy- siert. Das führt angesichts der Tatsache, dass viele Rufe der untersuchten Froschlurche sehr schnelle Amplitu- den-Modulationen aufweisen, zu einer kaum angemes- senen Darstellung der ‚eigentlichen’ Rufstruktur in den Lautspektrogrammen. Dazu seien hier nur zwei Beispiele angeführt: Abb. 27 (Paarungsruf) und Abb. 28 (Befrei- ungsrufe) von Bufo bufo (S. 40). In beiden Fällen wird die in den Oszillogrammen gut sichtbare Struktureigen- schaft einer schnellen Amplituden-Modulation in der ersten Hälfte der Rufe im Spektrogramm in der Fre- quenzdomäne dargestellt. Daher ist auch die Beschrei- bung der Struktur der letztgenannten Rufe im Text als „.... aus Impulsen bestehende erste Teil weist ein breites Spektrum auf, das bis 2500 Hz reicht und Frequenzbän- der erkennen lässt, ....* (S. 41) sachlich nicht wirklich zutreffend. Auch Unstimmigkeiten kommen vor. So wird in der Beschreibung der Struktureigenschaften der Paarungsrufe von Männchen und Weibchen der Ge- burtshelferkröte (S. 23 — 24, Abb. 12, 13) festgestellt, dass bei gleicher Körpergröße und Lufttemperatur die Grundfrequenz der Rufe (der Weibchen?, G.P.) um 60 — 80 Hz niedriger ist. Der in Abb. 12 gezeigte Paarungsruf eines Männchens (keine Größenangabe) bei 10 °C Luft- temperatur hat eine Grundfrequenz von um 1200 Hz, derjenige (Abb. 13) eines Weibchens (keine Größenan- gabe) bei 17 °C eine von um 700 Hz (Frequenzen je- weils aus dem abgebildeten Spektrogramm geschätzt). Die eigene lautspektrographische Analyse der beiden betreffenden Hörbeispiele auf der CD entspricht exakt den jeweiligen Abbildungen in dem Buch. Trotz der genannten Kritikpunkte darf man abschlie- Bend festhalten, dass diese Neuerscheinung für interes- sierte Laien und Fachwissenschaftler eine Fülle interes- santer Information bietet, die sie in vielfältiger Weise nutzen können. Angesichts des Buchtitels „Bioakustik der Froschlurche”” werden aber Leser mit spezifischen Erwartungen in dieser Hinsicht bioakustische Analysen und darauf fußende Beschreibungen der Lautstrukturen ‚lege artis’ bei vielen Taxa vermissen. G. 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(2005): Bioakustik der Froschlurche — Einheimische und verwandte Arten Titelbild/ Cover illustration: Polyommatus theresiae Schurian, van Oorschoot & van den Brink, 1992 (Insecta: Lepidoptera: Lycaenidae), from the collection of Sigbert Wagener, now at Zoologisches Forschungsmuseum Alexander Koenig, Bonn (see the con- tribution of Sigbert WAGENER, pp. 3-23) FORSCHUNGS mus ENIG Herausgegeben vom Bonne [ Zoologischen Forschungsmuseum 720010 oische En . Bonn eiträge Heft 3 2006 200s) ral Leibniz Gemeinschaft Bonner zoologische Beiträge werden publiziert 1m Eıgenverlag Zoologisches Forschungsmuseum Alexander Koenig, Adenauerallee 160, D-53113 Bonn (Germany) Die Zeitschrift erscheint mit vier Heften um Jahr, zum Preis von 11,50 € je Heft bzw. 46,- € je Band incl. Versand. Korres- pondenz betreffend Abonnement, Kauf oder Tausch bitten wir zu richten an die Bibliothek, Zoologisches Forschungsmu- seum Alexander Koenig, Adenauerallee 160, D-53113 Bonn (Germany). E-mail: d.stemebach.zfmk(@uni-bonn.de. © Zoologisches Forschungsmuseum Alexander Koenig (ZFMK ), Bonn, Germany ISSN 0006-7172 Druck: Druck Center Meckenheim Bonner zoologische Beiträge Editorial Board (in brackets: editorial competence) Prof. Dr. Michael SCHMITT (editor-in-chief, Coleoptera, theory), Tel./Fax: +49 228-9122 286, E-mail: m.schmitt(@uni-bonn.de Dr. Renate VAN DEN ELZEN (Vertebrata, except Mammalia), Tel. +49 228-9122 231, Fax: +49 228-9122 212, E-mail: r.elzen.zfmk(@uni-bonn.de Dr. Bernhard A. HUBER (Invertebrata, except Insecta), Tel. +49 228-9122 294, Fax: +49 228-9122 212, E-mail: b.huber.zfmk(@uni-bonn.de Dr. Gustav PETERS (Mammalia, bioacoustics), Tel. +49 228-9122 262, Fax: +49 228-9122 212, E-mail: g.peters.zfmk(@uni-bonn.de Dr. Bradley J. SINCLAIR (Diptera, language advisor), Tel. +49 228-9122 292, Fax: +49 228-9122 212, E-mail: b.sinclair.zfmk(@uni-bonn.de Dr. Dieter STUNING (Insecta, except Coleoptera and Diptera), Tel. +49 228-9122 220, Fax: +49 228-9122 212, E-mail: d.stuening.zfmk(@uni-bonn.de Editorial office: Adenauerallee 160, D-53113 Bonn, Germany Advisory Board Prof. Dr. Miguel Angel ALONSO-ZARAZAGA, Museo nacional, Ciencias Naturales, E-28006 Madrid, E-mail: zarazaga(@mncn.csic.es Prof Dr. Ulrike Aspöck, Naturhistorisches Museum, 2. Zoologische Abteilung (Insekten), Postfach 417, A-1014 Wien, E-mail: ulnke.aspoeck(@nhm-wien.ac.at Prof Dr. Paolo AUDISIO, Universita di Roma ,,La Sapienza”, Dip.Biol Anim. e dell'Uomo (Zool. ), Viale dell'Universita 32, 1-00185 Roma, Tel.: +39 6-49914744, E-mail: paolo.audisio(@uniroma 1 .1t Prof Dr. Aaron M. BAUER, Villanova University, Department of Biology, 800 Lancaster Avenue, Villanova, PA 19085-1699, USA. 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Box 50007, SE-104 05 Stockholm, E-mail: sven.kullander(@nrm.se Prof Dr. Steven PERRY, Rheinische Friedrich-Wilhelms- Universitat, Institut fur Zoologie, Poppelsdorfer Schloss, D-53115 Bonn, Tel. +49 228-73 3807, E-mail: perry(@uni-bonn.de Dr. Wolfgang SCHAWALLER, Staatliches Museum fur Naturkunde, Rosenstem 1, D-70191 Stuttgart, Germany, Tel. +49 711-8936 221, Fax +49 711-8936 100, E-mail: schawaller.smns(@naturkundemuseum-bw de Dr. W. David Sissom, Dept. of Life, Earth and Environ- mental Sciences, W. Texas A. & M. University, WTAMU Box 60808, Canyon, Texas 79016, USA, E-mail: dsissom(@wtamu.edu/ Dr. Miguel VENcES, University of Amsterdam, Zoological Museum, Mauritskade 61, PO Box 94766, NL-1090 GT Amsterdam, The Netherlands, Tel. +31 20-525 7319, E-mail: vences(@science.uva.nl PD Dr. Heike WÄGELE, Rheinische Fnedrich- Wilhelms- Universität, Institut für Evolutionsbiologie und Okologie, D-53121 Bonn, Tel.: +49 228-73 5159, Fax: +49 234-322 4114, E-mail: hwaegele(@evolution.unı-bonn.de Dr. Erich WEBER, Eberhard-Karls-Universitat, Zoologische Schausammlung, Sigwartstr. 3, D-72076 Tubingen, Germany, E-mail: erich. weber(@uni-tuebingen.de Bonner zoologische Beitráge Band 54 (2005) | Heft 3 | Seiten 97-102 Bonn, August 2006 A Revision of Porocallus Sharp. New Synonyms} (Insecta: Coleoptera: Staphylinidae: Aleocharit Volker ASSING Hannover, Germany nd New Species ae: Oxypodit Platysmarthrusa Pace, 1999 syn. n. = Ischyradelia Pace, 1999 syn. n.; Porocallus insignis Sharp, 1888. =. Platys- ~ marthrusa chinensis Pace, 1999 syn. n. Two new species are described and illustrated: Pofgcalhıs hfävaci sp. n. and_P. ligo sp. n. Porocallus tianmuensis (Pace) comb. n. (previously in /schyradelia) is briefly re@escribed. New records of P. insignis from China and a key to the four species of the genus are presented. The distributiof f the genus is mapped. Key words. Eastern Palaearctic region - China, taxonomy, description, new records, distribution, key to species 1. INTRODUCTION In a recent paper on the systematics of Porocallus Sharp, 1888 one species was recognized: the type spe- cies P. insignis Sharp, 1888 (ASSING 2001). The species is very widespread in the Eastern Palaearctic region, its distribution ranging from China to Japan. Based on ad- ditional material, which has become available in the meantime, and on a study of type material, three new synonymies are proposed and two new species are de- scribed. An additional undescribed species from Hubei is not named, as it is represented only by a single fe- male; the external characters distinguishing it from other species of the genus are not pronounced and the spermatheca is not very distinctive. 2. MATERIALS The material treated in this paper is deposited in the fol- lowing collections: MHNG Museum d Histoire (G. Cuccodoro) NHMW Naturhistorisches Museum Wien (H. Schill- hammer) Naturelle, Geneve cAss author’s private collection 3. POROCALLUS SHARP, 1888 Porocallus Sharp, 1888 (SHARP 1888: 286f). Platysmarthrusa Pace, 1999 (PACE 1999: 107f) syn. n. Ischyradelia Pace, 1999 (PACE 1999: 108) syn. n. Comments: When PACE (1999) described Platys- marthrusa, he did not compare it with Porocallus, a ge- nus at that time known only from Japan. The type species of both genera are conspecific (see below), so that Platysmarthrusa is consequently a junior synonym of Porocallus. According to the original description of /s- chyradelia, this taxon is distinguished from Platys- mathrusa by the shape and chaetotaxy of the ligula, as well as by the stouter second joint of the maxillary palpus and the slightly larger first joint of the labial palpus. An examination of the holotypes of both type species and of material of P. insignis from various localities re- vealed the following: The shape and chaetotaxy of the ligula are subject to considerable intraspecific variation (Figs. 1-2), so that differences seen in the ligulae of just two specimens do not justify a distinction at the generic level (in this case not even at the species level). The dif- ference in the shape of the second joint of the maxillary palpus is clearly based on an artefact: in the holotype of the Platysmathrusa chinensis, it 1s much wider than il- lustrated by PACE (1999: 109); the impression of a slen- der joint is caused by the fact that it is turned on the sli- de, so that, when viewed from above, it is not seen at its widest aspect. Finally, the first joint of the labial palpus is in fact slightly less oblong, but this extent of variation is quite normal within genera; there are numerous ex- amples of Staphylinidae with a much more pronounced intraspecific variation of the shape of the labial palpi. Most importantly, however, Porocallus is characterized by numerous highly conspicuous and very distinctive apomorphic characters states, all of them shared also by the holotype of /schyradelia tianmuensis: especially the long, massive, and densely pubescent antennae, the con- spicuous triangular flattened third joint of the maxillary palpi, the coarse and extremely dense puncturation of the forebody, and finally also the similar morphology of the primary and secondary sexual characters (see PACE 1999: 109-110). In addition, the facies is most similar (especially the shape of head and pronotum, the mor- phology of the abdomen), so that there is no doubt that Ischyradelia tianmuensis is in fact a species of Porocal- lus and that consequently /schyradelia is a synonym of that name. e y as A u 98 Bonner zoologische Beitráge 54 (2005) Figs. 1-10. Porocallus insignis Sharp (1-4), P. hlavaci sp. n. (5-6), P. ligo sp. n. (7-8), and P. tianmuensis (Pace) (9-10): ligulae of specimen from the Russian Far East (1) and of holotype of Platysmarthrusa chinensis Pace (2); forebody (3, 5); habitus (7); elytra (4, 6, 8, 9); abdominal segments VI and VII in dorsal view (10). Scale bars: 3-9: 1.0 mm; 10: 0.5 mm; 1-2: 0.08 mm. Types examined: P. insignis: see ASSING (2001). 3.1. Porocallus insignis Sharp, 1888 (Figs. 1 - 4, 11, Map 1) P. chinensis: Holotype 3: China Beijing, Songshan [ca. 80 km NW Bejing city centre], 15.V.1993, G. de Porocallus insignis Sharp, 1888 (SHARP 1888: 287). Rougemont / Holotypus Platysmarthrusa chinensis m., det. R. Pace 1995 / Platysmarthrusa chinensis gen. n. sp. n. det. R. Pace 1995 / Porocallus insignis Sharp det. V. Assing 2003 (MHNG). Platysmarthrusa chinensis Pace, 1999 (PACE 1999: 108) syn. n. Volker ASSING: Porocallus Sharp: New synonyms and new species 99 i 16 20 Figs. 11-21. Porocallus insignis Sharp (11), P. hlavaci sp. n. (12-16), P. ligo sp. n. (17-20), and P. tianmuensis (Pace) (21): an- tenna (11, 12, 17, 21); median lobe of aedeagus in lateral and in ventral view (13, 14); male tergite VII (8); male sternite VIII (16); female tergite VIII (18); female sternite VIII (19); spermatheca (20). Scale bars: 11, 12, 17, 21: 1.0 mm; 13-16, 18-19: 0.5 mm; 20: 0.2 mm. Comments: A comparison of the holotype of P. chinen- sis with P. insignis from various regions revealed no differences suggesting that it should represent a distinct species. Consequently, P. chinensis is here placed in the synonymy of P. insignis. For illustrations of the aedeagi of a male from Japan and of the holotype of P. chinen- sis, see ASSING (2001, figs. 6-7) and PACE (1999, figs. 2-3), respectively. The type locality of P. chinensis is situated within the known range of P. insignis. The forebody, elytra, ligula, and antenna of this species are illustrated in Figures 1-4 and 11. Additional material examined: China: 13, Jiangxi prov., Wuyi Shan Nat. Res., Hu- angganshan, 1800-2050 m, 5.V1.2001, leg. Hlavác € Cooter (cAss). Distribution: In China, Porocallus insignis was previ- ously known from only two localities, one in Shaanxi (Qin Ling Shan) and one in Sichuan (Qingcheng Shan) (ASSING 2001). It is now known also from two localities in eastern China (Map 1). The specimen from Songshan was collected from moss and litter on the bank of a stream (Guillaume de Rougemont, Londinieres, pers. comm. 2003). 100 Bonner zoologische Beitráge 54 (2005) 3.2. Porocallus tianmuensis (Pace, 1999) comb. n. (Figs. 9-10, 21, Map 1) Ischyradelia tianmuensis Pace, 1999 (PACE 1999: 108 ff). Type examined: Holotype Í: China: Zhejiang Prov., Lin’an County, 1000 m, W. Tianmu Shan N. R. 18.V.1996, J. Cooter / Holotypus Ischyradelia tian- muensis m., det. R. Pace 1996 / Ischyradelia tianmuen- sis gen. n. sp. n. det. R. Pace 1996 / Porocallus tian- muensis (Pace) det. V. Assing 2003 (MHNG). Diagnosis: The species is somewhat similar to P. in- signis, but readily distinguished by the following char- acters: Antenna shorter; antennomeres III - X of similar length and approximately as wide as long (Fig. 21). Head, pro- notum, and elytra with relatively long, dense, erect pu- bescence; this pubescence visible especially at lateral margins of pronotum and elytra (Fig. 9). Elytra with ve- ry coarse and somewhat granulose puncturation (Fig. 9). Abdomen with very long and dense setae both on dor- sal, lateral, and ventral surfaces; tergite VII with rela- tively coarse puncturation (Fig. 10). Aedeagus with weakly pronounced crista apicalis and with apex of ven- tral process of characteristic shape especially in lateral view (see PACE 1999, figs. 8-9). Distribution: Porocallus tianmuensis is known only from the type locality in Zhejiang province, eastern China (Map 1). 3.3. Porocallus hlavaci sp. n. (Figs. 5-6, 12-16, Map 1) Type material: Holotype @: China: Fujian prov., Wuyi Shan Nat. Res., Sangan env. (900m), 30.v.-12.vi.2001, Hlavác & Cooter lgt. / Holotypus 4 Porocallus hlavaci sp. n. det. V. Assing 2003 (cAss). Description: 5.3 mm. Forebody as in Fig. 5. Head and pronotum blackish; elytra castaneous with the sides weakly infuscated and the scutellum blackish; abdomen blackish brown with the posterior margins of the seg- ments narrowly lighter; legs castaneous; antennae dark brown. Head about 1.15 times as wide as long (length measured from anterior margin of clypeus); eyes large and promi- nent; puncturation dense, coarse, and granulose; pubes- cence short and depressed, barely noticeable (Fig. 5). Antennae relatively short and massive; antennomere II short; IV - VI weakly transverse; X weakly oblong, and XI almost as long as the combined length of IX + X (Fig. 12). Pronotum 1.22 times as wide as long and 1.25 times as wide as head; puncturation similar to that of head; pu- bescence depressed and short, but slightly longer than that of head (Fig. 5). Elytra at suture almost as long as pronotum (Fig. 5); puncturation ill-defined, irregular and confluent every- where (Fig. 6). Hind wings fully developed. Legs rela- tively short; metatibia of holotype 0.98 mm; metatarsus very short, only slightly more than half the length of metatibia; metatarsomere I longer than the combined length of II - IV, but shorter than the combined length of MY. Abdomen without distinct microsculpture and, espe- cially anteriorly, not very sparse fine puncturation; pos- terior margin of tergite VII with palisade fringe. Í: posterior margin of tergite VIII weakly convex (Fig. 15), that of sternite VIII obtusely pointed (Fig. 16); me- dian lobe of aedeagus as in Figs. 13 - 14. 2: unknown. Etymology: | dedicate this species to Peter Hlavaé (Kosice), who, together with Jon Cooter, discovered it and to whom I am grateful for the generous gift of the holotype. Comparative notes: From P. insignis, which, too, was found in the Wuyi Shan Nature Reserve, P. hlavaci is readily distinguished especially by the shorter and more massive antenna, the slightly more finely punctate and completely matt appearance of the head and pronotum, the irregular, confluent, and ill-defined puncturation, the matt elytra, the shorter metatibia and metatarsus, the more numerous and longer setae at the sides of the ab- domen, and by the morphology of the aedeagus, espe- cially the different shape of the apex of the ventral proc- ess in lateral view. From P. tianmuensis, it is separated by the much more massive antenna with slightly oblong antennomeres X, the matter appearance of the forebody, the completely different puncturation of the elytra, the shorter metatibia and metatarsus, and by the morphol- ogy of the median lobe of the aedeagus. For distinction from the similar P. /igo see the following section. Distribution: The new species is currently known only from one locality in the Wuyi Shan, Fujian Sheng, in southeastern China (Map 1). 3.4. Porocallus lige sp. n. (Figs. 7 - 8, 17 - 20, Map 1) Type material: Holotype Y: China: Guizhou, Leishan Co., SE Kaili, NE Leishan, Leigong Shan, E-slope, 26°23.39’N 108°13.33’E / 2.5 km E of pass, 19.6.2001, ca. 1800 m, leg. Schillhammer (14) / Holotypus Y Po- rocallus ligo sp. n. det. V. Assing 2003 (NHMW). Description: 6.6 mm. Facies as in Fig. 7. Coloration as in P. hlavaci, but antennae reddish brown. Volker ASSING: Porocallus Sharp: New synonyms and new species 10] Map 1. Known distribution of the genus Porocallus Sharp: P. insignis Sharp (filled circles), P. hlavaci sp. n. (open circle), P. ligo sp. n. (large square), and P. tianmuensis (Pace) (small square). Head about 1.12 times as wide as long (length measured from anterior margin of clypeus); eyes large and promi- nent; puncturation dense, coarse, and granulose; pubes- cence short and depressed, barely noticeable. Antennae relatively long and slender; antennomeres IV-X all clearly oblong; XI shorter than the combined length of IX+X (Fig. 17). Pronotum about 1.25 times as wide as long and 1.25 ti- mes as wide as head (Fig. 7); puncturation similar to that of head: pubescence depressed and short, but longer than that of head. Elytra at suture almost as long as pronotum; punctura- tion ill-defined, irregular and confluent everywhere (Fig. 8). Hind wings fully developed. Legs slender; metatibia of holotype 1.18 mm; metatarsus relatively long and slender; metatarsomere I almost as long as the combined length of II - V. Abdomen shining; posterior margin of tergite VII with palisade fringe. 4: unknown. ©: posterior margins of tergite and sternite VIII weakly convex (Figs. 18 - 19); spermatheca with slender cap- sule (Fig. 20). Etymology: The name (Lat.) is a noun in apposition and refers to the hoe-like shape of the spermatheca. Comparative notes: This species is readily distin- guished from all its congeners by the slender antennae and by the shape of the spermatheca, from P. insignis and P. tianmuensis also by the completely matt fore- body and by the irregular confluent puncturation of the elytra. Distribution: Porocallus ligo is known from only one locality, the Leigong Shan in Guizhou Sheng, southern China (Map 1), where the holotype was found at an alti- tude of 1800 m. Its ovaries contained two mature eggs. 4. KEY TO THE SPECIES OF POROCALLUS 1. Antenna short and not conspicuously massive, an- tennomere X about as wide as long (Fig. 21). Elytra with very coarse and dense puncturation (Fig. 9). Forebody with conspicuously dense and (sub-)erect pubescence. Abdominal tergite VII with relatively coarse puncturation (Fig. 10). Aedeagus and sper- matheca as figured by PACE (1999: 110). China: Zhejiang (Map 1)..................... P. tianmuensis (Pace) — Antennae longer and/or conspicuously massive; an- tennomere X longer than wide. Elytra either with well-defined finer puncturation with shining intersti- 102 Bonner zoologische Beitráge 54 (2005) ces, or with ill-defined, irregular, confluent punctura- tion. Forebody with depressed and less dense pubes- cence. Abdominal tergite VII usually with finer puncturation. Aedeagus of different morphology..... 2 2. Elytra with well-defined puncturation and with shin- ing interstices (Fig. 4); head and pronotum with very weak shine (Fig. 3). Antenna as in Fig. 11. Aedeagus and spermatheca as figured by ASSING (2001, fig. 17). Widespread species known from Japan, the Rus- sian Far East, North Korea, and China (Map 1). ........ PT EDERTRLRR P. insignis Sharp — Elytra with ill-defined confluent puncturation; head and pronotum completely matt. Antenna either dis- tinctly shorter and more massive, or longer and more slender. Species known only from southern China... 3 3. Antennae long and siender, antennomeres IV-X all longer than wide (Fig. 17). Elytra with very subdued shine and with coarser puncturation (Fig. 8). Meta- tibia and metatarsus long and slender; first metatar- somere almost as long as the combined length of the remaining four tarsomeres. Spermatheca as in Fig. 20. China: Gouizhou (Map 1). ................ P. ligo sp. n. — Antennae short and massive, antennomeres IV-X as wide as long or weakly transverse (Fig. 12). Elytra without shine and with finer puncturation (Fig. 6). Metatarsus ‘ short; first metatarsomere distinctly shorter than the combined length of the remaining four tarsomeres. Aedeagus as in Figs. 13-14. China: Fanan (Map lineal P. hlavaci sp. n. Acknowledgements. I am much indebted to Giulio Cuc- codoro, Peter Hlavac, and Harald Schillhammer for the loan and gift, respectively, of the material which this study is based on. REFERENCES ASSING, V. 2001. On the systematics and distribution of Porocallus SHARP, 1888 (Coleoptera: Staphylinidae, Aleocharinae, Oxypodini). Beitráge zur Entomologie, Keltern 51: 215-221. PACE, R. 1999. Aleocharinae della Cina: Parte V (conclu- sione) (Coleoptera, Staphylinidae). Revue Suisse de Zoologie 106: 107-164. SHARP, D. 1888. The Staphylinidae of Japan. Annals and Magazine of Natural History, Series 6 2: 277-295. Author’s address: Dr. Volker ASSING, Gabelsbergerstr. 2, D-30163 Hannover, Germany; E-mail: vassing.hann@ t-online.de Received: 22.09.2003 Accepted: 10.12.2004 Corresponding editor: M. Schmitt Bonner zoologische Beitráge Band 54 (2005) | Heft 3 | Seiten 103—172 Bonn, August 2006 Atlas of the Microscopic Hair Structure of Southern African Shrews, Hedgehogs, Golden Moles and Elephant-shrews (Mammalia) f Michael F. SCHNEIDER & Victorino A. BURAMUGE University Eduardo Mondlane, Maputo, Mozambique Abstract. A hair atlas of 41 southern African Macroscelididae, Soricidae, Erinaceidae and Chrysochloridae species is presented and a simple key to identifying those species on the basis of hair characteristics is compiled. Light and elec- tron microscopic hair analyses revealed diagnostic species characteristics such as medullary structure, cross sectional shape and cuticular scale patterns. Additional characteristics like hair colouration, diameter and length as well as geo- graphical distribution have been considered. Hairs of all examined species of golden moles and elephant-shrews show sufficient specific characteristics to allow an unequivocal identification of the species. Shrew hairs, however, lack spe- cific features and in most cases only a definite identification to generic level is possible. The diagnostic criteria of simi- lar keys by other authors as well as limitations of the key are discussed. Key words. hair fine structure, identification key, hair atlas, small mammals 1. INTRODUCTION Microscopic analyses of mammalian hair find practical applications in law enforcement and forensic sciences (DE BOOM & DREYER 1953), epidemiology (KEOGH 1979), taxonomy (KEOGH 1975; DUCOMMUN et al. 1994), ecological studies (e.g. DAY 1966; BRUNNER & WALLIS 1986), species conservation, fur and textile in- dustry (WILDMAN 1954) and are also relevant for the identification of food contaminations (KEOGH 1979). The examination of hairs extracted from regurgitated pellets of birds of prey (MARZ 1987) and scats provide information on dietary habits of the respective predators and the existence of mammalian prey (e.g. DAY 1966; GILBERT & NANCEKIVELL 1982; BRUNNER & WALLIS 1986). Since hairs suffer lesser degradation from masti- cation and digestion than skulls and bones, hairs retain valuable characteristics and are often the only usable remains of an ingested prey species for identification (DAY 1966; BRUNNER & COMAN 1974). Hairs are usually composed of a cuticula with a layer of scales, the cortex containing pigmentary granules and the medulla (BRUNNER & COMAN 1974). The variations in shape, arrangement and type of these structures as well as characteristics such as hair diameter, distal tip tapering, size ratio between medulla and cortex, etc. are specific for certain taxonomic groups and thus allow the identification of the genus or species from a hair sample when compared to a reference system based on micro- scopic hair structure such as a hair atlas or an identifica- tion key (BRUNNER & COMAN 1974). Certain types of body hairs such as overhairs are of particular diagnostic importance because of the presence of such specific characteristics and structures. Comprehensive photographic reference systems and identification keys were developed for many mammal species, e. g. of Australia (LYNE & MCMAHON 1951; BRUNNER & COMAN 1974), Asia (AMERASINGHE 1986), northern America (MATHIAK 1938; MAYER 1952; ADORJAN & KOLENOSKY 1969) and Europe (DAY 1966; DZIURDZIK 1973, DEBROT et al. 1982; TEERINK 1991). These also exist for most southern African Bovidae (DREYER 1966; KEOGH 1983), Muridae and Cricetidae (KEOGH 1985) and mammals of the Andries Vosloo Kudu Reserve, South Africa (PERRIN & CAMPBELL 1980). However, photographic reference systems do not exist for small prey species such as shrews and elephant- shrews as well for most rodent families of southern Af- rica. The objective of the present study is to compile an atlas of the characteristics and structure of hairs of southern African Macroscelididae, Soricidae, Erinacei- dae and Chrysochloridae species as well as a key for their identification based on hair characteristics. 2. MATERIALS AND METHODS 2.1. Collection of reference hair specimens for identification Hairs were taken from dried skins of the Natural History Museum (MNH), Maputo, Mozambique, the collection of Scientific Services (NKW, KNP), Kruger National Park, Skukuza, South Africa and the Transvaal Museum 104 Bonner zoologische Beitráge 54 (2005) (TM), Pretoria, South Africa. Origin and specimen numbers are stated in the Hair Atlas. By means of ex- amination of the teeth, only adult individuals with fully developed pelage were chosen for hair sampling. Hair samples were taken from behind the neck on the back, from the flanks and the belly of the animal. In order to obtain entire unfragmented hairs, a tuft of hairs was clasped with forceps at the base and carefully pulled against the direction of the undisturbed hair. Hair samples were transferred on transparent sticky tape to maintain the original hair orientation and removed for examination by applying a drop of 70% ethanol on the hair sample. Forty-one species of southern African Macroscelididae (7 spp.), Soricidae (20 spp.), Erinaceidae (1 sp.), and Chrysochloridae (13 spp.) were examined. Hair samples of Chrysochloris visagiei, Chryptochloris zyli (Chry- sochloridae) and Elephantulus fuscus (Macroscelididae) were not available from the three collections and there- fore not examined. Taxonomic classification and data on distribution are based on HUTTERER (1993), SCHLITTER (1993), BRON- NER (1995) and BOITANI et al. (1999). 2.2. Microscopic analysis of hair samples Length measurement, description of hair profile and colour was made with the help of a stereo microscope (Wild 195040, Germany) and an electronic calliper (Mi- tutoyo Digimatic CD-6 CS, Japan). Whole mounts, cross sections and scale casts of hairs were prepared as described in great detail e. g. by BRUNNER & COMAN (1974) and KEOGH (1983, 1985) and examined using a microscope (Olympus T 041, Japan) with phase contrast condenser (Olympus ULW CD 0.30) and phase contrast objectives (Olympus S Plan 10x PL, LWD CD Plan 20x PL, LWD CD Plan 40x PL). Prior to examination, hair samples were cleaned in 70% ethanol and dried between absorbent tissue paper. For the study of medulla types and pigmentation, temporary mounts were made. Sample hairs were mounted in par- affin oil on cover slip (76 mm x 26 mm x | mm) and examined in phase contrast. Cross sections of the hairs were obtained by the plate method using a stainless steel slide (76 mm x 25 mm x 0.5 mm) with holes of 0.8 mm in diameter. A loop of nylon thread was pushed through a hole and several threads of bright cellulose acetate yarn (335 DTEX, Bembergcell S.P.A., Italy) inserted into the loop as packing material. The nylon thread was pulled through the hole until the cellulose acetate yarn formed a funnel- like structure. A hair sample was inserted into the centre of the “funnel” and pulled into the hole. The protruding parts of the hairs and packing material were cut off on both sides of the slide with a razor blade held at an an- gle of about 35°. Paraffin oil and a cover slip were placed on the side upon which the second cut was made and the obtained cross section was examined in phase contrast in order to study the shape of cross sections and medulla. Cuticular scale casts were prepared in order to study cu- ticular scale patterns. Individual hairs were placed on a thin layer of transparent nail polish on a glass slide. Af- ter drying, hairs were removed with forceps and the re- sulting casts examined in bright field. Cuticular scale patterns were also examined by the use of a scanning electron microscope. Hair samples were mounted on stubs, coated with gold for two minutes us- ing a sputter coater (Edwards S 150, Germany) at 20 mA and 2.0 x 10° Pa and scanned with an electron mi- croscope (Zeiss DSM 940A, Germany) at 15.0 KV. 2.3. Measurement of hair diameter and angle of distal hair tip Photographs of the square compartments (1/400 mm?) of a counting chamber (Neubauer Lumicyte, Germany) were taken to produce 50 um scales for all magnifica- tions used in microscopic analyses. The respective scales were used to measure the diameters of hairs on photographs. For the trigonometric determination of the angle of the distal tip of golden mole hairs, a right-angled triangle was drawn from the respective photographs, measuring 200 um from the tip of the hair (or the extrapolated point of intersection for hairs with blunt tip) to the right angle. 2.4. Documentation of results The results were documented with an analogous still camera (Olympus OM-2, Japan) attached to the micro- scope using colour negative film rated at 100 ISO. The aperture was chosen to obtain relatively long exposure times of about 10 seconds. Thus blurring due to vibra- tions caused by setting off the shutter could be mini- mized. During the study of whole mounts and cross sec- tions, additional drawings of hair profiles, medulla types and cross sections shapes were made. For the descrip- tion and classification of hair characteristics, a simpli- fied scheme was used based on WILLIAMS (1938), BRUNNER & COMAN (1974), PERRIN & CAMPBELL (1980) and KEOGH (1985). Michael F. SCHNEIDER & Victorino A. BURAMUGE: Atlas of Microscopic Hair 105 Fig. 1. Basic hair profiles of elephant-shrews (a), shrews (b), hedgehogs (c) and golden moles (d); bars equivalent to | mm. 2.5. Hair profiles Hair profiles were classified according Figure | and as suggested 1. a. by WILLIAMS (1938) and DREYER (1966). Heavy overhairs are usually long and stiff. Curly overhairs also referred to as shield hairs (BRUN- NER & COMAN 1974) or intermediate hairs (PERRIN & CAMPBELL 1980), possess a distally enlarged “shield” region, which is grooved in the case of shrews, golden moles and many Rodent species. The “shield” region of- ten bears specific characteristics and is therefore of di- agnostic importance. Furhairs, also referred to as under- hairs (BRUNNER & COMAN 1974; PERRIN & CAMPBELL 1980), are shorter, softer and more abundant than over- hairs and lack specific features. Other hair types like whiskers were not considered. Unless otherwise stated, the results refer to overhairs having a “shield” region. 2.6. Medulla types The medulla is the central part of a hair and is embed- ded in the cortex. The cortical cells, visible only with an electron microscope, grow faster than medullar cells, re- sulting in medullar air spaces (KEOGH 1983). These air spaces are intransparent for transmission light and thus appear dark, unless “infiltrated”, 1. e. the contained gas or air is substituted with a suitable medium. Medulla types were studied from whole mounts of hair and clas- sified according to BRUNNER & COMAN (1974), shown in Figure 2. The following medulla types characterize the species examined in the present study: Wide medulla lattice: This type is the common me- dulla type of the wider, distal part of golden mole hairs and of some elephant-shrews. The network or lattice is composed of shrunken medulla cells and mostly small, but in the case of a few golden mole species, larger en- closed air spaces. The lattice stretches across the entire diameter of the hair. i fragmental medulla 3 air’ east Ry h > Ld Bw h b » 6 \ Gn! Ya wide medulla narrow simple lattice medulla ar eayt wide simple medulla DDDDDDD adadaaı interrupted medulla different types of uniserial ladder Fig. 2. Medulla types in longitudinal view. Simple medulla: The medulla lacks an obvious struc- ture and can be narrow or wide. This type is common in hairs of hedgehogs and might be found at the base and tip of shrew hairs. Fragmental: In this type, the usually simple medulla is repeatedly interrupted by longer sections of cortical ma- terial. A fragmental medullary column can be often found at the base of hairs. 106 circular medium size medulla circular large medulla oval narrow medulla oval large medulla concavo-COonvex reniform with bilobed medulla Fig. 3. Shapes cross sections of widest part of shield hairs. Interrupted: The usually simple medulla is interrupted by one or several short sections of cortex material and appears more continuous than the fragmental medulla. Uniserial ladder: A uniserial ladder medulla is com- posed of one column of alternating rows of medullar material and air spaces. The “rungs” of the ladder are e1- ther regular with parallel elongate, spiral-like, “L” or crescent shaped rungs or irregular with rungs forming letters like “N”, “V” or “M”. Uniserial ladders are common in underhairs, along the entire overhairs of many small mammals like of shrews and most elephant- shrews and the basal part of golden mole overhairs (DAY 1966). 2.7. Cross sectional shape at widest part of hair The outer cortex and the central medulla characterize the internal structure of a hair, studied best on cross sec- tions. Unless containing pigment granules, the cortex is of little diagnostic value due to the invisibility of the cortical cells under the light microscope (BRUNNER & COMAN 1974). The cross sectional and medullar shapes, however, are important hair characteristics and were categorized as shown in Figure 3, following WILLIAMS (1938) and BRUNNER & COMAN (1974). According to the medulla width, the shapes can be further divided Bonner zoologische Beitráge 54 (2005) circular medulla absent circular small medulla oblong large medulla ra dumb-bell star-shaped H-shaped shaped oblong medium size medulla into large, medium size or small medulla or the medulla can be absent. Circular shapes are characteristic for cross sections of underhairs, thin basal sections of overhairs and the en- tire overhair of elephant-shrew hairs and hedgehogs. Oval and oblong shapes characterize cross sections of hairs of hedgehogs, some elephant-shrews and the be- ginning of the shield region (before the groove) of hairs of some golden moles. Concavo-convex shapes are typical for cross sections of the shield region of golden mole hairs. Reniform with bilobed medulla is the shape of cross section found in the wider distal part of overhairs of some elephant-shrews. The concave side is the grooved side. Dumb-bell shaped cross sections occur in the shield region of shield hairs of shrews and golden moles. The emarginations are the result of shallow longitudinal grooves on both sides of the hair. Star- and ‘H’-shaped cross sections are typical for the shield region of shrew hairs and may have deep emargi- nations and prominent extensions of the apices of one or both grooved sides. Michael F. SCHNEIDER & Victorino A. BURAMUGE: Atlas of Microscopic Hair 107 a) form of scale margins Mere re NN - smooth crenate rippled dentate b) distance between scale margins | = distant near ==> TZ = wae == c) scale patterns AY E simple diamond narrow diamond club shaped cupped coronal petal petal regular wave irregular wave double chevron single chevron transitional Fig. 4. Cuticular scale patterns. 2.8. Shapes and arrangement of cuticular scales The cuticle is composed of keratinised overlapping scales, whose patterns, shape, size and types of margins are of diagnostic importance. The classification of scale margins according to BRUNNER & COMAN (1974) refers to the free distal margin of an individual scale according to Figures 4a, b. Smooth margins form a straight line without indenta- tions. Crenate margins have shallow but pointed indenta- tions. Rippled margins are characterized by deeper but rounded indentations. Dentate margins have tooth-like projections and are typical for petal scales. The distance between scale margins is characterized by the ratio between width and length of a scale: Distant scale margins are those with a width of no more than three times of the length. Near scale margins have a ratio between three and eight. Close scale margins have a width of more than eight times of the length. 2.9. Cuticular scale patterns Scale patterns were classified according to Figure 4c following WILLIAMS (1938), BRUNNER & COMAN (1974), PERRIN & CAMPBELL (1980) and KEOGH (1985). The patterns of Lipotyphla, Chrysochloridea and Mac- roscelidea can be divided into: 108 Bonner zoologische Beitráge 54 (2005) Petal scale patterns are formed by a series of overlap- ping scales with the free distal scale margin slightly raised above the following scale. Coronal scale patterns are the common type of the slender base of hairs. They are made of a single or two scales that completely encir- cle the entire shaft of a hair. The scale margins are smooth and distant. Diamond petals are the common scale patterns of the basal part of shrew and elephant- shrew and golden mole hairs, following the coronal hair base. The scale margins are dentate and distant. Club- shaped scales are made of a single or two scales reach- ing across the entire width of a hair and commonly found at the basal part of golden mole hairs. The scale margins are smooth and distant. Cupped scale patterns characterize the grooved wider part of overhairs of shrews. This type is made of single scales reaching across the entire diameter. The scale margins are smooth and usually near. Transitional scale patterns occur at the transition zone between the thinner basal (often diamond petal) and the wider distal part of hairs. Waved scale patterns commonly occur along the distal wider part of hairs. Regular waves usually have smooth and near scale margins. Irregular waves have rippled or crenate margins with deeper emarginations, the dis- tance between the scale margins can be near or close. Chevron scale patterns have more or less regular, paral- lel scale margins with one (single) or two (double) ‘V’- like crests and are composed of two or more scales. This type can be found in the thinner basal part of some ele- phant-shrew and golden mole hairs. 3. RESULTS AND DISCUSSION This Hair Atlas provides a key for the identification of 41 species of southern African Lipotyphla, Chrysochlo- ridea and Macroscelidea based on their hair characteris- tics. Hair characteristics allowing definite species-level identification was found for the hedgehog A/elerix fron- talis, some Macroscelid species (Petrodromus tetradac- tvlus, Macroscelides proboscideus, Elephantulus brachyrhynchus) and some golden moles species (Chrysopalax spp., Amblysomus spp., Chrysochloris asiatica, Calcochloris obtusirostris, Cryptochloris win- toni). The unmistakable identification of all Mac- roscelidea and all golden moles was possible, when geographic distribution data were used in addition to hair characteristics. However, some shrew species were not identified unequivocally, even when combining hair characteristics with geographic distribution. Other authors also reported of the unavailability of spe- cific hair characteristics for some species (PERRIN & CAMPBELL 1980; KEOGH 1985). Therefore, it has been suggested that a more reliable identification can be ob- tained by combining geographic distribution of a spe- cies with hair characteristics (BRUNNER & COMAN 1974; PERRIN & CAMPBELL 1980). Most species examined in the present study showed common generic characteristics, e. g. the hair length of the shrew genera Myosorex (7 mm to 9 mm), of Suncus (3 mm to 6 mm) and of Sylvisorex (about 7 mm). Al- most all species of Crocidura have hairs with coronal scale patterns at their distal tip whereas this characteris- tic 1s less consistent in the other examined shrew genera. Additionally, shield hairs of the genus Myosorex usually have five constrictions and the hair length was more homogenous than in other shrew genera. Some genera of golden moles also showed distinct genus specific fea- tures, e.g., much longer hairs in the case of Chrysopalax. Within the elephant-shrews, the medulla type was not specific for the genus Elephantulus. Whole mounts of overhairs of E. myurus and E. brachyrhyn- chus revealed medulla lattice, whereas all other species of this genus had a medulla with uniserial ladder. Hair characteristics of the four studied families are al- ways very typical and allow a definite identification of the family. Such family specific characteristics for in- stance were the hair profile and the shape of hair cross sections for golden moles and shrews as well as the cupped cuticular scale patterns of the distal part of overhairs of all shrew species. This is in accordance with specific features of moles and shrews found by WILLIAMS (1938) and shrews described by DANNELID (1986). The results of the present study are also in con- formity with the study of the hair morphology of the shield region of curly overhairs of various shrew genera (DUCOMMUN et al. 1994). Shallow V- or U-shaped notches were also found along the shield region of Cro- cidura and Suncus in the present study (e.g., Pl. 8N, 9N, 10K, 11N, 12N and Pl. 25N & O, 26L), whereas the Myosorex shield is deeply grooved (Pl. 20F, 21G, 22N). Additionally, both studies show similar scale patterns on the shield region of C. luna and Sylvisorex megalura. Hair characteristics of some of the species included in the present study have already been described by PERRIN & CAMPBELL (1980). Apart from using different terms to classify hair profiles and scale patterns, the study of these authors was mainly based on cuticular scale patterns. Despite the differences in methodology, both studies show similar results; e.g., for Atelerix fron- talis, Crocidura cyanea and C. flavescens. Regarding Michael F. SCHNEIDER & Victorino A. BURAMUGE: Atlas of Microscopic Hair 109 the study of PERRIN & CAMPBELL (1980), the petal scale pattern in the fine region and the flattened mosaic at the midpoint of the shield region of Macroscelides probos- cideus hair correspond with the transitional scale pattern between the narrow diamond petal and regular wave patterns (distal part of Pl. 6 M) and the regular scale pat- tern (Pl. 6 N) of the present study. PERRIN & CAMPBELL (1980) solely used cuticular scale patterns of the shield region to differentiate between golden moles. However, in the present study additional characteristics like cross sections and geographical distribution were used to dis- criminate more reliably between these species. The hair characteristics used in the present study are based on the longitudinal view of the medulla, cross sectional shape of hair and medulla, medulla and cortex pigmentation, cuticular scale pattern, angle of distal hair tip, constrictions of hairs as well as hair profiles, colouration, length, minimum and maximum diameter. The consideration and use of certain hair characteristics is controversial and largely depends on the author. In the present study a combination of various hair charac- teristics along the entire hair was studied in order to overcome the shortcomings of the one or the other fea- ture. Regarding characteristics related to the medulla struc- ture, KEOGH (1983) argues that the medulla varies con- siderably along the length of the hair and between indi- viduals of the same species, and therefore this feature was not included in her study. On the other hand, WIL- LIAMS (1938), DAY (1966) and BRUNNER & COMAN (1974) stress the importance of a number of medulla characteristics. Some authors (BRUNNER & COMAN 1974; KEOGH 1983) even include the “infiltrated” me- dulla in which the air or gas of the air spaces has been substituted with mounting medium. In the present study, a simplified classification system of the longitudinal view of the medulla according to BRUNNER & COMAN (1974) has proven to be of diagnostic value, particularly in combination with other hair characteristics. The cross sectional shape of a hair and the medulla are widely used in hair analyses (WILLIAMS, 1938; DREYER 1966; BRUNNER & COMAN 1974; KEOGH 1983). As de- scribed by BRUNNER & COMAN (1974) and found in the present study, cross sectional shapes in particular are prominent features characterizing hair of shrews and golden moles. Furthermore, with the help of cross sec- tions the longitudinal grooves along the shield region of shield hairs of shrews and golden moles can be easily studied. Grooves were also considered of diagnostic im- portance for the identification of Cricetidae and Muri- dae by PERRIN & CAMPBELL (1980) and KEOGH (1985). However, the preparation of cross Sections is delicate and requires a skilled operator (BRUNNER & COMAN 1974), particularly when dealing with short hairs as those of shrews. Cuticular scale patterns are another diagnostic feature used in the present and many other studies (e.g., DREYER 1966; BRUNNER & COMAN 1974; PERRIN & CAMPBELL 1980; KEOGH 1983; KEOGH 1985). Accord- ing to DAY (1966), cuticular scale patterns of the basal region of hairs are characteristic and thus useful for hair identification. In the present study, however, scale pat- terns were rarely found to be species-specific but were in most cases characteristic for genera or the entire fam- ily. Even electron microscopic scans did not reveal more detail of the cuticular scales than did light micros- copy, as the comparison shows (PI. 42). Therefore, the much faster, cheaper and simpler light microscopy was used for the examination of scale patterns. Even though the use of hair length is a controversial characteristic, length together with minimum and maximum hair diameter were used in the present study and specifically for some of the golden mole species and shrew genera. Hair length was also used by MATHIAK (1938), MAYER (1952) and KEOGH (1983 & 1985), but since this feature might be subject to large individual variations, DREYER (1966) and other authors only considered the minimum and maximum hair di- ameter. For similar reasons, pigmentation of hairs has not been used widely as a characteristic; however, there are a number of authors employing this feature (e.g., DREYER 1966; KEOGH 1983). According to BRUNNER & COMAN (1974), hair pigmentation is not of primary diagnostic importance, but can be used to confirm an identification made on the basis of other features. In the present study, colouration of entire hairs as well as of the cortex and medulla in cross sections were considered, despite the fact that colour might vary according to the age of the animal and the season of the year (PERRIN & CAMPBELL 1980). In order to minimize colour variations of differ- ent body parts, hairs from the back, flanks and belly of the animal were described. Additionally, colour charts can be used to minimize subjective interpretation of the hair colour (MAYER 1952). Identification keys based on hair characteristics have a number of limitations (e.g., DAY 1966; BRUNNER & COMAN 1974; PERRIN & CAMPBELL 1980). The most criticised shortfalls are the variations of hair characteris- tics with age, sex, diet, body part, geographic origin, season of the year, etc. 110 Bonner zoologische Beitráge 54 (2005) BRUNNER & COMAN (1974) studied the different stages of hair development and showed that the hair, once pro- truded above the skin, 1s fully differentiated and no longer subject to further morphological changes. KEOGH (1975) found that season, sex and diet had no effect on cuticular scale patterns of rodents. According to the same author, variations of scale patterns solely de- pended on the age of an individual and after six months of age, the scale pattern remained constant. According to BRUNNER & COMAN (1974), the development of the pelage of Rattus norvegicus 1s completed after three months of age. A comprehensive study of the Bovidae Tragelaphus strepsiceros, Aepyceros melampus, Sylvi- capra grimmia and Gorgon taurinus did not reveal any differences in cuticular scale patterns, pigmentation and hair profiles between adult males and females (DREYER 1966). However, this author found higher variations of the cuticular scale patterns in subadult Kudus. Different to this, DAY (1966) noticed that subadults of small mammals showed similar cuticular scale patterns to adult animals, but these patterns were expressed in a simplified form. In order to minimize the influence of age on certain hair characteristics, only adult animals were used in the present study for hair sampling and the subsequent hair analyses. Morphological differences between hair types and sam- ples taken from different body parts can be considerable and are particularly pronounced in whiskers, ornamental hairs, hairs of legs, tail, etc. (DE BOOM & DRYER 1953; DAY 1966; BRUNNER & COMAN 1974). Hairs of the body trunk, however, show more uniform characteristics and therefore are more suitable for hair studies. As done in the present study, the difficulty arising from varia- tions in hairs of different body parts can be overcome by including hairs of several parts of the trunk. In the pre- sent study hair samples of different body parts com- monly vary in colour. A hair characteristic typical for hair of a certain part of the trunk could be found only in a single instance, the concavo-convex cross sections of ventral hairs of Chrysochloris asiatica (Pl. 35 D). Individual morphological variations of hair structure render an unequivocal identification of a certain species more difficult (BRUNNER & COMAN 1974). Therefore, PERRIN & CAMPBELL (1980) suggest the sampling of hairs derived from different individuals of the same spe- cies. Additionally, hair development and the resulting hair characteristics can vary with the geographic origin of a hair sample, particularly with the prevailing climate and habitat (e.g., DREYER 1966; PERRIN & CAMPBELL 1980) so that samples of various origins are recom- mended (PERRIN & CAMPBELL 1980). The process of digestion usually causes little alteration of the hair structure of a consumed mammal (BRUNNER & WALLIS 1986) and thus, photographic reference sys- tems and identification keys can be based on hair sam- ples; e. g., taken from museum skins (BRUNNER & CO- MAN 1974). DAY (1966) suggests the additional analysis of teeth and skeletal remains contained in predator scats would confirm the identification of the prey species based on hair examinations. Finally, the revision of the genus Amblysomus (BRON- NER 1995) is supported by the findings of the present study: A. hottentotus and A. iris differ from Neambly- somus gunningi and N. julianae in terms of hair diame- ter and shape of cross section. The same author also separated Carpitalpa arendsi from the genus Chloro- talpa. Even though having some similar hair character- istics such as maximum diameter and shape of cross section, C. arendsi has double chevron scale patterns and thus differs from Chlorotalpa duthieae and C. sclateri, as shown by the present study. The comparison of hair samples of Crocidura bicolor with C. fus- comurina showed identical hair characteristics and therefore supports the assemblage of the two species as C. fuscomurina (HUTTERER 1993). The revision of the formerly separated species Crocidura olivieri occiden- talis and C. olivieri martiensseni (HUTTERER 1993), is not supported by the findings of the present studies, since the examined hairs differ quite considerably in hair length, diameter and scale patterns. 4. HAIR ATLAS OF SOUTHERN AFRICAN ELEPHANT-SHREWS, SHREWS, HEDGEHOGS AND GOLDEN MOLES 4.1. Order Macroscelidea Family Macroscelididae (elephant-shrews, sengis) Elephant-shrews have long overhairs that are wider at their distal part (Fig. la). Underhairs are long, soft, sometimes constricted and more abundant than over- hairs. Hairs occasionally have an intumescence at the distal part. Cross sections of hairs are almost always cir- cular, the medulla has a uniserial ladder and some spe- cies have wide lattice at the distal part of hair. Scale pat- terns of distal hair part are regular or irregular waves, the distal tip often is of coronal type. Unless indicated, the described hair characteristics refer to both types of hairs. Elephantulus brachyrhynchus (A. Smith, 1836) Short- snouted elephant-shrew (PI. 1) Michael F. SCHNEIDER & Victorino A. BURAMUGE: Atlas of Microscopic Hair 11] Origin of hair sample: MHN: junction Sabi-Lundi south bank, Mozambique (Nasilio brachyrhynchus), NKW 4: Mwambia Pan, KNP, South Africa. Hair colour: Dorsal hairs dark red at base and dark brown at the distal part; hairs of the flanks dark at base and dark red at the distal end; ventral hairs dark grey basally and brownish-yellow at distal part. Hair types, shape and length: Overhairs long and thick, underhairs relatively thin and constricted. Hairs sometimes with intumescences (F, G) at distal part. Length of overhairs about 15.5 + 1.1 mm (n = 9); maximum hair diameter 58 um, minimum diameter about 19 um. Description of whole mounts: Overhairs basally with uniserial ladder (B, C), wide medulla lattice at distal part (D-F) and distally ending with simple medulla (H). Generally, the distal parts have transparent and slightly thickened areas (D, F). Underhairs with uniserial ladder with crescent-shaped rungs distally (A). Form of scale margins: Basally smooth (J, K), dentate (L-N), smooth (O), crenate (P) and smooth (Q-S) at dis- tal hair tip. Distance between scale margins: Scale margins near at base (basal part of J), distant (distal part of J-N), near (O) and close (P-S) at distal hair tip. Cuticular scale patterns: Basally simple coronal scale patterns (J, K), narrow diamond petal (L, M), diamond petal/transitional (N), regular waves (O), irregular waves (P) and regular waves (Q-S) at distal hair tip. Description of cross sections: Circular cross sections with large medulla (la) or medium size medulla (Ib); medulla always dark; cortex of some cross sections or- ange or gold, resulting in an intense brilliant sheen of the cortex in transmitted light of microscope (Ib); cortex and medulla of other cross sections dark (Ic). Elephantulus edwardi (A. Smith, 1839) Cape rock ele- phant-shrew (PI. 2) Origin of hair sample: TM 687: Hannover, Central Prov., South Africa. Hair colour: Dorsal hairs dark grey basally and light brown at distal part; hairs of flanks and belly basally grey and light brown at distal part. Hair types, shape and length: Overhairs relatively long and thick and less abundant than underhairs; un- derhairs thin and sometimes constricted. Length of overhairs about 14.3 + 0.8 mm (n = 4); maximum hair diameter 30 um, minimum diameter about 15 um. , Description of whole mounts: Medulla with uniserial ladder (A-E) along entire hair, distally rungs crescent- like (C) and L-shaped (D). Form of scale margins: Smooth at base (G, H), dentate (I, basal part of J) and smooth (distal part of J-N) at dis- tal hair tip. Distance between scale margins: At base near (basal part of G), distant scale margins (distal part of G, H-J) and near (K-N) at distal hair tip. Cuticular scale patterns: Basally simple coronal scale patterns (G, H), narrow diamond petal (I, J), transitional (distal part of J), regular waves (K-M) and coronal (N) at distal hair tip. Description of cross sections: Circular shapes of cross sections with medium size to large medulla (Fa); me- dulla and cortex of some cross sections light brown, some cross sections with lighter medulla and darker cor- tex (Fa) or vice versa, or medulla and cortex of other cross sections black (Fb). Elephantulus intufi (A. Smith, 1836) Bushveld ele- phant-shrew (Pl. 3) Origin of hair sample: TM 15239: Nguia, Botswana Reserve, Botswana. Hair colour: Dorsal hairs dark grey at basal part and dark red distally; hairs of flanks and belly dark grey at basal part and light red distally. Hair types, shape and length: Overhairs relatively long and wider along distal part; underhairs thinner and sometimes constricted. Hairs sometimes with intumes- cence (D) at distal part. Length of overhairs about 14.7 + 0.5 mm (n = 6); maximum hair diameter 53 um, minimum diameter about 21 um. Description of whole mounts: Overhairs basally with uniserial ladder /simple medulla (B, C) and simple at distal part (D-F). Medulla of underhairs with uniserial ladder with spiral-like rungs (A). Form of scale margins: Smooth at base (H, I) dentate (J, basal part of K) smooth/crenate (distal part of K, L, M) and smooth (N, O) at distal hair tip. Distance between scale margins: Scale margins near at base (H), distant (distal part of H, I-K) and near (L-O) at distal hair tip. 112 Bonner zoologische Beitráge 54 (2005) Cuticular scale patterns: Basally simple coronal scale patterns (H and I), narrow diamond petal (J), diamond petal/transitional (K), regular waves (L, M) and coronal (N, O) at distal hair tip. Description of cross sections: Circular shapes of cross sections with medium size medulla or large medulla (Gc); medulla and cortex of some cross sections light brownish-grey (Gb, c), medulla and cortex of other cross sections dark (Ga). Elephantulus myurus Thomas & Schwann, 1906 Rock elephant-shrew (Pl. 4) Origin of hair sample: NKW 18: Pafuri, KNP, South Africa; NHM: junction Sabi-Lundi, south bank, Mo- zambique; TM 39049: Moshanens, Botswana. Hair colour: Dorsal hairs grey at basal part and dark red distally; hairs of flanks grey at basal part and dis- tally dark red with a brown hue; ventral hairs grey at basal part and light yellow distally. Hair types, shape and length: Overhairs relatively long, underhairs thinner and constricted. Hairs some- times with intumescences along distal part (F). Length of overhairs about 12.0 mm + 0.8 (n = 9); maximum hair diameter 46 um, minimum diameter about 15 um. Description of whole mounts: Overhairs basally with uniserial ladder (C, D), wide medulla lattice (E) and simple medulla (F, G) at distal part and occasionally with protuberances at distal part (F). Underhairs of uniserial ladder along the entire hair (A, B), sometimes constricted (A). Form of scale margins: Smooth (H, I) at base, dentate (J) and smooth (K-N) at distal hair tip. Distance between scale margins: Scale margins near at base (H), distant (distal part of H, I, J), near (K) and close (L-N) at distal part of hairs. Cuticular scale patterns: Basally simple coronal scale patterns (H, I), narrow diamond petal (J), regular waves (K-M) and coronal (N) at distal hair tip. Description of cross sections: Circular shapes of cross sections with large medulla (Oa); medulla and cortex of some cross sections light brown (Oa), medulla and cor- tex of other cross sections dark (Ob). Elephantulus rupestris (A. Smith, 1830) Smith’s or Western rock elephant-shrew (PI. 5) Origin of hair sample: TM 10218: Tsabis, West of Ro- chboth, Botswana. Hair colour: Dorsal hairs grey at basal part and dark red distally; hairs of flanks grey at basal part and light red distally; ventral hairs dark grey at basal part and light yellow distally. Hair types, shape and length: Overhairs relatively long and distal part wider; underhairs thinner and quite long. Length of overhairs about 18.2 + 0.8 mm (n = 6); maximum hair diameter 34 um, minimum diameter about 18 um. Description of whole mounts: Medulla of underhairs (A) and overhairs (B-H) with uniserial ladder along the entire hair; rungs distally crescent-shaped (F, G, H) and occasionally with transparent parts (D, F); distal tip without medulla (H). Form of scale margins: Smooth at base (J-L), dentate (M-O), smooth (P, Q), crenate (R) and smooth (S) at distal hair tip. Distance between scale margins: Scale margins near (J) at base, distant (K-P) and near (Q-S) at distal hair tip. Cuticular scale patterns: Basally simple coronal scale patterns (J-L), narrow diamond petal (M-O), transitional (P), regular waves (Q, R) and coronal (S) at distal hair tip. Description of cross sections: Circular shapes of cross sections with medium size (la) to large medulla (Ib); medulla and cortex of some cross sections light brown, resulting in a weak golden sheen of the cortex in trans- mitted light of microscope (Ib), some cross sections with more or less transparent cortex and dark medulla (la) or dark cortex and medulla. Macroscelides proboscideus (Shaw, 1800) Round-eared or short-eared elephant-shrew (Pl. 6) Origin of hair sample: TM 28146: Faron Kongras, Springbok Game Reserve, Transvaal, South Africa. Hair colour: Dorsal and lateral hairs dark grey at basal part and dark red distally; ventral hairs dark grey at basal part and light yellow distally. Hair types, shape and length: Hairs relatively long and thin, distal part of overhairs wider, underhairs thin- ner. Hairs sometimes with intumescence (F) at distal part. Length of overhairs about 18.1 + 0.9 mm (n = 4); maximum hair diameter 26 tm, minimum diameter about 9 um. | Michael F. SCHNEIDER & Victorino A. BURAMUGE: Atlas of Microscopic Hair 113 Description of whole mounts: Medulla of all hairs with uniserial ladder (A-F) along the entire hair; distally rungs “L”-shaped (D, E), occasionally with transparent parts (C), with protuberances (F) and with intransparent distal hair tip (G). Form of scale margins: Smooth (I, J) at base, dentate (K, L, basal part of M) and smooth (distal part of M-Q) at distal hair tip. Distance between scale margins: Scale margins near at base (I), distant (distal part of I-M), near (N, O) and close (P, Q) at distal hair tip. Cuticular scale patterns: Basally simple coronal scale patterns (I, J), narrow diamond petal (K-L), transitional (M), regular waves (N-P) and coronal (Q) at distal hair tip. Description of cross sections: Circular shapes of cross sections with medium size medulla or large medulla (Ha); medulla and cortex of some cross sections trans- parent (Hb) or with more or less transparent cortex and dark medulla, resulting in a weak sheen of the cortex in transmitted light of microscope (Ha); some cross sec- tions with dark cortex and medulla. Petrodromus tetradactylus Peters, 1846 Four-toed ele- phant-shrew (PI. 7) Origin of hair sample: NKW10: Tussen, Magabene, Makembane KNP, South Africa; NHM: north bank Sabi, Mozambique. Hair colour: Dorsal hairs grey at basal part and dark red distally; hairs of flanks grey at basal part and red with a yellow hue distally; ventral hairs vary from dark to light red. Hair types, shape and length: Overhairs relatively long and thick, distal part wider; underhairs thinner and occasionally with constrictions. Hairs sometimes with intumescence (B, F) at distal part. Length of overhairs about 17.5 + 0.5 mm (n = 9); maximum hair diameter 59 um, minimum diameter about 21 um. Description of whole mounts: Medulla at base of over- hairs with uniserial ladder with oblique, “V-shaped rungs (E, G); further distally, rungs resembling “letters” (H) and with simple medulla towards distal hair tip (1, J). Medulla of underhairs with uniserial ladder (A-C), at distal part with crescent-shaped rungs (C). Form of scale margins: Smooth at base (L), dentate (M-O), smooth/crenate (P), rippled (Q) and smooth (R, S) at distal hair tip. Distance between scale margins: Scale margins near (L) at base, distant (distal part of L-O) and close (P-S) at distal hair tip. Cuticular scale patterns: At base simple coronal scale patterns (L), narrow diamond petal (M, N); diamond petal/transitional (O), regular waves (P), irregular waves (Q) and regular waves (R, S) at distal hair tip. Description of cross sections: Circular shapes of cross sections with large medulla (Ka) or medium size me- dulla (Kb); medulla and cortex of some cross sections light brown, resulting in an intense brilliant sheen of the cortex in transmitted light of microscope (Ka, b); other cross sections dark medulla and dark cortex (Kc). 4.2. Order Soricomorpha Family Soricidae (shrews) The pelage of shrews is soft and plush-like, the hairs (Fig. 1b) are relatively short. Curled overhairs (shield hairs) usually have one or two constrictions and a flat- tened, expanded distal part of the hair with a pro- nounced longitudinal groove on one or both surfaces. Heavy overhairs are straight, longer, lack constrictions and are rare in number. Underhairs have a thin distal end and several constrictions, at which the hair changes its direction (Pl. 13 H), resulting in a zigzag-like shape. Cross sections of hairs are typically rectangular or quad- rangular with emarginations of several sides resulting in a dumb-bell shaped, “H”-like or star-like outline (Fig. 3); the medulla has uniserial ladder. Scale patterns of grooved part of shield region are always cupped, the hair tip has with coronal or regular waves. Unless indi- cated, the described hair characteristics refer to over- hairs with shield region (shield hair). Crocidura cyanea (Duvernoy, 1838) Reddish-grey musk shrew (Pl. 8) Origin of hair sample: NK W 2: Malelane, KNP, South Africa. Hair colour: Dorsal and lateral hairs light grey at basal part and greyish-brown distally; ventral hairs light grey. Hair types, shape and length: Heavy overhairs straight and without constrictions; curled overhairs (shield hairs) with thickened and grooved distal part and two constric- tions; underhairs thinner and undulating at constrictions. Length of overhairs about 6.2 + 0.5 mm (n = 7): maxi- mum diameter of shield region 29 um, minimum diame- ter about 7 Lum. 114 Bonner zoologische Beitráge 54 (2005) Description of whole mounts: At base simple medulla (A), further distally uniserial ladder (B-F) with regular and straight rungs (D, E) or less regular, ‘mushroom’ -shaped rungs (F) ending in a transparent distal hair tip (G). Form of scale margins: Smooth (I, J) at base, dentate (K, L), smooth (M, N), crenate (O, P) and smooth (Q) at distal hair tip. Distance between scale margins: Scale margins at base close, further distally distant (I-M), near (N) and close (O-Q) at distal hairs tip. Cuticular scale patterns: At base simple coronal (I, J), further distally narrow diamond petal (K), diamond petal (L), regular waves (M), cupped (N), irregular waves (O, P) and coronal (Q) at distal hair tip. Description of cross sections: Rectangular shaped out- line of cross section with slightly invaginated sides (Ha), “H”-shaped outline with more or less pronounced extensions (Hb), both with large medulla, other outlines of cross sections circular with medium size medulla (Hc); medulla black or grey, cortex brown, light or dark grey. Crocidura flavescens (Geoffroy, 1827) Greater red musk shrew (PI. 9) Origin of hair sample: TM 9000: Krysna Distr. Deep- walls, Cape Province, South Africa. Hair colour: Hairs light grey at basal part and dark brown to greyish-brown distally. Hair types, shape and length: Heavy overhairs straight and without constrictions; curled overhairs (shield hairs) with thickened and grooved distal part and two or three constrictions; underhairs thinner and undulating at con- strictions. Length of overhairs about 5.5 + 0.2 mm (n = 5); maximum diameter of shield region 35 um, mini- mum diameter about 6 pum. Description of whole mounts: At base simple medulla (A, B); further distally uniserial ladder (C-E), some- times irregular forming “letters” (D, E) and ending in a thin transparent distal tip (F). Form of scale margins: At base smooth (H, I), dentate (J-L), smooth (M, N) and rippled (O, P) at distal hair tip. Distance between scale margins: Scale margins near (H) at base, distant (I-L), near (M, N) and close (O, P) at distal hair tip. Cuticular scale patterns: At base simple coronal (H, I), further distally narrow diamond petal (J, K), diamond petal (L), transitional (M), cupped (N) and irregular waves (O, P) at distal hair tip. Description of cross sections: Cross sections predomi- nantly rectangular shaped with slight emarginations of all sides resulting in short extensions of one or both short sides (Ga), dumb-bell shaped (Gb), trapezoidal (Gc, d) or circular outlines (Ge, Qa); all cross sections with medium size to large medulla; medulla black or grey, cortex brown, light or dark grey, sometimes with a brilliant sheen in transmitted light of microscope (G). Crocidura fuscomurina (Heuglin, 1865) Tiny musk shrew (PI. 10) Origin of hair sample: NKW 1: Skukuza, KNP, South Africa; TM 45738: Vryburg Distr., Mopolo Nat. Res., North West Province, South Africa; TM 37158 (C. bi- color): Pietermaritzburg Distr., Natal, South Africa. Hair colour: Hairs dark grey at basal part and light brown to dark brown distally. Hair types, shape and length: Heavy overhairs straight and without constrictions; curled overhairs (shield hairs) with thickened and grooved distal part and one or two constrictions; underhairs thinner and undulating at con- strictions. Length of overhairs about 2.7 + 0.1 mm (n = 5); maximum diameter of shield region 26 um, minimum diameter about 6 um. Description of whole mounts: At base simple medulla (A); further distally uniserial ladder (B-D), at distal part with slightly oblique rungs (C, D), ending in a thin transparent distal hair tip (E). Form of scale margins: Smooth at base (G, H), further distally smooth/crenate (I-K), rippled (L) and smooth (M) at distal hair tip. Distance between scale margins: Scale margins near (basal part of G), distant (distal part of G, H) at base, near (I, J) and close (K-M) at distal hair tip. Cuticular scale patterns: At base simple coronal (G, H), at distal part regular waves (I, J), cupped (K), ir- regular waves (L) and coronal (M) at distal hair tip. Description of cross sections: Cross sections predomi- nantly quadrangular or rectangular shaped with slight emarginations of the short sides resulting in prominent extensions of these sides (Fa) or circular (Fb) shapes in outline; all cross sections with small or medium size medulla; medulla black or grey, cortex brown, light or dark grey. Michael F. SCHNEIDER & Victorino A. BURAMUGE: Atlas of Microscopic Hair 115 Crocidura hirta Peters, 1852 Lesser red musk shrew (aM) Origin of hair sample: NK W 13: Pafuri, KNP, South Africa; MHN: Maringa, Sabi north bank, Mozambique. Hair colour: Hairs dark grey at basal part and light brown to brownish-grey distally. Hair types, shape and length: Heavy overhairs straight and without constrictions; curled overhairs (shield hairs) with thickened and grooved distal part and three con- strictions; underhairs thinner and undulating at constric- tions. Length of overhairs about 5.5 + 0.2 mm (n = 7); maximum diameter of shield region 33 um, minimum diameter about 6 um. Description of whole mounts: At base simple medulla (A), further distally uniserial ladder (B-G), rungs some- times irregular and “V”-shaped (E) or “mushroom”-like (D, F), ending in a thin transparent distal hair tip (G). Form of scale margins: Smooth at base (I), further dis- tally dentate (J, K), smooth (L), rippled (M, N), crenate (O) and smooth (P) at distal hair tip. Distance between scale margins: Scale margins at base close; distant (I-L), near (M, N) and close (O, P) at distal hair tip. Cuticular scale patterns: At base simple coronal (1), further distally narrow diamond petal (J, K); transitional (L), cupped (M), irregular waves (N, O) and coronal (P) at distal hair tip. Description of cross sections: Cross sections dumb- bell shaped in outline with emarginations of the long sides with prominent extensions (Ha) or round short sides (Hb) and circular in outline (Hc); cross sections with medium size to large medulla, sometimes emargi- nated, reflecting the shape of the cross section; medulla black or grey, cortex brown, light or dark grey, some- times with a brilliant sheen in transmitted light of mi- croscope (Ha). Crocidura luna Dollman, 1910 Greater grey-brown musk shrew (PI. 12) Origin of hair sample: TM 7728: Mt. Selinda, Melset- ter Distr., Zimbabwe. Hair colour: Dorsal and ventral hairs dark grey at basal part and light brown distally, lateral hairs dark grey at basal part and dark brown distally. Hair types, shape and length: Heavy overhairs straight and without constrictions; curled overhairs (shield hairs) with thickened and grooved distal part and three con- strictions; underhairs thinner and undulating at constric- tions. Length of overhairs about 6.7 + 0.5 mm (n = 5); maximum diameter of shield region 31 um, minimum diameter about 11 um. Description of whole mounts: Hairs lack medulla at base (A), further distally fragmental simple medulla (B) and uniserial ladder (C-F), sometimes with irregular and “N” or “V”-shaped rungs (E), ending in a thin transpar- ent distal hair tip (G). Form of scale margins: Smooth at base (I, J), dentate (K-M), smooth (N), rippled (O) and smooth (P) at distal hair tip. Distance between scale margins: Scale margins near (1) at base, distant (J-M), near (N) and close (O, P) at distal hair tip. Cuticular scale patterns: At base simple coronal (I-K), further distally narrow diamond petal (L), diamond petal/transitional (M), cupped (N), regular waves (O) and coronal (P) at distal hair tip. Description of cross sections: Cross sections predomi- nantly rectangular shaped in outline with emarginations of all sides with prominent extensions (Ha, Hb, Qa), dumb-bell shaped (Qb), quadrangular shaped with con- cave sides resulting in a star-like shape (Hc) or trape- zoidal (Qc), others circular in outline (Hd); cortex and medulla clearly separated; all cross sections with me- dium size or large medulla, sometimes emarginated (Hb), reflecting the shape of the cross section; medulla black or grey, cortex brown, light or dark grey, some- times with a brilliant sheen in transmitted light of mi- croscope (Ha, b, Qa-c). Crocidura maquassiensis Roberts, musk shrew (Pl. 13) 1946 Maquassie Origin of hair sample: TM 40460: Kosi Lake, Dept. of Health Camp, Natal, South Africa. Hair colour: Hairs dark grey at basal part and light brown distally. Hair types, shape and length: Heavy overhairs straight and without constrictions; curled overhairs (shield hairs) with thickened and grooved distal part and two constric- tions; underhairs thinner and undulating at constrictions (H). Length of overhairs about 4.3 + 0.3 mm (n = 5); maximum diameter of shield region 31 um, minimum diameter about 9 um. Description of whole mounts: Hair margins at base strongly serrated with simple medulla (A, B), further dis- 116 Bonner zoologische Beitráge 54 (2005) tally uniserial ladder (C-E), with regular (D) or slightly oval rungs (E), ending in a thin transparent distal hair tip (F). Form of scale margins: Smooth at base (I, J), dentate (K-M), smooth (N), crenate (O) and smooth (P) at distal hair tip. Distance between scale margins: Scale margins near (basal part of I) at base, distant (distal part of I-M), near (N) and close (O, P) at distal hair tip. Cuticular scale patterns: At base simple coronal (I, J), further distally narrow diamond petal (K, L), diamond petal/transitional (M), cupped (N), irregular waves (O) and coronal (P) at distal hair tip. Description of cross sections: Cross sections predomi- nantly rectangular shaped in outline with emarginations of the long sides and prominent extensions of short sides (Ga) or dumb-bell shaped with rounded short sides (Gb); other cross sections with emarginations of all sides resulting in “star”-like shapes (Gc); all cross sec- tions with medium size or large medulla, sometimes emarginated, reflecting the shape of the cross section; medulla black or grey, cortex brown, light or dark grey. Crocidura mariquensis (A. Smith, 1844) Swamp musk shrew (PI. 14) Origin of hair sample: NKW 16: Punda, KNP, South Africa. Hair colour: Dorsal and lateral hairs dark grey, ventral hairs brownish-dark grey at basal part and dark brown distally. Hair types, shape and length: Heavy overhairs straight and without constrictions; curled overhairs (shield hairs) with thickened and grooved distal part and three con- strictions; underhairs thinner and undulating at constric- tions. Length of overhairs about 6.0 + 0.6 mm (n = 7); maximum diameter of shield region 31 tm, minimum diameter about 7 um. Description of whole mounts: Hairs at base with sim- ple medulla (A) and uniserial ladder (B-E), further dis- tally with individual rungs touching one another (D, E), ending in a thin transparent distal hair tip (F). Form of scale margins: Smooth at base (H, I), dentate (J, K), smooth (L), rippled (M), crenate (N) and smooth (O) at distal hair tip. Distance between scale margins: Scale. margins near (basal part of H) at base, distant (distal part of H-L), near (M) and close (N, O) at distal hair tip. Cuticular scale patterns: At base simple coronal (H, I), further distally narrow diamond petal (J, K), transi- tional (basal part of L), cupped (distal part of L), irregu- lar waves (M, N) and coronal (O) at distal hair tip. Description of cross sections: Cross sections predomi- nantly trapezoidal (Ga), others square in outline with emarginations of all sides resulting in “star”-like shapes (Gb) or circular in outline (Gc); all cross sections with medium size or large medulla, sometimes emarginated, reflecting the shape of the cross section; medulla black or grey, cortex brown, light or dark grey. Crocidura olivieri martiensseni (Lesson, 1827) Oliv- ier’s shrew (Pl. 15) Origin of hair sample: TM 1026: Mamba, Tanzania (Sorex martiensseni). Hair colour: Hairs dark grey at basal part and dark brown distally. Hair types, shape and length: Heavy overhairs straight and without constrictions; curled overhairs (shield hairs) with thickened and grooved distal part and three con- strictions; underhairs thinner and undulating at constric- tions. Length of overhairs about 9.5 + 0.6 mm (n = 3); maximum diameter of shield region 67 um, minimum diameter about 12 um. Description of whole mounts: Hairs at base with sim- ple medulla (A, B), uniserial ladder (C-F) with rungs very close to one another (C) and further distally with irregular rungs forming ‘letters’ (D), ending in a thin transparent distal hair tip (F). Form of scale margins: Smooth at base (I, basal part of J), dentate (distal part of J-L), smooth/crenate (M-P) at distal hair tip. Distance between scale margins: Scale margins near (basal part of I) at base, distant (distal part of I-L), near (M) and close (N-P) at distal hair tip. Cuticular scale patterns: At base simple coronal (I, basal part of J), narrow diamond petal (distal part of J- K), diamond petal/transitional (L), irregular waves (M), cupped (N), regular waves (O) and coronal (P) at distal hair tip. Description of cross sections: Cross sections square in outline and emarginations of short sides (Ha), oval (Hb), trapezoidal with (Ga) or without emarginations of the sides (Gb) or circular in outline (Qa); cross sections with small, medium size or large medulla, sometimes emarginated, reflecting the shape of the cross section; Michael F. SCHNEIDER & Victorino A. BURAMUGE: Atlas of Microscopic Hair 117 medulla black or grey, cortex brown, light or dark grey, sometimes with a brilliant sheen in transmitted light of microscope (Q). Crocidura olivieri occidentalis (Lesson, 1827) Olivier’s shrew (PI. 16) Origin of hair sample: TM 11413: Inyanga, Zim- babwe. Hair colour: Hairs grey at basal part and brownish-grey distally. Hair types, shape and length: Heavy overhairs straight and without constrictions; curled overhairs (shield hairs) with thickened and grooved distal part and one or two constrictions; underhairs thinner and undulating at con- strictions. Length of overhairs about 5.3 + 0.1 mm (n = 5); maximum diameter of shield region 38 tm, mini- mum diameter about 7 tm. Description of whole mounts: Hairs lack medulla at base (basal part of A), further distally simple medulla (distal part of A), uniserial ladder (B-D), sometimes with irregular rungs forming “letters” (C), distally with oblique rungs (D), ending in a thin transparent distal hair tip (E). Form of scale margins: Smooth at base (G, H), dentate (1), smooth (J), crenate (K) and smooth (L) at distal hair tip. Distance between scale margins: Scale margins near (basal part of G) at base, distant (distal part of G-I), near (J) and close (K, L) at distal hair tip. Cuticular scale patterns: At base simple coronal (G, H), narrow’ diamond petal (I), diamond petal/transitional, cupped (J), irregular waves (K) and coronal (L) at distal hair tip. Description of cross sections: Cross section rectangu- lar (Fa) or trapezoidal (Fb) in outline, with emargina- tions of long sides and small extensions or circular out- lines (Fc) of cross sections; all cross sections with small size, sometimes emarginated, reflecting the shape of the cross section; medulla black, brown or grey, cortex brown, light or dark grey. Crocidura silacea Thomas, 1895 Peters’ or lesser grey- brown musk shrew (Pl. 17) Origin of hair sample: KNP 13482: Skukuza Camp, KNP, South Africa. Hair colour: Dorsal hairs brownish-grey and dorsal hairs light grey with a brown hue. Hair types, shape and length: Heavy overhairs straight and without constrictions; curled overhairs (shield hairs) with thickened and grooved distal part and one or two constrictions; underhairs thinner and undulating at con- strictions. Length of overhairs about 4.3 + 0.4 mm (n = 7); maximum diameter of shield region 34 um, mini- mum diameter about 8 um. Description of whole mounts: Hairs lack medulla at base (basal part of A), further distally simple medulla (distal part of A), uniserial ladder (B-F) with regular rungs, ending in a thin transparent distal hair tip (F). Form of scale margins: Smooth at base (H), dentate (I- K), smooth (L, M), crenate (N) and smooth (O) at distal hair tip. Distance between scale margins: Scale margins near (basal part of H) at base, distant (distal part of H-K), near (L, M) and close (N, O) at distal hair tip. Cuticular scale patterns: At base simple coronal (H), narrow diamond petal (I, J), diamond petal/transitional (K), cupped (L, M), irregular waves (N) and coronal (O) at distal hair tip. Description of cross sections: Cross sections square (Ga) or trapezoidal (Pa) in outline with emarginations of sides, small extensions and medium size medulla, other cross sections circular in outline with large medulla (Gb); medulla black or grey, cortex brown, light or dark grey. Crocidura turba Dollman, 1910 Tumultuous shrew (Pl. 18) Origin of hair sample: TM 13797: N. R. Abercorn, L. Chiela, Zambia. Hair colour: Hairs dark grey at basal part and brownish-grey distally. Hair types, shape and length: Heavy overhairs straight and without constrictions; curled overhairs (shield hairs) with thickened and grooved distal part and two or three constrictions; underhairs thinner and undulating at con- strictions. Length of overhairs about 6.2 + 0.5 mm (n = 4); maximum diameter of shield region 36 um, minimum diameter about 6 tum. Description of whole mounts: Hairs at base with simple medulla (A, B), further distally uniserial ladder (C-E), at distal part with irregular rungs forming “letters” (D) or “mushroom”-like (E) rungs, ending in a thin transparent distal hair tip (F). 118 Bonner zoologische Beitráge 54 (2005) Form of scale margins: Smooth at base (H-J), dentate (K, E), smooth (M), crenate (N, O) and smooth (P) at distal hair tip. Distance between scale margins: Scale margins near (basal part of H) at base, distant (distal part of H-L), near (M) and close (N-P) at distal hair tip. Cuticular scale patterns: At base simple coronal (H-J), narrow diamond petal (K), diamond petal/transitional (L), cupped (M), irregular waves (N, O) and coronal (P) at distal hair tip. Description of cross sections: Cross sections predomi- nantly square (Ga), rectangular, dumb-bell shaped (Gb) or trapezoidal (Gc) in outline with emarginations of long sides and occasionally with small extensions, some with oval outline (Gd); cross sections with small or me- dium size medulla, sometimes emarginated, reflecting the shape of the cross section; medulla black or grey, cortex brown, light or dark grey, sometimes with a bril- liant sheen in transmitted light of microscope. Myosorex cafer (Sundevall, 1846) Dark-footed forest shrew (PI. 19) Origin of hair sample: TM 823: Port St. Johns, Pondo- land, Cape Province, South Africa. Hair colour: Hairs dark grey at basal part and light brown to dark brown distally. Hair types, shape and length: Heavy overhairs straight and without constrictions; curled overhairs (shield hairs) with thickened and grooved distal part and three con- strictions; underhairs thinner and undulating at constric- tions. Length of overhairs about 8.3 + 0.2 mm (n = 5); maximum diameter of shield region 40 um, minimum diameter about 12 um. Description of whole mounts: Hairs lack medulla at base (basal part of A), further distally fragmental me- dulla (distal part of A) and uniserial ladder (B-F), at dis- tal part sometimes with irregular rungs (D), ending in a thin transparent distal hair tip (FP). Form of scale margins: Smooth at base (H, I), dentate (J, K, basal part of L), smooth (distal part of L, basal part of M), crenate/rippled (distal part of M, basal part of N) and coronal (distal part of N) at distal hair tip. Distance between scale margins: At base scale mar- gins near (basal part of H), distant (distal part of H-L), near (M) and close (N) at distal hair tip. Cuticular scale patterns: At base simple coronal (H, 1), narrow diamond petal (J, K), diamond petal/transitional (basal part of L), cupped (distal part of L, basal part of M) and irregular waves (distal part of M, N) at distal hair tip. Description of cross sections: Cross sections predomi- nantly square in outline with emarginations of two sides, some with extensions forming the letter “H” (Ga, Oa), some cross sections circular (Gb, Ob) in outline, all cross sections with medium size or large medulla re- flecting the shape of the cross section; medulla black, dark or light grey, cortex black, brown or grey, some- times with a brilliant sheen in transmitted light of mi- croscope (Ga). Myosorex longicaudatus Meester & Dippenaar, 1978 Long-tailed forest shrew (PI. 20) Origin of hair sample: TM 32182: Langeberge, Boos- manskos Wilderness Area, Cape Province, South Africa. Hair colour: Hairs dark grey at basal part and brownish grey to light brown distally. Hair types, shape and length: Heavy overhairs straight and without constrictions; curled overhairs (shield hairs) with thickened and grooved distal part and five constric- tions; underhairs thinner and undulating at constrictions. Length of overhairs about 8.9 + 0.4 mm (n = 6); maxi- mum diameter of shield region 31 um, minimum diame- ter about 5 um. Description of whole mounts: Hairs lack medulla at base (basal part of A), further distally simple medulla (distal part of A) and uniserial ladder (B-D), at distal part with regular rungs (C, D), ending in a thin transpar- ent distal hair tip (E). Form of scale margins: Smooth at base (G), dentate (H, I), smooth (J, K), and rippled (L, M) at distal hair tip. Distance between scale margins: Scale margins near (basal part of G) at base, distant (distal part of G-J), near (K) and close (L, M) at distal hair tip. Cuticular scale patterns: At base simple coronal (G), narrow diamond petal (H, I), diamond petal/transitional (J), cupped (K) and irregular waves (L, M) at distal hair tip. Description of cross sections: Cross sections predomi- nantly square (Fa), occasionally trapezoidal (Fb) in out- line with slight emarginations of short sides, some with extensions forming the letter “H”, some cross sections circular in outline, medulla of all cross sections small or medium size; medulla black or grey, cortex brown or Michael F. SCHNEIDER & Victorino A. BURAMUGE: Atlas of Microscopic Hair 119 grey, sometimes with a brilliant sheen in transmitted light of microscope. Myosorex sclateri Thomas & Schwann, 1905 Sclater’s forest shrew (PI. 21) Origin of hair sample: TM 32418: Umlalazi Natl. Res. Mtunzini, Natal, South Africa. Hair colour: Hairs dark grey at basal part and brownish grey to light brown distally. Hair types, shape and length: Heavy overhairs straight and without constrictions; curled overhairs (shield hairs) with thickened and grooved distal part and five constric- tions; underhairs thinner and undulating at constrictions. Length of overhairs about 7.3 + 0.3 mm (n = 5); maxi- mum diameter of shield region 42 um, minimum diame- ter about 9 um. Description of whole mounts: Hairs lack medulla at base (basal part of A), further distally simple medulla (distal part of A), uniserial ladder with regular rungs (B- E), ending in a thin transparent distal hair tip (F). Form of scale margins: Smooth at base (H, I), dentate (J, K), smooth (L), crenate (M) and smooth (N) at distal hair tip. Distance between scale margins: Scale margins near (basal part of H) at base, distant (distal part of H-K), near (L) and close (M, N) at distal hair tip. Cuticular scale patterns: At base simple coronal (H, I), narrow diamond petal (J), diamond petal/transitional (K), cupped (L), irregular waves (M) and coronal (N) at distal hair tip. Description of cross sections: Cross sections predomi- nantly square (Ga), rectangular (Gb) and trapezoidal (Gc) in outline with slight emarginations of short sides, some with extensions forming the letter “H”, some cross sections circular (Gd, Oa) in outline, medulla of all cross sections small or medium size; medulla black or grey, cortex black, brown or grey. Myosorex tenuis Thomas & Schwann, 1905 Zuurbon forest or thin mouse shrew (PI. 22) Origin of hair sample: TM 43410: Wolkberg Wilder- ness Area, Transvaal, South Africa. Hair colour: Hairs dark grey at basal part and brownish grey to light brown distally. Hair types, shape and length: Heavy overhairs straight and without constrictions; curled overhairs (shield hairs) with thickened and grooved distal part and five constric- tions; underhairs thinner and undulating at constrictigns. Length of overhairs about 8.2 + 0.4 mm (n = 4); maxi- mum diameter of shield region 33 um, minimum diame- ter about 9 um. Description of whole mounts: Hairs lack medulla at base (basal part of A), further distally simple medulla (distal part of A), uniserial ladder with regular rungs (B, C), ending in a thin transparent distal hair tip (D). Form of scale margins: Smooth at base (F, G), dentate (H, basal part of I), smooth (distal part of I, J), rippled (K, L) and smooth (M) at distal hair tip. Distance between scale margins: Scale margins near (basal part of F) at base, distant (distal part of F, G-I), near (J) and close (K-M) at distal hair tip. Cuticular scale patterns: At base simple coronal (F), narrow diamond petal (G, H), diamond petal/transitional (1), cupped (J), irregular waves (K, L) and coronal (M) at distal hair tip. Description of cross sections: Cross sections predomi- nantly rectangular (Na), some quadrangular (Ea) in out- line with slight emarginations of two or all four sides, some cross sections circular in outline (Eb); medulla medium size or large, sometimes divided (Na) reflecting the shape of the cross section; medulla black or grey, cortex brown or grey, sometimes with a brilliant sheen in transmitted light of microscope (Eb). Myosorex varius (Smuts, 1832) Forest shrew (PI. 23) Origin of hair sample: TM 29409: Cathedral Peak For- est Reserve, Natal, South Africa. Hair colour: Hairs dark grey at basal part and light brown at distal hair tip. Hair types, shape and length: Heavy overhairs straight and without constrictions; curled overhairs (shield hairs) with thickened and grooved distal part and six constric- tions; underhairs thinner and undulating at constrictions. Length of overhairs about 7.3 + 0.9 mm (n = 6); maxi- mum diameter of shield region 38 um, minimum diame- ter about 6 um. Description of whole mounts: Hairs lack medulla at base (basal part of A), further distally simple medulla (distal part of A), uniserial ladder with regular rungs (B- D), ending in a thin distal hair tip (E). Form of scale margins: Smooth at base (G, H, basal part of I), dentate (distal part of I, J, basal part of K), 120 Bonner zoologische Beitráge 54 (2005) smooth (distal part of K, L), crenate (M) and smooth (N) at distal hair tip. Distance between scale margins: Scale margins near (basal part of G) at base, distant (distal part of G-K), near (L) and close (M, N) at distal hair tip. Cuticular scale patterns: At base simple coronal (G-I), narrow diamond petal (J), diamond petal/transitional (K), cupped (L), irregular waves (M) and coronal (N) at distal hair tip. Description of cross sections: Cross sections predomi- nantly rectangular (Fa), some quadrangular (Fb) in out- line with slight emarginations of short sides and exten- sions forming the letter “H”, some cross sections circular in outline (Fc); all cross sections with small to medium size medulla; medulla black or grey, cortex brown or grey, sometimes with a brilliant sheen in transmitted light of microscope (Fa, b). Suncus infinitesimus (Heller, 1912) Least dwarf shrew (Pl. 24) Origin of hair sample: TM 3494: Waterkloof, Pretoria, TV, South Africa. Hair colour: Hairs dark grey to brownish-grey. Hair types, shape and length: Heavy overhairs straight and without constrictions; curled overhairs (shield hairs) with thickened and grooved distal part and two constric- tions; underhairs thinner and undulating at constrictions. Length of overhairs about 2.7 + 0.2 mm (n = 7); maxi- mum diameter of shield region 29 um, minimum diame- ter about 4 um. Description of whole mounts: Hairs lack medulla at base (A); further distally simple medulla (B), uniserial ladder with regular rungs (C, D), ending in a thin trans- parent distal hair tip (E). Form of scale margins: Smooth at base (G, basal part of H), dentate (distal part of H, I), smooth (J) and cre- nate (K-M) at distal hair tip. Distance between scale margins: Scale margins near (basal part of G) at base, distant (distal part of G-I), near (J, basal part of K) and close (distal part of K-M) at dis- tal hair tip. Cuticular scale patterns: At base simple coronal (G), narrow diamond petal (H, I), diamond petal/transitional (basal part of J), cupped (distal part of J) and irregular waves (K-M) at distal hair tip. Description of cross sections: Cross sections predomi- nantly rectangular (Fa), some quadrangular (Fb) in out- line with slight emarginations of short sides and with extensions, some cross sections circular in outline (Fc); all cross sections with small to medium size medulla, sometimes divided (Fa) and reflecting the shape of the cross section; medulla black or grey, cortex brown or grey. Suncus lixus (Thomas, 1898) Greater dwarf shrew (Pl. 25) Origin of hair sample: TM 41827: Pietermaritzburg Natal, South Africa. Hair colour: Hairs light grey to dark grey. Hair types, shape and length: Heavy overhairs straight and without constrictions; curled overhairs (shield hairs) with thickened and grooved distal part and two or three constrictions; underhairs thinner and undulating at con- strictions. Length of overhairs about 3.7 + 0.3 mm (n = 7); maximum diameter of shield region 24 um, minimum diameter about 7 um. Description of whole mounts: Hairs lack medulla at base (basal part of A), further distally simple medulla (distal part of A, B), uniserial ladder (C-F), at distal part with regular (D), oval-shaped (E) or “mushroom”-shaped rungs (F), ending in a thin transparent distal hair tip (G). Form of scale margins: Smooth at base (1, J) and den- tate (K, L) and smooth (M-P) at distal part. Distance between scale margins: Scale margins near (basal part of I) at base, distant (distal part of I-M), near (N, O) and close (P) at distal end. Cuticular scale patterns: At base simple coronal (I, J), narrow diamond petal (K), diamond petal (L), transi- tional (M), cupped (N, O) and coronal (P) at distal end. Description of cross sections: Cross sections predomi- nantly rectangular (Ha), some trapezoidal (Hb) in out- line with slight emarginations of long sides and convex short sides; some cross sections circular in outline (Hc): all cross sections with medium size medulla, reflecting the shape of the cross section; medulla black, cortex lighter brown or grey. Suncus varilla (Thomas, 1895) Lesser dwarf shrew (Pl. 26) Origin of hair sample: TM 7842: Odendals Rest, OFS, SA, South Africa. Michael F. SCHNEIDER & Victorino A. BURAMUGE: Atlas of Microscopic Hair 121 Hair colour: Hairs dark grey at basal part and light brown distally. Hair types, shape and length: Heavy overhairs straight and without constrictions; curled overhairs (shield hairs) with thickened and grooved distal part and three con- strictions; underhairs thinner and undulating at constric- tions. Length of overhairs about 5.5 + 0.8 mm (n = 7); maximum diameter of shield region 27 um, minimum diameter about 7 um. Description of whole mounts: Hairs lack medulla at base (basal part of A), further distally simple medulla (distal part of A, B), uniserial ladder with regular rungs (C-E), ending in a thin transparent distal hair tip (E). Form of scale margins: Smooth at base (G, H), dentate (I, J, basal part of K), smooth (distal part of K, L) and crenate (M, N) at distal hair tip. Distance between scale margins: Scale margins near (basal part of G) at base, distant (distal part of G-K), near (L) and close (M, N) at distal hair tip. Cuticular scale patterns: At base simple coronal (G, H), narrow diamond petal (I, J), diamond petal/transitional (K), cupped (L) and irregular waves (M, N) at distal hair tip. Description of cross sections: Cross sections predomi- nantly rectangular (Fa) in outline with slight emargina- tions of long sides; some cross sections circular (Fb) or oval/oblong (Fc) in outline; all cross sections with small to medium size medulla, reflecting the shape of the cross section; medulla black, cortex lighter brown or grey. Sylvisorex megalura (Jentink, 1888) Climbing shrew (Pl. 27) Origin of hair sample: TM 34610: Mount Selinda. Chirinda Forest, Zimbabwe. Hair colour: Dorsal and lateral hairs dark brown at basal part and dark brown to brownish-grey distally; ventral hairs light grey at basal part and dark brown to brownish-grey distally. Hair types, shape and length: Heavy overhairs straight and without constrictions; curled overhairs (shield hairs) with thickened and grooved distal part and three con- strictions; underhairs thinner and undulating at constric- tions. Length of overhairs about 7.3 + 0.4 mm (n = 7); maximum diameter of shield region 32 um, minimum diameter about 6 um. Description of whole mounts: Hairs lack medulla at base (basal part of A), further distally simple medulla (distal part of A, B), uniserial ladder (C, D), distally sometimes with oblique rungs (D), ending in a {hin transparent distal hair tip (E). Form of scale margins: Smooth at base (G, H), dentate (I, basal part of J), smooth (distal part of J, K), crenate (L) and smooth (M) at distal hair tip. Distance between scale margins: Scale margins near (basal part of G) at base, distant (distal part of G-J), near (K) and close (L, M) at distal hair tip. Cuticular scale patterns: At base simple coronal (G, H), narrow diamond petal (1); diamond petal/transitional (J), cupped (K), irregular waves (L) and coronal (M) at distal hair tip. Description of cross sections: Cross sections predomi- nantly rectangular in outline with emarginations of long sides (Fa), some additionally with one plane short side and extensions on the opposite convex side (Na); some cross sections quadrangular (Fb) and circular (Fc, Nb) in outline; all cross sections with small to large medulla, sometimes divided (Fd) and reflecting the shape of the cross section; medulla black or grey, cortex black, brown or grey, sometimes with a brilliant sheen in trans- mitted light of microscope (Na, b). 4.3. Order Erinaceomorpha Family Erinaceidae (hedgehogs) The pelage of hedgehogs consists of straight, stiff hairs variable in size. Overhairs are relatively long or shorter, very thick and bristle-like (Fig. Ic); underhairs are thin- ner. Cross sections of hairs are circular, the medulla is simple or of uniserial ladder type. Scale patterns with ir- regular waves. The described hair characteristics refer to thicker overhairs. Atelerix frontalis (A. Smith, 1831) Southern African hedgehog (PI. 28) Origin of hair sample: TM 8019: Oshikango, Ovambo- land, Namibia. Hair colour: The colour of lateral hairs is dark brown alternating with light brown, hairs of head and some lat- eral hairs uniformly light brown. Hair types, shape and length: Length and width of hairs variable from very long and thick to shorter and thinner hairs. Overhairs wider at distal part. Length of overhairs about 24.8 + 5.9 mm (n = 5); maximum di- ameter of hair 160 um, minimum diameter about 29 um. 122 Bonner zoologische Beitráge 54 (2005) Description of whole mounts: Overhairs with black spot and fragmental medulla at base (A), further distally uniserial ladder (B, C) and simple medulla (D-F), distal hair tip transparent (G). Form of scale margins: At base dentate (1) and crenate (J-N) distally. Distance between scale margins: Scale margins distant at base (I), near (J-L) and close (M, N) at distal hair tip. Cuticular scale patterns: At base petal scale patterns (1); irregular waves (J-N) at distal part of hair. Description of cross sections: Predominantly circular shapes of cross sections with small medulla (Ha) or large medulla (Hc); medulla always dark brown or black, cortex of some cross sections light brown, result- ing in an intense brilliant sheen in transmitted light of microscope (Hc), cortex of other sections dark brown or black (Hb) 4.4. Order Chrysochloridea Family Chrysochloridae (golden moles) The pelage of golden moles consists of very characteris- tic soft, silky hairs with a golden sheen. The relatively long overhairs are thicker, grooved and flattened at the distal third and much thinner at their basal part (Fig. Id). Subtypes of overhairs are shorter and thinner at their distal third. Cross sections of wider part of hairs typically have oval, oblong or concavo-convex outline (Fig. 3); the medulla has wide medulla lattice. Scale pat- terns of grooved part of shield region always with regu- lar or irregular waves. The described hair characteristics refer to longer and thicker type of overhairs. Amblysomus hottentotus (A. Smith, 1829) Hottentot golden mole (PI. 29) Origin of hair sample: TM 40789: Grasskop Town, Transvaal, South Africa. Hair colour: Dorsal hairs grey at basal part and distally vary from dark brown to gold; lateral hairs are golden- brown at their distal part; ventral hairs light grey at base and light gold distally. Hair types, shape and length: Hairs long and very thick at distal third; distal hair tip forming an angle of about 11°. Subtypes distally thinner. Hair length about 13.0 + 1.3 mm (n = 5); maximum diameter of shield re- gion 153 um, minimum diameter about 18 tum. Description of whole mounts: At base fragmental me- dulla (basal part of A), uniserial ladder (distal part of A), at thicker distal third wide medulla lattice (B-E), oc- casionally with cells reaching from one margin to the other (D). Form of scale margins: At base smooth (G, H), dentate (1), smooth (J), rippled (K) and crenate (L) at distal hair tip. Distance between scale margins: Scale margins near (G, H) at base, distant (I, J); near (K) and close (L) at distal hair tip. Cuticular scale patterns: At base simple coronal (G, H), club-shaped/transitional (I, J), regular waves (K) and irregular waves (L) at distal hair tip. Description of cross sections: Cross section of shield region predominantly oblong with medium size or large medulla (Fa), some cross sections with one invaginated convex side (Fb); cross sections of thin basal part of hairs circular in outline with medium size medulla (Fc); medulla always dark brown or black and cortex light brown, resulting in a brilliant sheen in transmitted light of microscope (F). Amblysomus iris Thomas & Schwann, 1905 Zulu gol- den mole (PI. 30) Origin of hair sample: TM 26304: Diepwalle Forest Res., Krysna, Cape Province, South Africa. Hair colour: Hairs grey at basal part and greyish-brown distally. Hair types, shape and length: Hairs relatively short and very thick at distal third; distal hair tip forming an angle of about 4°. Subtypes distally thinner. Hair length about 7.9 + 0.5 mm (n = 4); maximum diameter of shield region 143 um, minimum diameter about 14 um. Description of whole mounts: At base fragmental me- dulla (distal part of A), further distally uniserial ladder, at thicker distal third wide medulla lattice (B, C), ending in a transparent thin tip (D). Form of scale margins: At base smooth scale margins (F), dentate (G, basal part of H), at wider distal third of hair smooth (distal part of H, I-M). Distance between scale margins: Scale margins close (basal part of F) at base, distant (distal part of F, G-I); near (J) and close (K-M) at distal hair tip. Cuticular scale patterns: At base simple coronal (F), narrow diamond petal (G, basal part of H), club-shaped/ transitional (H, I); regular waves (I-L) and irregular waves (M) at distal hair tip. Michael F. SCHNEIDER & Victorino A. BURAMUGE: Atlas of Microscopic Hair 123 Description of cross sections: Cross sections of shield region oblong or concavo-convex in outline with large medulla (Ea); cross section of thin basal portion of hair of circular outline with large medulla (Eb); medulla black and cortex brown or gold, resulting in an intense brilliant sheen in transmitted light of microscope (E). Calcochloris obtusirostris (Peters, 1851) Yellow golden mole (Pl. 31) Origin of hair sample: MHN: Tofo, Inhambane Prov., Mozambique; NKW 7: Machai Sandveld, KNP, South Africa. Hair colour: Hairs yellow-gold at basal part and dis- tally dark brown. Hair types, shape and length: Hairs relatively short and thick at distal third; distal hair tip forming an angle of about 15”. Subtypes distally thinner. Hair length about 6.3 + 0.3 mm (n = 5); maximum diameter of shield region 99 tum, minimum diameter about 19 um. Description of whole mounts: At base fragmental me- dulla (basal part of A), further distally uniserial ladder (distal part of A), at thicker distal third wide medulla lattice (B, C), ending in a blunt tip (C). Form of scale margins: At base with smooth scale margins (E); dentate (F, G); at wider distal third of hair smooth (H) and rippled/crenate (I-K) at distal hair tip. Distance between scale margins: Scale margins close at base, further distally distant (E-G); at wider distal part of hair near (H, I) and close (J, K) at distal hair tip. Cuticular scale patterns: At base coronal (basal part of E), further distally narrow diamond petal (F), club- shaped/transitional (G, H) and irregular waves (I-K) at thicker distal third of hair. Description of cross sections: Cross sections of shield region oblong in outline (Da), slightly dumb-bell shaped (Db) or concavo-convex shapes (Dc) with small me- dulla; cross section of thin basal part of hair circular in outline with small medulla (Dd); medulla often light brown or grey and cortex dark (Da). Carpitalpa arendsi Lundholm, 1955 Arend’s golden mole (Pl. 32) Origin of hair sample: TM 12778: Vumba Mt., Umtali Distr., Zimbabwe (Chlorotalpa arendsi). Hair colour: Dorsal and lateral hairs grey at basal part and dark brown further distally. Ventral hairs grey at basal part and distally light brown. Hair types, shape and length: Hairs relatively long and thick at distal third; distal hair tip forming an angle of about 3°. One subtype with thinner distal part. Hair length about 13.0 + 0.8 mm (n = 5); maximum diameter of shield region 101 um, minimum diameter about 5 um. Description of whole mounts: At base simple medulla, sometimes fragmental (basal part of A), further distally uniserial ladder (distal part of A); at thicker distal third wide medulla lattice (B-D), ending in a thin tip (E). Form of scale margins: At base with smooth scale margins (G); dentate (H, basal part of I); at wider distal third of hair smooth (distal part of I, J) and crenate (K- M) at distal hair tip. Distance between scale margins: Scale margins close at base, further distally distant (G-I); at wider distal part of hair near (J) and close (K-M) at distal hair tip. Cuticular scale patterns: At base simple coronal (G), further distally narrow diamond petal/composed double chevron (H), club-shaped/transitional (1), regular waves (J) and irregular waves (K-M) at distal hair tip. Description of cross sections: Cross sections of shield region oblong or oval in outline with large medulla (Fa) or concavo-convex outline with large medulla (Fb); cross section of thin basal part of hair circular in outline with large medulla (Fc); medulla grey or black, cortex of some cross sections light brown resulting in a weak sheen in transmitted light of microscope (Fa), cortex of other sections black. Chlorotalpa duthieae (Broom, 1907) Duthie’s golden mole (Pl. 33) Origin of hair sample: TM 19119: Krysna, Cape Prov- ince, South Africa. Hair colour: Hairs grey at basal part and brown dis- tally. Hair types, shape and length: Hairs relatively long and thick at distal third; distal hair tip forming an angle of about 10°. Subtypes with thinner distal part. Hair length about 12.1 + 1.1 mm (n = 5); maximum diameter of shield region 101 um, minimum diameter about 7 um. Description of whole mounts: At base fragmental me- dulla (A), further distally uniserial ladder (B); at thicker distal third wide medulla lattice (C-E). Form of scale margins: Scale margins at base smooth (G), further distally dentate (H); at wider distal third of 124 Bonner zoologische Beitráge 54 (2005) hair smooth (1), rippled (J) and crenate (K, L) at distal hair tip. Distance between scale margins: Scale margins near (G) at base, further distally distant (H); at wider distal part of hair near (1, J) and close (K, L) at distal hair tip. Cuticular scale patterns: At base coronal (G), further distally narrow diamond petal (H), regular waves (I) and irregular waves (J-L) at thicker distal third of hair. Description of cross sections: Cross sections of shield region oblong with large medulla (Fa) or concavo- convex outline with large medulla (Fb); cross section of thin basal part of hair circular in outline with large me- dulla; medulla dark grey or black, cortex light brown or grey resulting in a weak sheen in transmitted light of microscope (F). Chlorotalpa sclateri (Broom, 1907) Sclater’s golden mole (Pl. 34) Origin of hair sample: TM 39445: Mount View, Karoo N. P., Cape Province, South Africa. Hair colour: Dorsal hairs grey at basal part and brownish-gold distally. Lateral and ventral hairs grey at basal part and light gold at distal part. Hair types, shape and length: Hairs relatively short and thick at distal third; distal hair tip forming an angle of about 8°. Subtypes with thinner distal part. Hair length about 7.0 + 0.8 mm (n = 5); maximum diameter of shield region 86 um, minimum diameter about 10 um. Description of whole mounts: At base fragmental me- dulla (basal part of A), further distally uniserial ladder (distal part of A); at thicker distal third wide medulla lattice (B-D), ending in a thin transparent tip (D). Form of scale margins: At base smooth (F-H) scale margins, at wider distal third of hair rippled (1) and cre- nate (J, K) at distal hair tip. Distance between scale margins: Scale margins near at base (F, G), further distally near (H, I) and close (J, K) at distal hair tip. Cuticular scale patterns: At base simple coronal (F, basal part of G); at thicker distal third of hair regular waves (H) and irregular waves (I-K). Description of cross sections: Cross sections of shield region oblong with large medulla (Ea) or concavo- convex outline with large medulla (Eb); cross section of thin basal part of hair circular in outline with large me- dulla; medulla with brown, grey or black, cortex of some cross sections light brown resulting in an intense brilliant sheen in transmitted light of microscope (E). Chrysochloris asiatica (Linnaeus, 1758) Cape golden mole (Pl. 35) Origin of hair sample: TM 5808: Goudini, Worcester, Cape Province, South Africa. Hair colour: Dorsal hairs grey at basal part and light grey distally; ventral hairs entirely brownish-gold. Hair types, shape and length: Hairs relatively long and thick at distal third; distal hair tip forming an angle of about 4°. Subtypes shorter and with thinner distal part. Hair length about 11.2 + 0.5 mm (n = 5); maxi- mum diameter of shield region 92 um, minimum diame- ter about 9 um. Description of whole mounts: At base uniserial ladder, sometimes fragmental (A); at thicker distal third wide medulla lattice (B, C), ending in a thin transparent tip (C). Form of scale margins: Scale margins smooth (F-H) at base, at thicker distal third crenate (I-K). Distance between scale margins: Scale margins near (F-H) at base and close (I-K) at distal hair tip. Cuticular scale patterns: At base simple coronal (F); at thicker distal third of hair regular waves (G-I) and ir- regular waves (J, K) at distal hair tip. Description of cross sections: Cross sections of shield region oblong (Ea) or oval (Eb) with large medulla; cross sections of ventral hairs concavo-convex in out- line with medium size medulla (Da); cross section of thin basal part of hair circular in outline with medium size medulla (Db, Ec); medulla black, cortex light brown resulting in an intense brilliant sheen in transmit- ted light of microscope (D, E). Chrysospalax trevelyani (Gúnther, 1875) Giant golden mole (Pl. 36) Origin of hair sample: TM 746: Port St. Johns, Cape Prov., South Africa. Hair Colour: Hairs light yellow at basal part and dark brow distally. Hair types, shape and length: Hairs very long and very thick at distal third; distal hair tip forming an angle of about 9°. Subtypes with thinner distal part. Hair length about 20.6 + 0.4 mm (n = 5); maximum diameter of shield region 190 um, minimum diameter about 25 um. Michael F. SCHNEIDER & Victorino A. BURAMUGE: Atlas of Microscopic Hair | Description of whole mounts: At base fragmental me- dulla (A), further distally uniserial ladder (B); at thicker distal third wide medulla lattice (C-E). Form of scale margins: At base with smooth (G) scale margins, dentate (H, I), smooth (J, K) and crenate (L, M) at distal hair tip. Distance between scale margins: Scale margins close (G) at base, further distally distant (H-J), near (K) and close (L, M) at distal hair tip. Cuticular scale patterns: Simple coronal (G) at base, narrow diamond petal (H), club-shaped/transitional (I, J), regular waves (K, L) and irregular waves (M) at distal hair tip. Description of cross sections: Cross sections of shield region predominantly oblong with large medulla (F); cross section of thin basal part of hair circular in outline with medium size medulla; medulla black, cortex light brown resulting in an intense brilliant sheen in transmit- ted light of microscope (F). Chrysospalax villosus (A. Smith, 1833) Rough-haired golden mole (Pl. 37) Origin of hair sample: TM 22416: Pietermaritzburg Natal, South Africa. Hair colour: Hairs grey at basal part and dark brow dis- tally. Hair types, shape and length: Hairs very long and thick at distal third; distal hair tip forming an angle of about 2°. Subtypes with thinner distal part. Hair length about 20.8 + 1.0 mm (n = 5); maximum diameter of shield region 105 um, minimum diameter about 11 um. Description of whole mounts: At base fragmental simple medulla (A), at distal third of hair wide medulla lattice (B, C); ending in a thin, transparent distal hair tip (D). Form of scale margins: At base smooth (G) scale mar- gins, dentate (H); at wider distal third of hair smooth (1), smooth/crenate (J), rippled (K) and crenate (L, M) at distal hair tip. Distance between scale margins: Scale margins near at base (G), further distally distant (H, I), near (J) and close (K-M) at distal part of hair. Cuticular scale patterns: At base simple coronal (G), narrow diamond petal (H), club-shaped/transitional (1), regular waves (J) and irregular waves (K-M) at distal hair tip. m N Description of cross sections: Cross sections of shield region predominantly oblong (Fa) and oval (Ea, fb) with large medulla; cross section of thin basal part of hair circular in outline with large medulla (Eb); medulla dark grey or black, cortex light brown, resulting in an intense brilliant sheen in transmitted light of microscope (E, F). Cryptochloris wintoni (Broom, 1907) de Winton’s golden mole (PI. 38) Origin of hair sample: TM 8235: Port Nolloth, Nama- qualand, Cape Prov, South Africa. Hair colour: Hairs dark grey at basal part and of gold colour distally. Hair types, shape and length: Hairs relatively short and thick at distal third; distal hair tip forming an angle of about 12°. Subtypes with thinner distal part. Hair length about 6.9 + 0.6 mm (n = 5); maximum diameter of shield region 113 um, minimum diameter about 9 tm. Description of whole mounts: At base fragmental uniserial ladder (A); at thicker distal third wide medulla lattice with large aerial spaces (B-D). Form of scale margins: At base smooth (F) scale mar- gins, further distally dentate (G); at wider distal third of hair smooth (H) and crenate (I-K) at distal hair tip. Distance between scale margins: Scale margins close at base (F), further distally distant (G), near (H, I) and close (J, K) at distal hair tip. Cuticular scale patterns: At base coronal (F), further distally narrow diamond petal (G); at thicker distal third of hair regular waves (H) and irregular waves (I-K) at distal hair tip. Description of cross sections: Cross sections of shield region predominantly oblong with small medulla (Ea) and some cross sections with one invaginated convex side (Eb); cross section of thin basal part of hair circular in outline with medium size medulla (Ec); medulla light or dark grey, cortex light brown, resulting in an intense brilliant sheen in transmitted light of microscope (E). Eremitalpa granti (Broom, 1907) Grant’s golden mole (PL 39) Origin of hair sample: TM 8237: North Nolloth Distr., Namaqualand, Cape Province, South Africa; TM 15154: Sossusvlei, Namib Desert, Namibia. 126 Bonner zoologische Beitráge 54 (2005) Hair colour: Dorsal hairs grey and dark brown at small portion of distal tip, lateral and ventral hairs grey at basal part and distally of light gold colour. Hair types, shape and length: Hairs long and moder- ately thick at distal third; distal hair tip forming an angle of about 6°. Subtypes with thinner distal part. Hair length about 15.5 + 0.6 mm (n = 5); maximum diameter of shield region 78 um, minimum diameter about 9 um. Description of whole mounts: At base fragmental uniserial ladder (A), at thicker distal third wide medulla lattice (B-E) with larger clear spaces (D). Form of scale margins: Scale margins smooth (G, H) at base, and crenate (I-K) at distal third of hair. Distance between scale margins: At base scale mar- gins near (G, H) and close (I-K) at distal third of hair. Cuticular scale patterns: At base simple coronal (basal part of G), regular waves (distal part of G, H); at thicker distal third of hair irregular waves (H-K). Description of cross sections: Cross sections of shield region predominantly oblong (Fa) or dumb-bell shaped with large medulla (Fb), occasionally concavo-convex in outline; cross section of thin basal part of hair circular in outline with medium size medulla (Fc); medulla light or dark grey, cortex light brown, resulting in a weak sheen in transmitted light of microscope (F). Neamblysomus gunningi (Broom, 1908) Gunning’s golden mole (PI. 40) Origin of Hair Sample: TM 40780: Magoebas Kloof, de Hoek Nat. Res. Transvaal, South Africa (Amblyso- mus gunning). Hair colour: Dorsal hairs light grey at basal part and distally brown, lateral and ventral hairs light grey at basal part and distally of gold colour. Hair types, shape and length: Hairs moderately long and thick at distal third; distal hair tip forming an angle of about 6°. Subtypes with thinner distal part. Hair length about 8.1 + 1.1 mm (n = 4); maximum diameter of shield region 102 um, minimum diameter about 9 um. Description of whole mounts: At base uniserial ladder (A); at thicker distal third wide medulla lattice (B, C), sometimes with clear spaces (B), ending in a thin tip (C). Form of scale margins: At base smooth (E-G) scale margins and crenate (H, I) at distal hair tip. Distance between scale margins: At base scale mar- gins near (E-G) and close (H, I) at distal hair tip. Cuticular scale patterns: At base coronal (basal part of E), club-shaped/transitional (F); at thicker distal third of hair regular waves (H) and irregular waves (H, 1) at dis- tal hair tip. Description of cross sections: Cross sections of shield region predominantly dumb-bell shaped with large me- dulla (Da); cross section of thin basal part of hair circu- lar in outline with medium size (Db) or small medulla (Dc); medulla black, light or dark grey, cortex dark or light brown, resulting in a brilliant sheen in transmitted light of microscope (D). Neamblysomus julianae Meester, 1972 Juliana’s golden mole (PI. 41) Origin of hair sample: NKW 4: Manchulane, KNP, South Africa (=4mblysomus julianae). Hair colour: Hairs grey at basal part and distally light to dark brown. Hair types, shape and length: Hairs relatively long and thick at distal third; distal hair tip forming an angle of about 7°. Subtypes shorter and with thinner distal part. Hair length about 13.8 + 0.7 mm (n = 5); maxi- mum diameter of shield region 86 um, minimum diame- ter about 14 um. Description of whole mounts: At base uniserial ladder (A); at thicker distal third wide medulla lattice (B-D) with some aerial spaces (C), ending in a thin tip (D). Form of scale margins: At base scale margins smooth (F-J) and crenate/rippled (K-N) at distal hair tip. Distance between scale margins: At base scale mar- ems near (F, G), further distally distant (H-J), near (K, L) and close (M, N). Cuticular scale patterns: At base simple coronal (F, G), further distally club-shaped/transitional (H), regular waves (I, J) and irregular waves (K-N) at distal hair tip. Description of cross sections: Cross sections of shield region oblong (Ea) and dumb-bell shaped (Eb) with large medulla; cross section of thin basal part of hair circular in outline with medium size medulla (Ec); me- dulla black, light or dark grey, cortex light brown, re- sulting in an intense brilliant sheen in transmitted light of microscope (E). Michael F. SCHNEIDER & Victorino A. BURAMUGE: Atlas of Microscopic Hair 127 KEY TO SOUTHERN AFRICAN ELEPHANT-SHREWS, SHREWS, HEDGEHOGS AND GOLDEN MOLES BASED ON HAIR CHARACTERISTICS Pelage composed of distally enlarged overhairs and slightly curved underhairs with or without constric- tions (Fig. la); hairs occasionally with intumescence (e.g., Pl. IF & G, 4F, 6F, 7B & F); length of dorsal hairs varies between 11 mm and 18 mm; medulla with uniserial ladder, medulla of some species dis- tally with wide lattice; scale patterns of distal part of hair with regular or irregular waves, hair tip often of coronal type; cross section almost always circular NE elephant-shrews Pelage composed of straight, long heavy overhairs without constrictions, a second type of distally enlarged, grooved and slightly curved overhairs (shield hairs) and undulating underhairs (Fig. 1b); the latter two types with one or several constrictions along the hair (Pl. 13 H); hairs very short varying between 3 mm and 10 mm; except from immediate base and tip, medulla always with uniserial ladder; scale patterns of grooved part of shield region al- ways cupped, hair tip with coronal or regular waves; cross section of enlarged part of shield hair typically rectangular, or “H”-like with two or more emargina- te REA A shrews Pelage composed of straight, stiff, distally enlarged overhairs lacking constrictions and dilatations (Fig. lc) and thinner underhairs; length of hairs up to 25 mm; medulla simple or with uniserial ladder; scale patterns with irregular waves; cross section circular idolo Secs duuwatceneeas tie hedgehogs Pelage composed of long overhairs (shield hairs) with distal third expanded, flattened and club-like (Fig. 1d) and shorter sub-types with thinner distal part; length of dorsal hairs between 6 mm and 21 mm; medulla at thickened part of hair always with wide medulla lattice; scale patterns of grooved part of shield region always with regular or irregular waves; cross section of enlarged part of hair oblong, oval or CONCAVO-CONVEX .........::cseeceeeeee golden moles Elephant-shrews (or sengis) la Maximum diameter of hairs between 40 um and 1b Maximum diameter of hairs smaller than 40 um .. 5 2a Cross sections with golden sheen in transmitted Ins hitg omen CROSCOPS! ne eeesnctes dtc eteed 3 2b 3a 3b 4a 4b 5a 5b Sc Cross sections not with golden sheen in transmitted lichtof MICTOSCO PC nes y 4 Hair length about 16 mm; wide medulla lattice at distal part of overhair; maximum diameter of hairs approx. 58 um; circular cross sections with bright sheen. Distribution: N South Africa and NE Na- mibia; Angola; S Zaire; Zimbabwe, Mozambique to Kenya and Uganda...Elephantulus brachyrhynchus Hair length about 18 mm; medulla with uniserial ladder; maximum diameter of hairs approx. 59 um; circular cross sections with bright sheen. Distribu- tion: Mozambique; Tanzania (including Mafia and Zanzibar); SE Kenya; S Uganda; Zambia, Malawi; SE Zimbabwe; Zaire; Namibia; NE Angola; N Natal and E Transvaal, South Aira .2..).ccccseedecccessseodoocaesss lisas Petrodromus tetradactylus Medulla with uniserial ladder; hair length about 15 mm; maximum diameter of hairs approx. 53 um; cross sections circular in outline. Distribution: SW Angola; Namibia; Botswana; NW Transvaal and N Cape Prov., South Africa .......... Elephantulus intufi Hair length about 12 mm; medulla with wide me- dulla lattice; maximum diameter of hairs approx. 46 um; cross sections circular in outline. Distribution: Zimbabwe, Namibia, E Botswana, E and N South Africa, Mozambique .............. Elephantulus myurus Hair length about 18 mm; medulla with uniserial ladder; maximum diameter of hairs about 34 um; circular cross sections with bright sheen. Distribu- tion: Namibia; Cape Prov., South Africa................... A anc ORTE Elephantulus rupestris Hair length about 14 mm; medulla with uniserial ladder; maximum diameter of hairs approx. 30 um; circular cross sections without sheen. Distribution: SW and C Cape Prov.; South Afriea. unten. DE Elephantulus edwardi Hair length about 18 mm; medulla with uniserial ladder; maximum diameter of hairs approx. 26 um; circular cross sections with bright sheen. Distribu- tion: W and NW Cape Prov., South Africa to SW Namibia. runs Macroscelides proboscideus Shrews la 1b 2a Length of overhairs longer than 7 mm ................... 2 Length of overhairs shorter than 7 mm .................. 5 Cross sections of widest part of shield hair in the formof an Fin outline ne laa eae ites 3 128 2b 3a 3b 3c 3d 3e 4a 4b 6b 6c Bonner zoologische Beitráge 54 (2005) Cross sections of widest part of shield hair not in the form of an H in outline Hair length about 8.3 mm; maximum diameter of hairs about 40 um. Distribution: South Africa, east- ern escarpment and north to the Transvaal; extreme W Mozambique and E Zimbabwe . Myosorex cafer Hair length about 8.9 mm; maximum diameter of hairs about 31 um. Distribution: SE Cape Province, South Africa Myosorex longicaudatus Hair length about 7.3 mm; maximum diameter of hairs about 42 um; distribution: Natal, South Africa; Swaziland; Mozambique; Zimbabwe Myosorex sclateri Hair length about 8.2 mm; maximum diameter of hairs about 33 um. Distribution: Transvaal, South Africa; possibly W Mozambique .. Myosorex tenuis Hair length about 7.3 mm; maximum diameter of hairs about 38 um. Distribution: South Africa, from NW Cape Province to E Transvaal; Lesotho and Or- ange Free State Myosorex varius Hair length about 9.5 mm; maximum diameter of hairs about 67 um; medulla typically with ‘letters’ (Pl. 15D). Distribution: Egypt; Senegal to Ethiopia and southwards to N South Africa Crocidura olivieri martiensseni Hair length about 7.3 mm; maximum diameter of hairs about 32 um; cross sections quadrangular or rectangular in outline with plane and convex short sides (Pl. 27 Na). Distribution: Upper Guinea to Ethiopia and south to Namibia, Angola, Mozam- bique and Zimbabwe Sylvisorex megalura Hair length shorter than 4 mm Hair length between 4 mm and 7 mm Hair length about 2.7 mm; maximum diameter of hairs about 26 tm; cross sections rectangular in out- line with a deep invagination of one side resulting in prominent extensions of the margin (Pl. 10Fa). Dis- tribution: Sudan and Guinea, from Senegal to Ethio- pia and south to South Africa Crocidura fuscomurina Hair length about 2.7 mm; maximum diameter of hairs about 29 um. Distribution: South Africa to Kenya; Central African Republic; Cameroon Suncus infinitesimus Hair length about 3.7 mm; maximum diameter of hairs about 24 um. Distribution: Kenya, Tanzania, Ta 7b 8a 8b Sc 8d 8e 9a 9b Ic 9d Malawi, Zaire, Zambia, Angola, Botswana, Na- mibia, Zimbabwe and Transvaal, South Africa ........ Suncus lixus Cross sections dumb-bell shaped in outline; rungs of uniserial ladder irregular and often like “letters” ... 8 Cross sections quadrangular or rectangular in out- line; rungs of uniserial ladder more regular Hair length about 5.5 mm; maximum diameter of hairs about 33 um. Distribution: Angola, Zaire, Uganda, Kenya, Somalia, Tanzania, Malawi, Zim- babwe, Zambia, Mozambique, Botswana, Namibia, South Africa Crocidura hirta Hair length about 4.3 mm; maximum diameter of hairs about 31 um. Distribution: Transvaal, South Africa; Swaziland, Nyamaziwa Falls, Matopo Hills, Zimbabwe Crocidura maquassiensis Hair length about 6.7 mm; maximum diameter of hairs about 31 tm. Distribution: Mozambique, Zambia, Zimbabwe, E Angola, Zaire, Malawi, Tan- zania, Kenya, Uganda, Rwanda Crocidura luna Hair length about 6.2 mm; maximum diameter of hairs about 36 um. Distribution: Angola, Zambia, Zaire, Malawi, Tanzania, Kenya, Uganda, Camer- Crocidura turba Hair length about 5.5 mm; maximum diameter of hairs about 35 um. Distribution: southern and east- ern South Africa, southern Mozambique Crocidura flavescens Hair length about 6.2 mm; maximum diameter of hairs about 29 um. Distribution: South Africa, Na- mibia, Angola, Botswana, Mozambique, records further north uncertain Crocidura cyanea Hair length about 5.3 mm; maximum diameter of hairs about 38 um; medulla typically with “letters” (Pl. 16C). Distribution: Egypt; Senegal to Ethiopia, and southwards to N South Africa Hair length about 6.0 mm; maximum diameter of hairs about 31 um. Distribution: South Africa to Mozambique, W Zimbabwe and Zambia; NW Bot- swana and NE Namibia to SC Angola; perhaps SE Zaire Crocidura mariquensis Hair length about 4.3 mm; maximum diameter of hairs about 34 um. Distribution: most of South Af- rica, parts of Botswana, Mozambique and Zim- babwe; possibly wider distribution Crocidura silacea 9e Michael F. SCHNEIDER & Victorino A. BURAMUGE: Atlas of Microscopic Hair Hair length about 5.5 mm; maximum diameter of hairs about 27 um. Distribution: Cape, South Africa to Zimbabwe, Mozambique, Zambia, Tanzania, E Zaire, Malawi; an isolated record from Nigeria Suncus varilla Hedgehogs 1 Hair length about 25 mm; maximum diameter of hairs about 160 um; overhairs with black spot at base (Pl. 28A). Distribution: Cape Province, South Africa to E Botswana and W Zimbabwe, Namibia to SWeAngola, Zambia ................ Atelerix frontalis Golden moles la 1b 2a 2b 3a 3b da 4b 5a 5b 6a Hair length about 21 mm Hair length much less than 21 mm Hair length about 21 mm; maximum diameter of hairs about 190 um; distal hair tip forming an angle of about 9°. Distribution: Cape Prov., South Africa Chrysospalax trevelyani Hair length about 21 mm; maximum diameter of hairs about 105 um; distal hair tip forming an angle of about 2°. Distribution: Transvaal and Natal, SouthrAftical.... Anh: Chrysospalax villosus Maximum diameter of hairs between 125 um and 160 um Maximum diameter of hairs smaller than 125 um 5 Distal hair tip forming an angle of about 11°; haır length about 13 mm; maximum diameter of hairs about 153 um. Distribution: Natal, Transvaal to S Cape Prov., also NE Orange Free State, South Af- rica; Lesotho; Swaziland; S Mozambique Amblysomus hottentotus Distal hair tip forming an angle of about 4°; hair length about 8 mm; maximum diameter of hairs about 143 um. Distribution: S Cape Prov. to Tran- skei, Natal, including Zululand, and SE Transvaal, South Africa; S Mozambique ....... Amblysomus iris Distal hair tip forming an angle of about 15°; hair length about 6 mm; maximum diameter of hairs about 99 um. Distribution: Zululand and E Trans- vaal, South Africa; S Zimbabwe; S Mozambique .... Calcochloris obtusirostris Distal hair tip forming an angle < 15° Medulla lattice with large aerial spaces; hair length about 7 mm; maximum diameter of hairs about 113 um; distal hair tip forming an angle of about 12°. 6b Ta 7b 8a 8b Sc 8d 8e sf 129 Distribution: Little Namaqualand; Cape Prov., South Africa Cryptochloris wintoni Medulla lattice not with large aerial spaces Cross sections of extended part of ventral hairs pre- dominantly showing concavo-convex outline (Pl. 35Da); hair length about 11 mm; maximum diame- ter of hairs about 92 pum; distal hair tip forming an angle of about 4°. Distribution: W Cape Prov. and Robben Isl., South Africa; perhaps Damaraland, Namibia Chrysochloris asiatica Cross sections of extended part of ventral hairs pre- dominantly showing other than concavo-convex outline The following species can be distinguished ac- cording to their distribution: Hair length about 8 mm; maximum diameter of hairs about 102 um; distal hair tip forming an angle of about 6°. Distribution: E Transvaal, South Africa scuuese state, Soraaneeceeteedsmencee seeds Neamblysomus gunningi Hair length about 14 mm; maximum diameter of hairs about 86 um; distal hair tip forming an angle of about 7°. Distribution: Pretoria, Nylstroom/Nylsvley and Kruger Nat. Park (Transvaal), South Africa Neamblysomus julianae Hair length about 13 mm; maximum diameter of hairs about 101 um; distal hair tip forming an angle of about 3°. Distribution: E Zimbabwe and adjacent Mozambique Carpitalpa arendsi Hair length about 12 mm; maximum diameter of hairs about 101 um; distal hair tip forming an angle of about 10°. Distribution: S Cape Prov., South Af- Chlorotalpa duthieae Hair length about 7 mm; maximum diameter of hairs about 86 um; distal hair tip forming an angle of about 8°. Distribution: Cape Prov., E Orange Free State, S Transvaal, South Africa; Lesotho ................ Chlorotalpa sclateri Hair length about 16 mm; maximum diameter of hairs about 78 um; distal hair tip forming an angle of about 6°. Distribution: Cape Prov., South Africa to Namib Desert, Namibia; Botswana ....................... Eremitalpa granti Acknowledgements. The authors wish to thank the Maputo Natural History Museum, Scientific Services of Kruger National Park and Transvaal Museum, Pretoria for providing hair samples. Further thanks to Dr. Chris Tide- mann, Australian National University, Canberra for provid- 130 Bonner zoologische Beitráge 54 (2005) ing materials; Bembergcell SPA, Milano, Italy for provid- ing cellulose acetate yarn; Mr. Ralf Loeper, University Mainz, Germany, for his support during the initial phase of the study; Mr. Ottmar Fischer, University Freiburg, Ger- many for the electron microscopy of hair samples; Dr. Esteban Sarmiento, American Natural History Museum, New York and Dr. Rick Bein, University Indiana Univer- sity Purdue University, Indianapolis, for their critical comments on the manuscript and Centrum ftir Internation- ale Migration und Entwicklung (CIM) for funding. REFERENCES ADORJAN, A. S. & KOLENOSKY, G. B. 1969. A manual for the identification of hairs of selected Ontario mam- mals. Research Report (Wildlife) No. 90. Department of Lands and Forests, Ontario. AMERASINGHE, F. P. 1986. The structure and identification of the hairs of mammals of Sri Lanka. Ceylon Journal of Science (Biological Science) 16 (1983): 76-125. BOITANI, L., CORSI, L., DE BIASE, A., D’INZILLO CAR- RANZA, [., RAVAGLI, M., REGGIANI, G., SINIBALDI, I., & TRAPANESE, P. (eds.) 1999. A databank for the conser- vation and management of the African mammals. European Commission & Instituto di Ecologia Appli- cata, Rome. DE BOOM, H. P. A. & DREYER, J. H. 1953. The possibility of identifying hair from S.A. game for forensic pur- poses. South African Journal of Science 49: 233-234. BRONNER, G. N. 1995. Systematic revision of the golden mole genera Amblysomus, Chlorotalpa & Calcochloris (Insectivora: Chrysochloromorpha; Chrysochloridae). Ph.D. Thesis, University of Natal, Durban. BRUNNER, H. & COMAN, B. J. 1974): The Identification of Mammalian Hair. Inkata Press, Melbourne. BRUNNER, H. & WALLIS, R. 1986. Roles of Predator Scat Analysis in Australian Mammal Research. Victorian Naturalist 103: 79-87, DANNELID, E. 1986. Observation of an H-profile in lead hairs of shrews (Mammalia, Soricidae. Zeitschrift für Säugetierkunde 51:188-190. DAY, M. G. 1966. Identification of hair and feather remains in the gut and faeces of stoats and weasels. Journal of Zoology, London 148: 201-210. DEBROT, S., MERMOD, C., FIVAZ, G. & WEBER, J.-M. 1982. Atlas des poils de mammiferes d’Europe. Institut de Zoologie, Université de Neuchatel, Switzerland. DREYER, J. H. 1966. A study of hair morphology in the family Bovidae. Onderstepoort Journal of Veterinary Research 33: 379-472. DUCOMMUN, M.-A., JEANMARIE-BESANCON, F. & VOGEL, P. 1994. Shield morphology of curly overhair in 22 genera of Soricidae (Insectivora, Mammalia). Revue Suisse de Zoologie 101: 623-643. DZIURDZIK, B. 1973. Key to the Identification of Hairs of Mammals from Poland. Acta zoologica cracoviensia 18: 73-92. GILBERT, F. F. & NANCEKIVELL, E. G. 1982. Food habits of mink (Mustela vison) and otter (Lutra canadensis) in northeastern Alberta. Canadian Journal of Zoology 60: 1282-1288. HUTTERER, R. 1993. Order Insectivora. Pp. 69-130 in: WILSON, D. E. & REEDER, D. M. (eds.) Mammal Spe- cies of the World. Smithsonian Institution Press, Washington DC. KEOGH, H. J. 1975. The study of hair characteristics of forty-two species of South African Muridae and the taxonomic application of these as a definite criteria. MSc Thesis, University of Cape Town, Rondebosch. KEOGH, H. J. 1979. Applications of Hair Studies in Epide- miology. South African Journal of Science 75: 151. KEOGH, H. J. 1983. A photographic references system of the microstructure of the hair of southern African bo- vids. South African Journal of Wildlife Research 13: 89-131. KEOGH, H. J. 1985. A photographic references system based on the cuticular scale patterns and groove of the hair of 44 species of southern African Cricetidae and Muridae. South African Journal of Wildlife Research 15: 109-159. LYNE, A. G. & MCMAHON, T. S. 1951. Observations on the surface structure of the hairs of Tasmanian Monot- remes and Marsupials. Papers and Proceedings of the Royal Society of Tasmania 1950: 71-84. MARZ, R. 1987. Gewóll und Rupfungskunde. Akademie- Verlag, Berlin. MATHIAK, H. A. 1938. A key to the hairs of mammals of Southern Michigan. Journal of Wildlife Management 2: 251-268. MAYER, W. V. C. 1952. The hair of Californian mammals with keys to the dorsal guard hairs of Californian mammals. American Midland Naturalist 48: 480-512. PERRIN, M. R. & CAMPBELL, B. S. 1980. Key to the mam- mals of the Andries Vosloo Kudu Reserve (eastern Cape), based on their hair morphology, for use in predator scat analysis. South African Journal of Wild- life Research 10: 3-14. SCHLITTER, D. A. 1993. Order Macroscelidea. Pp. 829-830 in: WILSON, D. E. & REEDER, D. M. (eds.) Mammal Species of the World. Smithsonian Institution Press, Washington DC. TEERINK, B. J. 1991. Hair of West-European mammals. At- las and identification key. Cambridge University Press, New York, Port Thester, Melbourne, Sydney. WILDMAN, A. B. 1954. The microscopy of animal textile fibres. Wool Industry Research Association, Leeds. WILLIAMS, C. S. 1938. Aids to the identification of mole and shrew hairs with general comments on hair struc- ture and hair determination. Journal of Wildlife Man- agement 2: 239-250. Authors’ address: Michael F. SCHNEIDER* (*corres- ponding author), Pfeiffermuehle 3, 87497 Wertrach, Germany; E-mail: mfschneider@vr-web.de; Victorino A. BURAMUGE, Forestry Department, University Eduardo Mondlane, P. O. Box 257, Maputo, Mozambique. Received: 26.10.2004 Revided: 02.04.2005 Accepted: 15.05.2006 Corresponding editor: G. Peters 131 Michael F. SCHNEIDER & Victorino A. BURAMUGE: Atlas of Microscopic Hair xx ae ae ANG pet ALES Lidia Ad E — Liu ap AAA \| a rum nn Fran 5 er EEE A en Be EB ek MP a a Er ii ir ad shed wl GA ee o, wil ae cil A ibas pi sal 2 = a alae Sap 2 He a ge NA Pe “e ern u Plate 1. Whole mounts, cross sections and cuticular scale casts of Elephantulus brachyrhynchus (bars equivalent to 50 um). Bonner zoologische Beitráge 54 (2005) PD DIDIDIDIDDADA A Plate 2. Whole mounts, cross sections and cuticular scale casts of Elephantulus edwardi (bars equivalent to 50 um). Michael F. SCHNEIDER & Victorino A. BURAMUGE: Atlas of Microscopic Hair q. E GE? + ¢ ‘wig? E me + Plate 3. Whole mounts, cross sections and cuticular scale casts of Elephantulus intufi (bars equivalent to 50 um). Uy Bonner zoologische Beitrage 54 (2005) 134 ee E WUC Cceescacweccarcaccn KANE eee see AAN an < EN PT TU ANI Ns SE en ee) lado má = verirrt re ee] INN A 1 u a u a nn ici il Bass Ba OF, q soe ose bad qe qe O Sins AA ct dd a ta id tl a ds J ALA sia a IM Plate 4. Whole mounts, cross sections and cuticular scale casts of Elephantulus myurus (bars equivalent to 50 um). Michael F. SCHNEIDER & Victorino A. BURAMUGE: Atlas of Microscopic Hair Cor UCHESHAIB ATE BACEAdGea: nn $$$ TT ey LLLLLCCLCALTLSITEL k Pp es] ON J > A EN JA \ TERN \ 3 y UN 2 \ at AN AA PY Adal : \ { , } \ \ ‘ { ite: dregs 4 | Q oR aa eb Plate 5. Whole mounts, cross sections and cuticular scale casts of Elephantulus rupestris (bars equivalent to 50 um). Bonner zoologische Beitráge 54 (2005) 136 Plate 6. Whole mounts, cross sections and cuticular scale casts of Macroscelides proboscideus (bars equivalent to 50 um). 137 Michael F. SCHNEIDER & Victorino A. BURAMUGE: Atlas of Microscopic Hair Plate 7. Whole mounts, cross sections and cuticular scale casts of Petrodromus tetradactylus (bars equivalent to 50 um). Bonner zoologische Beitráge 54 (2005) 138 ee A a II ‘OD OB 0D i fp >O999900M0Q000 | aaa EH ee ey, ve" >) E 5 EN A es URIS Y 5s Pye Plate 8. Whole mounts, cross sections and cuticular scale casts of Crocidura cyanea (bars equivalent to 50 um). 139 Michael F. SCHNEIDER & Victorino A. BURAMUGE: Atlas of Microscopic Hair O R By PAN / diia dde O SEE r Aloe a ROA o a TS ): Plate 9. Whole mounts, cross sections and cuticular scale casts of Crocidura flavescens (bars equivalent to 50 um Bonner zoologische Beitráge 54 (2005) 140 ee Haan ng QU Se DR ST er tf VN AA ES x SAN Zins ae Be f Aina SAR DORA ARE es nn ee od a . . >— HO TAN se Ap Se aout O Pe € aS 1 3 ANA aS E SA OA oes CP | el ISAS Plate 10. Whole mounts, cross sections and cuticular scale casts of Crocidura fuscomurina (bars equivalent to 50 um). Michael F. SCHNEIDER & Victorino A. BURAMUGE: Atlas of Microscopic Hair 14] 2 4 D rey DNA 3 4 —A ) Lae Plate 11. Whole mounts, cross sections and cuticular scale casts of Crocidura hirta (bars equivalent to 50 um). Bonner zoologische Beitráge 54 (2005) 142 TT o A u Se Sl ese namen ES E a O a ae ne A oO Plate 12. Whole mounts, cross sections and cuticular scale casts of Crocidura luna (bars equivalent to 50 um). Michael F. SCHNEIDER & Victorino A. BURAMUGE: Atlas of Microscopic Hair 143 pew y 1, po) Hh Seats ) Jf: \ la) ee ) woke gg ko hen » Bi nah, SEN EN Sn al ray Ji) ty ge pr / 1 1] x / A O H Plate 13. Whole mounts, cross sections and cuticular scale casts of Crocidura maquassiensis (bars equivalent to 50 um). 144 Bonner zoologische Beitráge 54 (2005) Plate 14. Whole mounts, cross sections and cuticular scale casts of Crocidura mariquensis (bars equivalent to 50 um). Michael F. SCHNEIDER & Victorino A. BURAMUGE: Atlas of Microscopic Hair I DE TS ~~ Be NS m y y q N Plate 15. Whole mounts, cross sections and cuticular scale casts of Crocidura olivieri martiensseni (bars equivalent to 50 um). 146 Bonner zoologische Beitráge 54 (2005) DAA A Zi A 4 Seen SESE or I BAHU AVR OC Plate 16. Whole mounts, cross sections and cuticular scale casts of Crocidura olivieri occidentalis (bars equivalent to 50 um). 147 Michael F. SCHNEIDER & Victorino A. BURAMUGE: Atlas of Microscopic Hair e Y o da as ON, DN rr Plate 17. Whole mounts, cross sections and cuticular scale casts of Crocidura silacea (bars equivalent to 50 um). 148 Bonner zoologische Beitráge 54 (2005) _ Der é GG N vn Neyer Te le Plate 18. Whole mounts, cross sections and cuticular scale casts of Crocidura turba (bars equivalent to 50 um). Michael F. SCHNEIDER & Victorino A. BURAMUGE: Atlas of Microscopic Hair 149 Ma MN — > tie. BE we we: m4 , fj i 4 | | { ? N | | Y M zy ® Plate 19. Whole mounts, cross sections and cuticular scale casts of Mvosorex cafer (bars equivalent to 50 um). Bonner zoologische Beitráge 54 (2005) 150 Ñ ke > 4 { ra E kr > LECCE AA a el a ii a bok Sih Wii Wi Plate 20. Whole mounts, cross sections and cuticular scale casts of Myosorex longicaudatus (bars equivalent to 50 um). Michael F. SCHNEIDER & Victorino A. BURAMUGE: Atlas of Microscopic Hair Ion PAE Y "Ze —s Lig Cas WTA in Plate 21. Whole mounts, cross sections and cuticular scale casts of Myosorex sclateri (bars equivalent to 50 um). Bonner zoologische Beitráge 54 (2005) Beats eres OTe BIETE TE FEN, Lely IL Y LIA Se PESAS AN á. - A ang » rg AS IO JS E IA a wc ee Da o y RARA TATI TER CC 67 A AS EEE NN WE, a a ar TN AVES OS De Am rn bocha pal) 77 a Ay Pr —— da BE) = ti he ET 2: Plate 22. Whole mounts, cross sections and cuticular scale casts of Myosorex tenuis (bars equivalent to 50 um). Michael F. SCHNEIDER & Victorino A. BURAMUGE: Atlas of Microscopic Hair — =r — h \ nz Plate 23. Whole mounts, cross sections and cuticular scale casts of Mvosorex varius (bars equivalent to 50 um). in w 154 Bonner zoologische Beiträge 54 (2005) Plate 24. Whole mounts, cross sections and cuticular scale casts of Suncus infinitesimus (bars equivalent to 50 um). Michael F. SCHNEIDER & Victorino A. BURAMUGE: Atlas of Microscopic Hair Plate 25. Whole mounts, cross sections and cuticular scale casts of Suncus lixus (bars equivalent to 50 um). 155 156 Bonner zoologische Beitráge 54 (2005) \ - 1 ) sae ti bi) i. MN Plate 26. Whole mounts, cross sections and cuticular scale casts of Suncus varilla (bars equivalent to 50 um). Michael F. SCHNEIDER & Victorino A. BURAMUGE: Atlas of Microscopic Hair Plate 27. Whole mounts, cross sections and cuticular scale casts of Sy/visorex megalura (bars equivalent to 50 um). 157 ge 54 ( he Beiträge Oo Bonner zoologisc t 4 j a = Wt ane ik ae a eee ma Plate 28. Whole mounts, cross sections and cuticular scale casts of Atelerix frontalis (bars equivalent to 50 um). Michael F. SCHNEIDER & Victorino A. BURAMUGE: Atlas of Microscopic Hair sinter A!" AGRA ye Plate 29. Whole mounts, cross sections and cuticular scale casts of Amblysomus hottentotus (bars equivalent to 50 um). Bonner zoologische Beitrage 54 (2005) 160 cuticular scale casts of Amblysomus iris (bars equivalent to 50 um). 30. Whole mounts, cross sections and Plate Michael F. SCHNEIDER & Victorino A. BURAMUGE: Atlas of Microscopic Hair 16] A A | N N CT eee iy Plate 31. Whole mounts, cross sections and cuticular scale casts of Calcochloris obtusirostris (bars equivalent to 50 um). 162 Bonner zoologische Beitráge 54 (2005) Plate 32. Whole mounts, cross sections and cuticular scale casts of Carpitalpa arendsi (bars equivalent to 50 um). Michael F. SCHNEIDER & Victorino A. BURAMUGE: Atlas of Microscopic Hair e MTB { y Dr ay} ie RAY wala eet hem PS A a. AA Plate 33. Whole mounts, cross sections and cuticular scale casts of Chlorotalpa duthieae (bars equivalent to 50 um). 163 Bonner zoologische Beitráge 54 (2005) 164 Dy Plate 34. Whole mounts, cross sections and cuticular scale casts of Chlorotalpa sclateri (bars equivalent to 50 um). 165 Michael F. SCHNEIDER & Victorino A. BURAMUGE: Atlas of Microscopic Hair FU Pe pego! LAREDO MILT oN ae , rv if oy NE ARA SII eee Á ad SASS SNORT Bi daa > A X EN icons Ri aa a SS >, > ae En y A de A ESA y ¡E a IRAN, E 4 de 4 ‘ 4 a oi ER Rt: un 5 i ' A x 4 4 oF : y Je) e, rad A TR if el. gps 15% ft: Saeed pes DI wiki BORG AA IL ERE SER RE P= — Plate 35. Whole mounts, cross sections and cuticular scale casts of Chrysochloris asiatica (bars equivalent to 50 um). Bonner zoologische Beitráge 54 (2005) 166 =~ IM, MANN RN SIE a € > cular scale casts of Chrysospalax trevelvani (bars equivalent to 50 um). Whole mounts, cross sections and cuti Plate 36. Michael F. SCHNEIDER & Victorino A. BURAMUGE: Atlas of Microscopic Hair : ¥ 1 7 A q 7 ? } + Plate 37. Whole mounts, cross sections and cuticular scale casts of Chrysospalax villosus (bars equivalent to 50 um). 167 Bonner zoologische Beitráge 54 (2005) 168 Plate 38. Whole mounts, cross sections and cuticular scale casts of Cryptochloris wintoni (bars equivalent to 50 um). Michael F. SCHNEIDER & Victorino A. BURAMUGE: Atlas of Microscopic Hair 169 I Plate 39. Whole mounts, cross sections and cuticular scale casts of Eremitalpa granti (bars equivalent to 50 um). Bonner zoologische Beitráge 54 (2005) 170 Ar gunningi (bars equivalent to 50 tm). cross sections and cuticular scale casts of Neamblysomus Plate 40. Whole mounts, Michael F. SCHNEIDER & Victorino A. BURAMUGE: Atlas of Microscopic Hair 171 Plate 41. Whole mounts, cross sections and cuticular scale casts of Neamblysomus julianae (bars equivalent to 50 um). Bonner zoologische Beitráge 54 (2005) ww) we: sera Sara mas na re E Plate 42. Comparison of cuticular scale structure using electron microscopic scans and light microscopy of scale casts of Neam- blysomus iris (A), Chlorotalpa arendsi (B), C. sclateri (C), Crocidura flavescens (D) and Elephantulus rupestris (E) (bars of light microscopic scale casts equivalent to 50 um). 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Sinclair Dieter Stüning Bonn, Germany ho Bonner zoologische Beitráge vol. 54, Index ASLAN, IRFAN; BEENEN, RON & OZBET, HIKMET: Biological aspects of Galeruca circassica Reitter, 1889 (Coleoptera: Chrysomelidae: Galerucinae) in Relation to the Weed Cephalaria procera Fish. and Lall. PADIS SAG AA le NN ASSING, VOLKER: A Revision of Western Palaearctic Medon: the Species of the Atlantic Islands, the Western Mediterranean, and Europe, Except for the Southeast (Insecta: Coleoptera: Styphalinidae: Paederinae)....... ASSING, VOLKER: A Revision of Porocallus Sharp. New Synonyms and New Species (Insecta: Coleoptera: Stapkylinidae Aleocharmac: OXypodini) A sico BEENEN, RON: Translocation in Leaf Beetles (Coleoptera: Chrysomelidae) ...u..u.uue00. nennen DUNGELHOEF, SUSANNE & SCHMITT, MICHAEL: Functional Morphology of Copulation in Chrysomelidae- Crioeennae and Bruchidae (insecta? Colo pies a) ics sais urn een FURTH, DAVID G.: The Current Status of Knowledge of the Alticinae of Mexico (Coleoptera: Chrysomelidae) GEISER, ELISABETH: Frozen Chrysomelids from Alpine Glaciers — Witnesses of the Postglacial Resettlement . GROSS, JURGEN & FATOUROS, NINA E.: Striking Differences in Behaviour and Ecology Between Populations OE CHP VSCOM A TAD A A O RE HUBWEBER, LASSE & SCHMITT, MICHAEL: Parameres — Similarities and Differences in Chrysomelidae and (erambyeitastCokoper) nee ee hen AS mene ieee PETERS, GUSTAV: Besprechung von/review on Schneider, H. (2005): Bioakustik der Froschlurche — Einheimischeund' verwandte Arten ds e een SCHMITT, MICHAEL & BOPP, SIGRUN: Leaf Beetles (Insecta: Coleoptera: Chrysomelidae) Suffer From Feeding on Peri Lew. a ee SCHNEIDER, MICHAEL F. & VICTORINO A. BURAMUGE: Atlas of the Microscopic Hair Structure of Southern African Shrews, Hedgehogs, Golden Moles and Elephant-shrews (Mammalia) .............:::escesseeseeeeeeeeeneeeeees SCHOLLER, MATTHIAS: The Genus Achaenops Suffrian, 1857 (Chrysomelidae: Cryptocephalinae), Designation of Neotypes and Description of New Species. una VAN DEN ELZEN. RENATE: Andreas Helbie Y sida NAAA VENCL, FREDRIC V. & Allen, BENGT J.: Failure-time Analyses of the Effectiveness of Larval Shield Defenses in Tortoise Beetles (Chrysomelidas: Cassidmas anna ae ee VERMA, KRISHNA K. & JOLIVET, PIERRE: On Phyletic Closeness Between South American and New Caledonian Spilopyrines (Chrysomelidae, Eumolpinae, Tribe Spilopyrini) «...............::ssessesesseaseceentesccacsees VIG, KAROLY & MARKO, VIKTOR: Species Composition of Leaf Beetle Assemblages in Deciduous Tree Canopies: in Hungary (Coleoptera: ChrysomelldaS) .. urn a eae WAGENER, SIGBERT (7): Butterfly Diversity and Protection in Turks... na Publication dates: No. 1/2: 30. April 2006, No. 3: 30. August 2006, No. 4: 30. October 2006 173-177 25-95 97-102 179-199 201-208 209-237 239-245 247-252 253-259 96 261-269 103-172 271-286 1-2 287-295 297-303 305-312 3-23 o Bonner zoologische Beitráge Band 54 (2005) | Heft 4 | Seiten 173-177 Bonn, Oktober 2006 Biological Aspects of Galeruca circassica Reitter, 1889 (Coleoptera: Chrysomelidae: Galerucinae) in Relation to the Weed Cephalaria procera, Fish. and Lall. (Dipsacaceae) in Anatolia' Irfan ASLAN”, Ron BEENEN? & Hikmet OZBEK”? Atatürk University, Agricultural Faculty, Plant Production Department, Erzurum, Turkey Nieuwegein, The Netherlands Abstract. Biological aspects of Galeruca circassica, a herbivore feeding on Cephalaria procera, were studied in Tur- key. In the laboratory, adults oviposited on Cephalaria procera from 9 days after emergence and laid an average of 198 eggs per female. Development of eggs and larvae proved to be temperature dependant. Larvae passed through four in- stars and pupated in a pupal case in the soil. Pupal stage lasted 48 days. Laboratory tests indicated that this species can complete its development on several Cephalaria species and on Salvia staminea and Centaurea solstitialis. Only on Cephalaria procera and C. hirsute a substantial percentage reached the adult stage. Galeruca circassica should be pre- served in Eastern Anatolia because it most probably contributes in regulating the abundance of Cephalaria procera and because it might be used in biocontrol. The biology and host specificity of G. circassica indicates that it has a potential to be an agent for the biological control of C. procera in situations where this plant species occurs as an invasive plant. Key words. Chrysomelidae, leaf beetle, Galeruca circassica, Cephalaria procera, biological control, weeds 1. INTRODUCTION Cephalaria procera Fisch and Lall., 1840 (Dipsacaceae) is distributed in North Iran, Armenia and Turkey (DAVIS 1972). It is a perennial plant with subglabrous or sparsely pilose stems up to two meters. The flowers are pale yellow or cream. Cephalaria procera is very close to C. gigantea, possibly it has to be reduced to a sub- species of C. gigantea (DAVIS 1972). The Turkish name of Cephalaria procera is “Gevrek”. Cephalaria plants root deep and the roots are strong which can be prob- lematic in cultivating the ground (ploughing). In Eastern Anatolia “Gevrek” is regarded an important weed in meadows and pastures. Therefore a study was con- ducted of one of the herbivorous species associated with this plant and attempts were made to determine the in- fluence of this species on Cephalaria. Cephalaria is associated with several leaf beetle spe- cies. MEDVEDEV & ROGINSKAJA (1988) mention Gale- ruca pomonae (Scopoli) and G. tanaceti (L.) (Galeruci- nae) as polyphagous species feeding on Cephalaria. DOGUET (1994) mentions the polyphagous Longitarsus luridus (Scopoli) (Alticinae) feeding on Cephalaria mauritanica in Northern Africa and Cephalaria syriaca in Israel. In Turkey Galeruca is the only leaf beetle ge- nus associated with Cephalaria (ASLAN pers. obs.). The subfamily of Galerucinae comprises six tribes and approximately 5800 species (WILCOX 1971-1975) dis- 1 th Paper included here, but not presented to the 6” International Sym- posium on the Chrysomelidae, Bonn, Germany, May 7, 2004. tributed throughout the world but predominantly in the tropics. In Turkey, this subfamily is known with 14 genera and 52 species (ASLAN 1998; ASLAN et al. 2000). In this subfamily, larvae feed on leaves or on plant roots in the soil, rarely on fruits. The biology of most species in this subfamily is unknown, except for the biology of pest species such as Agelastica alni (L.), Aulacophora foveicollis (Lucas) and Xanthogaleruca luteola (Miiller) (ASLAN et al. 2000). On the other hand, there have been some attempts to study the biology of Galerucella calmariensis (L.) and G. pusilla in order to use them as biocontrol agents for management of Ly- thrum salicaria in the United States and some other countries (MCFAYDEN 1998). The genus Galeruca is represented in Turkey by ap- proximately 10 species (ASLAN et al. 2000). The ge- nus is currently revised (BEENEN 1999, 2002, 2003) and possibly this will affect the number of Turkish species. 2. MATERIAL AND METHODS 2.1. Taxonomy The specimens of Galeruca observed in this study be- long to the Galeruca pomonae-group. From this spe- cies-group ASLAN et al. (2000) include G. fuliginosa (Joannis) and G. pomonae in their preliminary list of Turkish species. The inclusion of G. fuliginosa is based only on the type specimen collected in “Grece et Cau- case”. BEENEN (1999) concluded that G. fuliginosa is to be treated as a junior synonym of G. littoralis (F.) and 174 Bonner zoologische Beitráge 54 (2005) that it is unlikely that it has been captured in the Cauca- sian mountains. The other species, G. pomonae, is a widely distributed and variable species. Many local forms have been described as different species and sub- sequent authors have synonymised these with G. pomonae. One of these, G. circassica Reitter, has been synonymised with G. pomonae by lABLOKOFF-KHNZO- RIAN (1968). ASLAN (1998) followed IABLOKOFF- KHNZORIAN in this respect. BEENEN (2005) gives argu- ments to remove G. circassica from synonymy and re- established it as a valid species. The results presented in the next paragraphs are based on the study of Turkish specimens of G. circassica. 2.2. Development Adults and larvae of G. circassica were collected from Gólyurt Pass (Ispir, Erzurum), Olur (Erzurum) and Askale (Erzurum). The altitudes in the localities differed between 1000 m. and 2450 m. The collected specimens were used to start laboratory studies. Larvae and adults were placed in cages (50 x 45 cm) and kept at 20 (+ 3) °C and 45 (+ 10) % humidity. Fresh leaves of Cephalaria procera taken from the same localities were placed in the cages. In order to keep the leaves fresh their petioles were dipped in glass vials (15 ml) filled with tap water. The leaves were replaced every four days. The bottom of the cages was covered with a layer of soil (three cm) in which larvae could pupate. Cages with pupal cases were retained in laboratory, under natural photoperiod (during July approximately 12 hours dark). Eggs were removed every 24 hours. —$— 25 > | I 3 | E larvae 3 20 + a = 45: HB | ¢ Se | E 10 - | 2 : al i oO a | 04 1 15 20 25 30 Temperature Fig. 1. Development period of larvae and egg in different temperature. To determine the effect of temperature on egg develop- ment, eggs less than 24 hours old were placed, still at- tached to C. procera, on moist filter paper in petri dishes in each of four constant environment chambers at 15: °C. (n= 25), 20 "C (n= 25); 25 °C (125 and 30°C (n=25) (RH 65 + 5 %; 12h L: 12 h D). Twice every day, at 8:30 and 19:30, newly hatched larvae were counted and removed. 1 2 3 4 Instar stage Fig. 2. Days taken for each instar stage to develop at 15, 20, 25 and 300C. To determine the effect of temperature on larval devel- opment newly hatched larvae were kept in constant en- vironment chambers at 15 °C (n= 20), 20 °C (n= 20), 25 °C (n=20) and 30 °C (n=20) (RH 65 + 5 %; 12h L: 12h D). Each larva was placed on a fresh leave. In order to keep the leaves fresh their petioles were dipped in glass vials (15 ml) filled with tap water. Larvae were moni- tored twice daily at 8:30 and 19:30. Leaves were re- placed every other day. After the final moult, the bottom of each cage was covered with soil as above. 2.3. Foodplants In the province of Erzurum the vegetation was studied in Ispir, Pasinler, Olur, Askale and Oltu. Plant species of the genera Cephalaria, Salvia, Centaurea, Cirsium and Scabiosa were intensively examined for the pres- ence of G. circassica eggs, larvae, and adults. Each plant species, regardless the number of individuals, was examined for 15 minutes. In addition also laboratory tests were conducted. Five species of Cephalaria including C. procera, two species of Centaurea and Salvia and one species of Cirsium and Scabiosa were tested in no-choice tests for feeding. Adults manually removed from pupal cases 25 to 35 days after pupation were used in tests for feeding. Six adults (3 males and 3 females) were placed on each leaf for the above mentioned species. In larval test three first instar larvae were placed on each plants leaf. In order to keep the leaves fresh their petioles were dipped glass vials filled with tap water. The leaves were replaced every four days. The no-choice tests were carried out in the laboratory under natural light condition at 20 + 3 40 and a relative humidity of 45 + 10%. All plants were tested separately using three replicates. 2.4. Predators and parasites In the province of Erzurum, eggs, larvae and pupae were observed to identify possible predators or para- Irfan ASLAN, Ron BEENEN & Hikmet ÓZBEK: Galeruca circassica in Anatolia 175 sites. Additionally eggs, larvae and pupae were col- lected and placed in a cage for emerging parasites. 3. RESULTS 3.1. Development Eggs of G. circassica were 1.9+0.015 mm (after the + sign the standard deviation is indicated) long 1.5+0.005 mm (n=25) wide and deposited in groups at the under- side, rarely at the upper side, of the leaves. Eggs were ini- tially translucent to dirty white, becoming darker during development. Egg development took 9.8+0.17 (n=30), 6.3+0.14 (n=30), 4.7+0.08 (n=30), and 4.2+0.04 (n=30) days at 15, 20, 25, and 30 °C, respectively (Fig. 1). Larvae emerged (head first) and immediately started feeding on the foliage. The black larvae passed through four instars, taking 19.5+0.17 (n=30), 16+0.14 (n=30), 13.4+0.11 (n=30) and 12.7+0.08 (n=30) days from emergence to the construction of the pupal case at 15, 20, 25, and 30 °C, respectively (Figs. 1, 2). Larval mor- tality was lowest at 20 °C and highest at 30 °C, espe- cially during the first instar stage. Mature larvae dropped to the ground where they pupated in a pupal case in the soil. In the laboratory 80.5 % (145/180) of the larvae that en- tered the pupal stage emerged as -adults. Pupal stage lasted 47.8 (+1.86) days and was independent of tem- perature. On the locations were field studies were conducted ac- tive G. circassica adults were present mainly from the beginning of May to the end of June. Galeruca circas- sica eggs and larvae were found on C. procera in from mid of May to the beginning of July. 3.2. Food plants Eggs, larvae and adults of G. circassica were recorded in the field on plant species of the families Dipsacaceae, Lamiaceae and Asteraceae (Table 1). The highest pres- ence occurred on plant species of Cephalaria, especially C. procera. Salvia staminea and Centaurea solstitialis were the only species in the subfamilies Lamiaceae and Asteraceae on which G. circassica were registered. Table 1. Plant species monitored in the field in Turkey for presence of G. circassica. Family Species Sites” Occasions” Present‘ Dipsacaceae Cephalaria aristata 10 5 2 C. hirsute 20 12 12 C. speciosa 15 10 3 C. anatolica 13 8 2 C. procera 48 1415 45 C. syriaca 12 12 4 C. media 8 4 I C. gigantean 18 9 2 Lamiaceae Salvia staminea 5 4 | S. verticillata 4 5 0 S. brachyantha 3 2 0 Asteraceae Centaurea depressa 8 9 0 C. solstitialis 23 39 3 C. iberica 11 5 0 C. virgata 9 2 0 Asteraceae Cirsium arvense 38 76 0 C. ciliatum 5 3 0 Dipsacaceae Scabiosa rotata 11 5 0 S. columbaria 4 2 0 “Sites monitored; "Occasions each site monitored; “Sites Galeruca circassica present. No-choice tests revealed that G. circassica laid eggs on all plant species on which this Galeruca species has been observed in the field. Most eggs were laid within nine days of the adults’ removal from pupal cases. Each female deposited on average 198 eggs. Although intense feeding was observed on only one species (C. procera showing severe damage) many adults survived for 45 days on most species tested (Table 2). Larvae completed their development on Cephalaria spp. and two other species, Salvia staminea and Centaurea solstitialis (Ta- ble 2). 176 Bonner zoologische Beitráge 54 (2005) Table 2. Plant species tested in larval and adult no-choice test. Larvae Test plants n” % Pupated” Damage" Cephalaria 10 85 S procera C. hirsute 8 50 S C. arsistata 5 20 M C. speciosa 5 10 M C. anatolica = 8 M Salvia staminea 3 2 NE S. verticillata 3 0 N Centaurea 3 0 N depressa C. solstitialis 3 3 M Cirsium arvense 3 0 N Scabiosa rotata 3 0 N Adults n* % Alive" Eggs“ Damage 9 80 250 S 5 78 175 M 4 70 60 M 5 72 20 NE 2 69 18 NE I 67 32 NE 0 60 0 N 1 69 0 N 2 58 29 NE 0 32 0 N 0 55 0 N “Number of replicates; "Mean percentage of larvae for replicates reaching the pupal stage; “Estimated damage (N, none; NE, neg- ligible; M, moderate; S, substantial); “Mean percentage of adults alive for all replicates after 45 days; “Total number of eggs laid after 45 days. 3.3. Predators and parasites In this study no natural enemies were found that were associated with G. circassica. However, one Miridae (Heteroptera) species, Deraecoris seranus Del, was seen feeding on the eggs of G. circassica, but their den- sity was low. On the other hand, this species is zoophy- tophagous and feeds generally on alfalfa, some other cultivated plants and weeds. From eggs, larvae or pupae no parasites emerged. 4. DISCUSSION Life history tables of species of Galeruca (subgenus Galeruca), as far as known, show some remarkable similarities. The minor differences seem to be related to the exact months of summer heat and severity of winter. In the North-western part of its distribution Galeruca pomonae shows adult aestivation during summer (July and August) and reproduction in late summer and au- tumn (from September to the end of October) (BEENEN 1998). In Sardinia Galeruca sardoa (Gené) show sum- mer aestivation from the end of June to the start of Oc- tober. Reproduction takes place from November to January (CROVETTI & USCIDDA 1978.). In Sicily Galeruca species (G. sicana (Reiche) and G. reichei (Joannis)) show summer aestivation during July and August and reproduction from September to December (SINACORI & MINEO 1993). All these species show egg diapause during winter. The life history of G. circas- sica, a species hitherto confused with other Galeruca species, is unknown. The results of this study however show remarkable differences with the above mentioned species. Galeruca circassica is known to overwinter in the adult stage, mate and deposit eggs in spring and completely develops during early summer. From this study 1t became clear that development takes place in approximately 70 days. Eggs were seen in the field from mid May onwards, consequently adults could be seen some 70 days later: from the end of July. The results from the laboratory studies fit the results from the field. Adults of G. circassica overwinter and reproduce in spring. This life history table is in concordance with the observations on Galeruca rufa Germar by BOURDONNÉ & MALDES (1995). In the south of France Galeruca rufa deposits eggs from mid May to the end of August. Lar- vae are active during summer. Adults can be seen al- most all year. Galeruca rufa is not related to Galeruca circassica. It is classified in the subgenus Emarhopa; G. circassica in the subgenus Galeruca. From eggs, larvae or pupae no parasites emerged. Only a single predator has been observed. Most probably this subject needs some additional research. Other Galeruca species are known to be parasitized by Hymenoptera and Diptera parasites (COX 1994) and predated by Zicrona (Heteroptera), Lebia (Carabidae) and Aiolo- caria (Coccinellidae) (COX 1996). It is unlikely that Galeruca circassica has no parasites. The distribution of G. circassica is insufficiently known due to the taxonomic entanglement. This study provides information on the occurrence in eastern Anatolia. The description of G. circassica was based on specimens collected in Fisht in the north-western part of the Cau- casian mountain range (REITTER 1889). Further study of collections and surveys in the field must provide knowl- edge about its distribution. Food plants of G. circassica mentioned in literature are not reliable because of the uncertain taxonomic position Irfan ASLAN, Ron BEENEN & Hikmet OZBEK: Galeruca circassica in Anatolia 177 of this species. The same may hold true for G. pomonae. Records of G. pomonae from the Caucasian area could refer to G. circassica. GRUEV & TOMOV (1986) and WARCHALOWSKI (1994) indicate that G. pomonae could feed on Centaurea, Salvia, Knautia, Cirsium, and Scabiosa. MEDVEDEV & ROGINSKAJA (1988) mention Mentha and Melissa as possible (marked with question mark) food plants for G. circassica. These authors men- tion Cephalaria as food plant for G. pomonae. The observation in our study show that G. circassica feeds primarily on C. procera and C. hirsute; plants which are widespread and abundant in eastern Anatolia. These plants are important weeds and G. circassica most probably contributes in regulating their abundance. However, because both Cephalaria and G. circassica are indigenous in this region it is unlikely that G. cir- cassica will control this food plant completely. In situa- tions were Cephalaria is introduced and causes prob- lems, the possibility of using G. circassica as a biological control agent should be seriously investi- gated. Our study revealed that both adults and larvae of G. circassica feed on C. procera and cause important damage. In Anatolia a decline of G. circassica (e.g., a decline due to the use of non-specific insecticides) should be prevented because this leaf beetle can help in restricting the nuisance caused by “Gevrek”. Furthermore it is a potential agent for the biological control of Cephalaria in other places. REFERENCES ASLAN, I. 1998. Erzurum [li Galerucinae (Coleoptera, Chrysomelidae) altfamilyası üzerinde faunistik ve sis- tematik bir calisma. Türkiye Entomoloji Dergisi 22: 285-298. ASLAN, I. & ÖZBEK, H. 1999. Erzurum Ili’nde bazı yabancı otlarda beslenerek önemli derecede zarar veren yaprak böcekleri (Coleoptera, Chrysomelidae). Türkiye IV. Biyolojik Mücadele Kongresi Bildirileri, 26-29 Ocak 1999, Adana, 75-86. ASLAN, I., WARCHALOWSKI, A. & ÖZBEK, H. 2000. A Pre- liminary Review of the Subfamily Galerucinae (Col- eoptera, Chrysomelidae) in Turkey. Journal of the En- tomological Research Society 2: 27-42. BEENEN, R. 1998. Patterns in the distribution of Galeruci- nae in The Netherlands. Proceedings of the Fourth In- ternational Symposium on the Chrysomelidae. Atti Museo Regionale di Scienze Natural 1998: 7-16. BEENEN, R. 1999. Revisional notes on Galeruca 1. Ento- mologische Blatter 95: 85-92. BEENEN, R. 2002. Revisional notes on Galeruca 2. Ento- mologische Blatter 98: 21-28. BEENEN, R. 2003. Galeruca (Galemira n. subgen.) sub- costata n. sp. from Pakistan. Revisional notes on Galeruca 3. Stuttgarter Beiträge zur Naturkunde (Serie A Biologie) 648: 1-9. BEENEN, R. 2005. The identity of Galeruca circassica Reit- ter and G. bicolor Kollar & Redtenbacher. (Revisional notes on Galeruca 4). Entomologische Blatter 101: 13- 20. , BOURDONNE, J. C. & MALDES, J. M. 1995. Galeyuca (Emarhopa) rufa Germar et Galeruca (Haptoscelis) melanocephala Ponza, especes rares en France. Bulle- tin ACOREP 21: 11-18. Cox, M. L. 1994. The Hymenoptera and Diptera parasi- toids of Chrysomelidae. Pp. 419-467 in: JOLIVET, P. H., Cox, M. L. & PETITPIERRE, E. (eds.) Novel aspects of the biology of Chrysomelidae. Kluwer Academic Publ., Dordrecht. etc. Cox, M. L. 1996. Insect predators of Chrysomelidae. Pp. 23-91 in: JOLIVET, P. H. A. & Cox, M. L. (EDS.) Chry- somelidae Biology, Vol. 2. SPB Academic Publ., Am- sterdam. CROVETTI, A. & USCIDDA, C. 1978. Riceche eco-etholo- giche su Galeruca sardoa (Gené). Frustula Entomolo- gica (Nuova Serie) 1: 45-97. Davis, P. H. 1972. Flora of Turkey and the East Aegean Is- lands 4. 657 pp., Edinburgh at the University Press. DOGUET, S. 1994. Coléopteres Chrysomelidae, volume 2 Alticinae. Faune de France 80: I-IX, 1-694. GRUEV, B. & Tomov, V. 1986. Fauna Bulgarica, 13 Col- eoptera, Chrysomelidae, Part Il, Chrysomelinae, Galerucinae, Alticinae, Hispinae, Cassidinae. 388 pp., In Aedibus Academia Scientiarum Bulgaricae. IABLOKOFF-KHNZORIAN, S. M. 1968. Notes sur les Chry- somelidae de l’Armenie Soviétique. Annales de la So- ciété entomologique de France (N. S.) 4: 259-277. MCFADYEN, R.E.C. 1998. Biological control of weeds. Annual Review of Entomology 43: 369-393. MEDVEDEV, L. N. & ROGINSKAJA, E. J. 1988. A catalogue of foodplants of USSR Chrysomelidae. 192 pp., Aka- demia Nauk SSSR, Moscow. LOPATIN, I. 1977. Leaf-beetles (Chrysomelidae) of Middle Asia and Kazakhstan. 268 pp., Nauka, Leningrad. REITTER, E. 1889. Neue Coleopteren aus Circassien. Wie- ner Entomologische Zeitung 8: 97-104. SINACORI, A. & MINEO, G. 1993. Nota preliminare su Galeruca spp. delle Madonie. Frustula Entomologica (Nuova Serie) 16: 97-110. WARCHALOWSKI, A. 1994. Chrysomelidae, Stonkowate (Insecta, Coleoptera) Czesé IV. Fauna Polski 16: 1- 301. WILCOX, J. A. 1971-1975. Chrysomelidae. Galerucinae. Coleopterorum Catalogus Supplementa 78: 1-770. Authors’ addresses: Irfan ASLAN and Hikmet ÖZBET, Atatúrk University, Agricultural Faculty, Plant Produc- tion Department, 25240 Erzurum, Turkey, E-mail: iaslan@atauni.edu.tr and aslanir@hotmail.com: Ron BEENEN (corresponding author), Martinus Nijhoffhove 51, NL 3437 ZP Nieuwegein, The Netherlands. E-mail: r.beenen@wxs.nl Bonner zoologische Beitráge j Band 54 (2005) | Heft 4 | Seiten 179-199 Bonn, Oktober 2006 Translocation in Leaf Beetles (Coleoptera: Chrysomelidae)' Ron BEENEN Nieuwegein, The Netherlands Abstract. An overview of leaf beetle translocations, and the possible effects related to endangerment of native leaf bee- tle species is presented. Translocation is defined as the movement of living organisms from one area to another across natural barriers. In the new area the translocated organism lives free. Translocations can be intentional or accidental. The database of records of translocated leaf beetles is compiled using published records as the main source of informa- tion. In this study 126 leaf beetle species have been recorded to have been translocated at least once. Most translocated species occur in the subfamilies Chrysomelinae, Galerucinae, Alticinae, Cassidinae and Hispinae. Most unintentional successful translocations occur in mono- and oligophagous species that develop on cultivated and non-indigenous plant species. At present, no negative effects of translocated Chrysomelidae on biodiversity have been reported. The success- ful control of imported weeds by introduced Chrysomelidae can be seen as a positive effect to biodiversity since some of these weeds prevent natural processes in ecosystems. Historical distribution patterns of Chrysomelidae are discussed in relation to the translocations. Keywords. Coleoptera, Chrysomelidae, invasive species, bioinvasion, alien species, translocation, introduction 1. INTRODUCTION One of the major threats to native biological diversity is now acknowledged by scientists and governments to be biological invasions caused by alien invasive species (IUCN 2000, UNION OF CONCERNED SCIENTISTS 2001). This is not a new item: the threat of invasive species for the conservation of biodiversity was already pointed out by ELTON in 1933 (cf. SIMBERLOFF’s foreword in ELTON 2000). Meanwhile it is supposed that every year a wide range of alien species are imported into many countries through international trade both intentionally as trade products (horticulture, pets, etc.) or uninten- tionally (for example by hitch-hiking on legitimate products). Little is known about the number of leaf bee- tles (Coleoptera, Chrysomelidae) that are translocated in this way or about their impact on biological diversity. Translocated species can be a threat in the way that a native leaf beetle is displaced by an alien leaf beetle species. On the other hand translocated species can re- duce indigenous plant species and thus affect the native herbivores depending on it. In this study we restrict con- sideration to leaf beetle translocations and their effects on native leaf beetle populations caused by competition. Interspecific competition is any interaction between two or more species populations which adversely affects their growth and survival (ODUM 1971). The importance of interspecific competition is not clear (STEWART 1996) but evidence for competition among phyto- ' Paper presented to the 6" International Symposium on the Chry- somelidae, Bonn, Germany, May 7, 2004. phagous species (including Chrysomelidae) exists (e.g., GONZALES-MEGIAS & GOMEZ (2003)). When a population of a non-indigenous species out competes a population of a native species, the invasive species must have reached a population level in which its competition can be effective. When the population level is too low, it is unlikely that there will be a large effect on the population of the indigenous species. CO- LAUTTI & MACISAAC (2004) developed a framework for defining different stages during invasions. After the new species is established a non-indigenous species may be localised and rare (a), widespread but rare (b) or wide- spread and dominant (c). In this last mentioned stage, out competing of indigenous species may be the result. Therefore, for our purpose, we need to know whether translocated Chrysomelidae have become established. That is the first step. Subsequently we would like to know the status of the established species. The aim of this study is to get an overview of leaf beetle introduc- tions, and the possible effects related to endangerment of native leaf beetle species. Natural range expansion of a species, as for example re- corded for Chaetocnema major Jacquelin-Duval by DOBERL (1994a), is not treated here. Here we consider all transport across natural barriers as translocation. Both intentionally (e.g., in biological control programs) as unintentionally. The IUCN (1987) defines transloca- tion more strictly and restricts to “introductions”, “re- introductions” and “re-stocking”. Introductions can be effective tools in the management of natural and man made environments. Re-introduction and re-stocking as an aim in nature management has not been used with 180 Bonner zoologische Beitráge 54 (2005) Chrysomelidae species. Thus, for this purpose we do not have to include the IUCN-restrictions. Here we define translocation as the movement of living organisms from one area to another across natural barri- ers. In the new area the translocated organism lives free. Translocations can be intentional or accidental. The role of man is not always clear in cases of translocation. 2. METHODS The database of records of intentionally or non- intentionally introduced leaf beetles is compiled using published records as the main source of information. Because not all published information was available for this study there is a chance that information is lacking; this is believed to be compensated by other publications in which similar information is available. Only the traditional leaf beetles have been studied, i.e. exclusive the Bruchidae. Subfamily names used in this study are the traditional as have been used by SEENO & WILCOX (1982). All species group names of Chrysomeli- dae in Appendix | are scientific names complete with au- thor. In the text they are without author names, except for those species that are not listed in Appendix 1. A translocation event is defined as a translocation from an area where the species is indigenous to an area where the species previously did not occur. In this respect the translocation of Chrysolina hyperici from England to Australia in 1930 and from France to Australia in 1980 are considered as two different translocation events. In the case of biocontrol, translocations with different tar- gets (host plants) are considered as different transloca- tion events. Excluded from the database are natural expansions. However we have to admit that it is not always clear to discriminate between natural range expansion and transport across natural barriers. This might be the case in species that are still in the phase of post-glacial ex- pansion. Also accidental translocations that have resulted in es- tablishment in artificial environments as in greenhouses are not included in this research. In the Netherlands for example the alticine Acrocrypta purpurea Baly has es- tablished in a greenhouse where during several genera- tions it damaged orchids (DOBERL 1994b). Acrocrypta purpurea originates from South East Asia and it is unlikely that it will survive in the Netherlands outside (green)houses. Sometimes it is difficult to define the exact moment of establishment. Leptinotarsa decemlineata, for example, was discovered in the Rocky Mountains and described in 1824. Suddenly, in 1859, it began devastating potato crops 100 miles west of Omaha, Nebraska, USA. Over the next few years, the beetle spread eastward to the At- lantic coast, which it reached in 1874. Leptinotarsa de- cemlineata became established in Europe following its introduction from the USA to Bordeaux, France in 1922 (after several unsuccessful attempts from 1876). The beetle spread rapidly in Europe despite intensive opera- tions to control it. It was first reported in Belgium and Spain in 1935, Luxembourg in 1936, the Netherlands and Switzerland in 1937, Austria in 1941, Hungary and the former Czechoslovakia in 1945, Poland and Roma- nia in 1947, and Turkey in 1949 (CAB-CPC 2003). In Appendix | the year 1922 is taken as the moment of es- tablishment in Europe although it has been found earlier in France and established many years later in some European countries. On the other hand a species may not have established in all parts of the country to which it is translocated. For example: all attempts to establish L. jacobaeae in On- tario, New Brunswick and Prince Edward Island were unsuccessful. In Canada it is at present established only in British Columbia. In the United States it is currently known from California and Oregon (LESAGE 1988). 3. RESULTS In this study 126 leaf beetle species have been recorded to have been translocated at least once. They are listed in Appendix 1. Because many species have been translo- cated more than once, a total number of 230 translocation events in Chrysomelidae is listed (Appendix 1). Of each event information is given on place of origin of the leaf beetle, place where it is translocated to, moment of trans- location, degree of success, host plant and source (refer- ence). In Figure 1 the length of each bar represents the to- tal number of species of each subfamily. Most translocated species occur in the subfamilies Chrysomeli- nae, Galerucinae, Alticinae, Cassidinae and Hispinae. In Figure 2 the length of the bar represents the number of translocations of each subfamily. The bar for the subfam- ily Hispinae is much larger than in Figure 1, indicating that some species have been translocated several times. Translocations can be intentional or unintentional. The number of intentional translocations exceeds the number of unintentional translocations (Table 1). This is not surprising. Unintentional translocations are hard to de- tect; they mostly have not been published. Publications on species that have been unintentionally translocated mostly will deal with species that have become estab- lished in the newly inhabited land. When established they are more likely to be observed. Of the unintentional translocated species 81 % (70 out of 86) have become established against 63 % (91 out of 144) of the inten- tionally translocated species. The successes of inten- tional translocations that are part of a biological control program are monitored. This means that not only infor- mation on successful translocations are available but also of the unsuccessful releases. On the other hand the intentional translocations mostly take place only after intensive study. In biological control programs the suc- cess (and the risk) of the introduction is carefully stud- ied before release. In that respect it is surprising to no- tice that 27 % (39 out of 144) of the intentionally translocated species have not become established. Ron BEENEN: Translocation in Leaf Beetles 18] 35 y Im 1 1 q „0 Do eer CRIO ZEUG CRYP EUMO CHLA CHRY GALE ALTI CASS HISP Fig. 1. Results of translocations in Chrysomelidae. Number of translocated species per subfamily. Subfamily abbreviations: CRIO = Criocerinae; ZEUG = Zeugophorinae; CRYP = Cryptocephalinae; EUMO = Eumolpinae; CHLA = Chlamisi- nae; CHRY = Chrysomelinae; GALE = Galerucinae; ALTI = Alticinae; CASS = Cassidinae; HISP = Hispinae. Results: UE = unintentional translocation followed by establishment; UN = unintentional, not established; UD = unintentional, establish- ment doubtful; IE = intentional and established: IN = inten- tional and not established; ID = intentional and doubtful. 80 5 70] [BID GIN 60 | (ale MUD 50 | MUN BUE 40 - 30 4 1 20 10 + ; E 0 a @ N HY Ei Pd EA . CRIO ZEUG CRYP EUMO CHLA CHRY GALE ALTI CASS HISP Fig. 2. Results of translocations in Chrysomelidae. Events per subfamily. Subfamily abbreviations as in Figure |. Results: UE = unintentional translocation followed by establishment; UN = unintentional, not established; UD = unintentional, es- tablishment doubtful; IE = intentional and established; IN = intentional and not established; ID = intentional and doubtful. Intentionally translocated species mostly have very re- stricted food preference. The chances that they become a problem as a result of feeding on non-target species is lowest in monophagous species. We analysed the food preference of the unintentionally translocated species. In Figure 3 the number of species is presented for the next groups of phytophages: monophages — species that de- velop on a single plant species or genus; oligophages — species that develop on more than one plant genus of a single plant family; polyphages — species that develop on plant species of more than one family. It shows that most of these successful translocated species are mono- or oligophagous; they develop on plant species within a single genus or within a single plant family. This more or less implies that, because of the food restriction of most leaf beetles, host plants from the same genus or family have to be available in the new country. It is interesting to find out what types of plants these successfully unintended translocated species feed on. Four categories of plants species have been distin- guished: cultivated plant species, non-indigenous “wild” plants, indigenous “wild plants” and a last category of which information was insufficient (Fig. 4). It is evident that the majority of unintended successful translocation of leaf beetles occurred on cultivated plants and on non- indigenous (introduced plant species). Only a minority occurs on plants that are indigenous. : species 30 + y H E events FF > HH HH HH] [ HEH mono oligo poly uncertain Fig. 3. Number of unintentionally translocated leaf beetles that have established per categories of food plants. Mono = mono- phagous, oligo = oligophagous, poly = polyphagous. Species = number of translocated species, events = number of transloca- tion events. 40 - 357 HH Dspecies 30 4 HHH B events 25 : 20 Fr 15 - | Het 10, | fea EE | 54 + = | | aH vi I - Tr | tt 0 PE E HH cultivated non-indig. indigen uncertain Fig. 4. Number of unintentionally translocated leaf beetles that have established per plant category: cultivated plants, non- indigenous plants, indigenous plants and plants of which this information is not available. 4. MISCELLANEOUS RESULTS Apart from the results described above, interesting in- formation became available on a variety of subjects. An extract of this is presented in this section. The following subjects are presented: description of species based on translocated specimens; unused biological control agents; the use of indigenous species in biological con- trol; anecdotal information on the ways unintentional translocations have taken place and finally non-target effects. 182 Bonner zoologische Beitráge 54 (2005) 4.1. Description of species based on translocated specimens Some species description were based on displaced specimens: Falsoplatyxantha diversicornis Pic, Lupe- rus marginalis Allard and Aulacophora pannonica Csiki (= Hoplasoma unicolor 11.) (all Galerucinae). The type specimen(s) of Falsoplatyxantha diversicornis were from Turkey, but the author already stated that the spe- cies most probably originally came from the Indies (“Acquis comme provenant d'Asie Mineure, mais origi- naire vraisemblablement des Indes’) (PIC 1931). Of Hoplasoma unicolor a single specimen was collected in Hungary and described as Aulacophora pannonica by CsIkI (1953). SILFVERBERG (1978) studied the type specimen of A. pannonica and reduced it to a junior synonym of H. unicolor. The type specimen of A. pan- nonica was probably imported from the Oriental region. Although there always remains the possibility of misla- belled specimens (see MOHAMEDSAID 2001). Luperus marginalis was described by ALLARD (1890) from the Falkland archipelago and according JOLIVET & VERMA (2002) evidently imported there in historical times. Apart from L. marginalis no Chrysomelidae have been mentioned from the Falkland archipelago. 4.2. Unused biological control agents Some species have been selected to be used as a bio- logical control agent but have not been used yet: Agen- iosa electoralis Vogel (Chrysomelinae) was selected to be introduced in Australia to control Chrvsanthemoides monilifera (JOLIVET 2001). Charidotes pygmaea Klug (Cassidinae) was selected to be released in Australia to control species of Lantana (JOLIVET 2001). Homichloda barkeri Jacoby (Alticinae) from Kenya was selected to control Acacia nilotica in Queensland (JOLIVET 2001). Trachyaphthona sordida (Baly) (Alticinae) a specialist herbivore on Paederia foetida (Skunk Vine) in China and Japan, which is a very invasive weed in Florida spreading in other parts of the Southern Unites States. Trachyaphthona sordida is of special interest to be be used as a biological agent (PEMBERTON & PRATT 2002). Several species have been selected for the control of in- vasive plants but finally introduction has been aban- doned because the species proved to be not selective enough in their food choice. This is the case for exam- ple for Platyphora biforis (Germar), P.conviva (Stal), P. nigronotata (Stal) and P. paraguana (Jacoby) for the control of Solanaceae (JOLIVET 2001). 4.3. The use of indigenous species in biological control Very rarely indigenous species are successful in control- ling nuisance herbs. Both Chelymorpha cassidea (F.) (Cassidinae) and Chirida guttata (Olivier) (Cassidinae) have been collected in 1979 in Saskatchewan and re- leased in Alberta in an attempt to extend their ranges for the control of Convolvulus arvensis, where they did not establish (JULIEN 1992). In order to control Calystegia sepium, native organism of Chirida guttata have been collected in Ontario in 1971 and released in British Co- lumbia. They did not establish either (JULIEN 1992). The same holds true for Metriona purpurata native organism collected in 1979 in Saskatchewan and released in Al- berta in an attempt to extend its range and control Con- volvulus arvensis (JULIEN 1992). However, Altica fovei- collis Jacoby (Alticinae) a native species of Thailand was used in this country to control Ludwigia adscen- dens. It caused considerable damage with satisfactory degree of control (JULIEN 1992). Gastrophysa atro- cyanea a native species introduced in Noto Peninsula (Japan) to control Rumex obtusifolius quickly estab- lished, increased its population and spread rapidly. Population of the beetle reached satisfactory levels within four years of release (JULIEN 1992). Metriona bi- color, a native organism collected in 1971 in Ontario and released in British Columbia in order to control Ca- lystegia sepium, established. In 1985 it was found 14 km from the release site (JULIEN 1992). Both Leptinotarsa defecta and L. texana are indigenous in the USA (Texas) and have been proposed as a bio- logical control agent against three invasive Solanum species. The results of the study suggest that of the in- vasive Solanum species, $. torvum may be included in the potential host range of L. texana. It is suggested that this “new association” approach might result in a sub- stantial control of one of Florida’s most invasive so- lanaceous weeds with acceptable ecological risks (CUDA et al. 2002). Native biological control agents offer potential advan- tages over non-native agents, because they may have lit- tle impact on non-target native species that have coex- isted with the control agent (SHELDON et al. 1995). 4.4. Registered unintentional translocations Very rarely aspects of unintentional translocations have been registered. In a few cases they can however be re- constructed. Sometimes unintentional introductions have been the result of intentional introduction during a biological control program. Aphthona nigriscutis was accidentally released in Canada in 1982 together with Aphthona cyparissiae. The release site was treated with herbicide and insecticide in order to eradicate this spe- cies (JULIEN 1992). And Chrysolina hyperici is believed to have been introduced accidentally in Hawaii together with C. quadrigemina in 1965. It was discovered in 1970 (JULIEN 1992) and JOLIVET (2001) assumes this species is established in Hawaii. In 1979 a Longitarsus species referred to as L. jacobaeae has been released in Australia from France. This species more closely re- sembles L. flavicornis. It is established and spreading, causing high reduction in weed density at some sites (JULIEN 1992). Longitarsus ganglbaueri, a European species, is re- cently found in Oregon. LESAGE (1988) assumed it to Ron BEENEN: Translocation in Leaf Beetles 183 be accidentally introduced with those of L. jacobaeae imported from Italy and introduced in Oregon. Xanthogaleruca luteola was accidentally introduced into Britain from Bolzano (Italy) in July 1986. Several specimens had accidentally been transported in camping equipment (SMITH 1990). The single specimen recorded by STERRENBURG (1989) for the Netherlands may also have been based on a displaced specimen. This species has been recorded for the Netherlands only from the very south of the country (BEENEN 1998) 4.5. Non-target effects Zygogramma bicolorata is an efficient biocontrol agent that can have significant negative impact on the growth and reproduction of Parthenium hysterophorus (DHILE- EPAN et al. 2000). Zygogramma bicolorata breeds in In- dia on small scale on Xanthium strumarium and feeds on Helianthus annuus. Feeding on H. annuus is re- garded unwanted non-target feeding. Parthenium pol- len, which contains parthenin, when deposited on Heli- anthus annuus, makes the latter attractive to beetles in absence of Parthenium. Continuous feeding on Helian- thus annuus retards ovarian development and thus af- fects fecundity (VIRAKTAMATH et al. 2004). JULIEN (1992) mentioned the case of Chrysolina hy- perici that was introduced in New Zealand in 1947 from England to control Hypericum perforatum but attacked Hypericum androsaemum. Despite some early damage the insect has not persisted on H. androsaemum and has not established. In this case the non-target effect did not persist. Galerucella calmariensis and Galerucella pu- silla have been introduced to North America in 1992 as a biological control agent for Lythrum salicariae and are now established in many US states and Canadian prov- inces. At some sites short-term attack on Rosa multi- flora, Potentilla anserina and Decodon verticillatus has been observed. This “spillover” does not constitute a host shift since the beetles are unable to complete de- velopment on these non-target plants (BLOSSEY et al. 2001). 5. DISCUSSION It is difficult to give an explanation of the observed rela- tive high number of species from the subfamilies Chry- somelinae, Galerucinae, Alticinae, Cassidinae and Hispinae that have been translocated. It cannot be ex- plained from the representation of these subfamilies within Chrysomelidae. Eumolpinae for example is a large subfamily and is represented in our database by only one species. Although JOLIVET (2001) attributed this to the endogenic life of the larvae of Eumolpinae, this can only be part of the explanation. KIMOTO (1988) showed remarkable differences in the geographical rep- resentation of subfamilies in Chrysomelidae. The trans- located species originate from all biogeographical re- gions (Palearctic [47 %], Neotropical [20 %], Nearctic [16 %], Australasian [12 %] and Afrotropical Region [2 %|). The observed representation of subfamilies among the translocated species seems to be a mix of the sub- family representations in these biogeographical regions. For exampled, in the Palearctic Eumolpinae are repre- sented by a low number of species. It is most likely that the number of unintentional trans- locations is highly underestimated. Unintentional trans- locations that did not result in establishment are rarely detected and rarely published. However, unintentional translocations that resulted in establishment and showed negative effects are likely to have been noticed. Such translocations are of importance for our main question, namely the impact of translocations on the biodiversity. Sometimes the reason why an introduced species has not established is clear. Altica carduorum was intro- duced in Canada in 1963 from Switzerland and France to control Cirsium arvense. It did not establish in Al- berta, British Colombia, Nova Scotia or Ontario. Slow development in cool summers exposed larvae to high predation (JULIEN 1992). It was also introduced to Great Britain in 1969 and 1970 from France. Several thou- sands of specimens were released at Silkwood Park, As- cot, Berkshire and at three sites west of Cardiff, Glamorganshire, South Wales (BAKER et al. 1972, COX 2000). They survived the winter in cages only. The cli- mate is too cold and wet to allow survival, except lo- cally (JULIEN 1992). Chrysolina varians was introduced in Australia from England in 1930 to control Hypericum perforatum and did not establish. It is considered to have suffered from heavy predation and unfavourable climate (JULIEN 1992). It was introduced in Canada from Sweden in 1957 to control Hypericum perforatum and did not establish in British Columbia. Release sites were too dry during summer (JULIEN 1992). Adults of the alticinae Disonycha glabrata (F.) have been released to control Amaranthus retroflexus from Massachusetts (U.S.A.) in Red River Valley, North Dakota in 1979 and 1980 but failed to overwinter (JULIEN 1992). Longitar- sus albineus was introduced in Australia in 1979 and 1981 from Greece and France to control Heliotropium europaeum. Establishment failed due to drought, which eliminated the host (JULIEN 1992). Physonota alutacea was introduced in Mauritius from Trinidad in 1947 to control Cordia curassavica. It did not establish due to interference by ants (JULIEN 1992). The observed low percentage of polyphagous species (7 %) that have established after unintentional transloca- tions is not surprising. It seems to be the result of the small proportion of polyphagous Chrysomelidae. Of Chrysomelidae (including Bruchidae) in Central Europe only 19 % are polyphagous, whereas monophagous spe- cies contribute 21 % and oligophagous species 60 % (SCHOLLER 1996). Although we know of some invasive species that cause a lot of harm, for example Leptinotarsa decemlineata and Diabrotica species, these species had effect on plants that were non-native culture plants. The spread of 184 Bonner zoologische Beiträge 54 (2005) the potato over the world was followed by the spread of the insect species associated to it. The same holds for Diabrotica virgifera and the increasing culture of corn (Zea mais) in Europe. The associated insects will follow the crop when environmental conditions are favourable for the leaf beetle species. Dicladispa armigera (Oliv- ier) (Hispinae) was recorded as a pest of rice in Bengal and Bangladesh in 1906. It is the most important hispid pest of rice in tropical Asia, and known from Bangla- desh, Myanmar, India, Nepal and China, frequently causing extensive losses of rice crops. Although D. ar- migera is widely distributed throughout India, and fre- quently causes considerable damage to rice crops, it is only considered as a major pest of rice in Andhra Pradesh, Assam, Bihar, Madhya Pradesh, Manipur, Kerala, Orissa, Indian Punjab, Uttar Pradesh and West Bengal. It is only a rice pest in the southern provinces of China and is present in western Iran and Sumatra, Indo- nesia and Taiwan (CAB-CPC 2003). Introduced leaf beetle species affecting native plants seem to be rare. The introduced Plagiodera versicolora, Agelastica alni and Xanthogaleruca luteola are known to damage ornamental trees, and they could cause dam- age to indigenous woody trees such as Salix, Populus, Alnus or Ulmus in North America. However this has never been reported. At present, Epithrix hirtipennis na- tive to North and Central America occurs in Italy on several Solanaceae, also on the wild and indigenous species (Maurizio Biondi, L’Aquila-Coppito, pers. comm. 2004) but there 1s no indication that this causes a threat to biodiversity in Italy. It seems that negative effects of introduced species on nature are barely recognised. However, this does not mean that they may never occur (DOWNIE 2001). SILFVERBERG (1995) listed the exotic beetle species that have colonized Finland and established themselves out- doors. No leaf beetle was included among the 14 spe- cies. This might be due to the climatic conditions in Finland. However, among the more than 200 invasive species listed in the internet invasive species database no Chrysomelidae have been listed (IUCN/SSC 2004). This database focuses on invasive species that are be- lieved to be a threat to biodiversity. From the absence of leaf beetles we might conclude that translocated leaf beetles do not threaten biodiversity yet. This is interest- ing because there are very alarming situations of leaf beetles attacking cultivated plants (Diabrotica virgifera, Leptinotarsa decemlineata) or species that have been ef- fectively used as control agent against introduced weeds (Chrysolina hyperici and C. quadrigemina). In these circumstances, where alien leaf beetle species eradicate a population of a host plant, the host plants grow vigor- ously because of two different situations: one is the cul- tivated situation in which the environmental (fertilisa- tion, hydration) and ecological conditions (absence of competition, diseases) favour the host plant very much. A monoculture of good growing plants is the result. The other is a situation in which an alien plant is invading a new environment, mostly in absence of natural enemies or diseases and mostly without competitors. Under more natural conditions the herbivorous beetles find less fa- vourable conditions. When a leaf beetle enters an eco- system it faces a complex of interrelations in which it first has to establish. It has been shown (KOVALEV 2004) in model studies that this process is more likely to occur in situations where host plants are evenly spread. Under natural conditions this situation rarely occurs. ELTON (2000) promoted in his famous work on ecologi- cal invasions, ecological stability to resist invading spe- cies. The threats caused by introduced species on native spe- cies are larger than can be concluded from this study on Chrysomelidae. Introduced plant species, introduced carnivorous species, introduced plant or animal dis- eases, may all have their effect on native leaf beetles. Many cases have been listed in which non-native insects (predators or parasitoids) have been released to control non-native pest species (for examples release of Oo- myzus gallerucae Fonscolombe (Hymenoptera, Eulo- phidae) in the United States for the control of Xan- thogaleruca luteola (PUTTLER & BAILEY 2003). The damage of intentionally introduced species to non- target, native species from these biological controls are rarely documented. No negative effects on biodiversity of translocated Chrysomelidae have, at present, been reported. The suc- cessful control of imported weeds by introduced Chry- somelidae can be seen as a positive effect to biodiver- sity since some of these weeds prevent natural processes in ecosystems. It is generally accepted that only a mi- nority of alien species become abundant. WILLIAMSON & FITTER (1996) concluded that as many as 80 — 90 % of the established non-indigenous species may actually have minimal detectable effects. Apart from that, intro- duction of non-indigenous species as a biological con- trol agent is preceded by intensive research to predict the effect of these introductions. BYERS et al. (2002), however, indicated that although there has been a lot of research, we still have little information on the effects on the communities and species we are attempting to protect. They propose key research questions to effec- tively prioritise and manage non-indigenous species. Although there are no cases known in which biodiver- sity is threatened by translocated Chrysomelidae this is no guarantee that it will not happen. Accidental translo- cations must be avoided as much as possible. No inten- tional translocations should be considered until the fac- tors, which limit its distribution and abundance in its native range, have been understood. The approach sug- gested by the IUCN (1987, 2000) will minimise the risks. The Convention on Biological Diversity (Rio de Janeiro 1992) called for prevention of the introduction of, con- trol or eradication of those alien species, which threaten ecosystems, habitats or species. However it proved to be Ron BEENEN: Translocation in Leaf Beetles 185 difficult to discriminate between alien species that are likely to be invasive and those that are not. Because post entry control of invasive alien species is much more costly than prevention of invasions, MACK et al. (2000) suggested that national and international quarantine laws should be altered by adopting a “guilty until proven in- nocent” approach. A rarely studied risk of alien species is the role of evo- lutionary processes. KOVALEV (2002) showed changes in the introduced Zygogramma suturalis which devel- oped flight ability and morphological differences within five generations. The differences where large enough to attribute the Palearctic population to a new subspecies: Z. suturalis volutes KOVALEV. About the risk of hy- bridisation between native leaf beetle species and alien species nothing is clear. Hybridisations between native and alien plants species or between genetically different populations of alien plants are believed to promote rapid evolution and further invasion as has been observed in Spartina anglica (polyploid hybrid of S. alterniflora from eastern America and the European S. maritima) and Rhododendron ponticum (introduced from several different Iberian populations) (PETIT 2004). Apart from natural processes as the post glacial spread of many species (including spread across barriers as the Alps and Pyrenees in Europe), human-induced translo- cations may have started as soon as man appeared. In Europe the invasion of alien species started in ancient times. The original landscape most probably was a mo- saic of forest and open areas. When man arrived large areas were cleared for agriculture. Plant and animal spe- cies from deserts and steppes invaded the European re- gion when new, cultivated, habitats appeared. Already at the time of Columbus the coastland of Western Europe was almost devoid of forests and transformed into an artificial steppe inhabited by man’s constant fol- lowers among plants and animals, invited or self-invited (LINDROTH 1957). Part of theses species have simply expanded their range but it is very likely that part of them were actively transported with seeds or with the transport of livestock (both plants and animals). It is very likely that leaf beetle species associated with crops or the herbs that grow between them have taken the same route as herbs associated with cereals. They are supposed to have arrived in Europe from the Near East (PINHASI ef al. 2005). This process is still going on. HAMMOND (1974) assumed that some of the relative re- cent immigrants to the British Isles may represent de- layed movements of the same kind. An example of such a species combination may be Buglossoides (= Lithos- permum) arvensis and Longitarsus fuscoaeneaus RED- TENBACHER. Both species are nowadays indigenous in Europe. It is believed that B. arvensis originated in southwest-Asia and spreaded with agriculture to large parts of the temperate parts of the Northern Hemisphere. In The Netherlands it already occurred during the Ro- man times (WEEDA er al. 1988). It is likely that L. fus- coaeneus spreaded with this herb. FRITZ-KÖHLER (1996) and FRITZLAR (1998) estimate that about 5% of the leaf beetles of the German states Rhineland and Thúringia came along with introduced plants (neophytes and archeophytes). In America conditions were different when transatlantic trade started. In northeastern North America the tribes were more or less migratory and had hardly progressed beyond the Neolithic stage. Therefore in North Amer- ica, the chance of native steppe plants invading perma- nent arable land and transforming into constant weeds was considerably less than in the old world. When the first Europeans arrived and permanent settlements were founded upon which agriculture started, there were few indigenous plants present able to intrude as weeds. This gave free entrance to the corresponding floral elements from Europe. Species introduced from Europe into North America are about ten times as numerous as those transported in the opposite direction (LINDROTH 1957). This is partly explained by the peculiar character of bal- last traffic in the 19th century. The small sailing vessels used for commercial traffic needed cargo or ballast to be sailed efficiently. On their way to Northern American ports they often were short in tonnage. The bottom of the lower holds were then filled with material available at the shore of departure: gravel, rocks or even moist sand. On arrival at their destination the ballast was dumped on the coast, including the plants and animals that came along with this material. Examples of leaf beetles that have been transported in this way are Cassida flaveola, Chrysolina staphylea and Gastro- physa polygoni (LINDROTH 1957). Not all translocations have to be human-induced. CLARKE & ZALUCKI (2004) suggested, based on histori- cal information, that a substantial population of the monarch butterfly (Danaus plexippus (L.), Danaidae) was carried to Australia on winds associated with cy- clones that hit the Queensland coast in early 1870. The leaf beetle Chaetocnema confinis, originally a North American species, has spread to tropical America, Af- rica, Asia and the Pacific, accidentally introduced in Hawaii and established, it is reported to be transported by a hurricane (JOLIVET 2001). Stegnaspa trimeni is common on Tristan da Cunha and originated from Southern Africa, most probably it was accidentally in- troduced or transported eolic (hurricane ?) (JOLIVET 1998). Epithrix hirtipennis from North and Central America was recorded in 1984 from the Azores and It- aly (DOBERL 1994a). Subsequently it spreaded to Greece (1988) and Turkey (1993). Although introduc- tion with cultivated plants is possible, spreading with passatwinds is also suggested as a possibility (DOBERL 1994a). Acknowledgments. S. M. Clark, M. Biondi, D. Erber (+) and M. A. Hielkema kindly supported this study either be supplying information or supplying specimens from which conclusions were drawn. Theresa Canning (CAB Interna- tional) kindly offered a free trial of the internet version of the Crop Protection Compendium. P. Jolivet stimulated this 186 Bonner zoologische Beitráge 54 (2005) study by many references to useful publications. D. G. Furth, T. Wagner and an anonymous reviewer made valu- able comments on a previous version of this article. Participating in the 6th International Symposium on the Chrysomelidae was possible with a grant from the “Uyttenboogaart-Eliasen Stichting tot bevordering van de entomologische wetenschap”. REFERENCES ALLARD, M. E. 1890. Troisieme note sur les Galérucides. Compte rendu des séances de la Société ento- mologique de Belgique 4(6): 80-94. BAKER, C. R. B., BLACKMAN, R. L. & CLARIDGE, M. F. 1972. Studies on Haltica carduorum Guerin and aliien beetle released in Britain as a contribution to the bio- logical control of creeping thistle, Cirsium arvense (L.) Scop. Journal of Applied Ecology 9: 819-830. BEENEN, R. 1998. Patterns in the distribution of Galeruci- nae in the Netherlands. Museo regionale di Scienze Naturali — Torino 1998: 7-16. BLOSSEY, B., CASSAGRANDE, R., TEWKSBURY, L., LANDIS, D. A., WIEDENMANN, R. N. 8 ELLIS, D. R. 2001. Non- target feeding of leaf beetles introduced to control Purple Loosestrife (Lythrum salicaria L.). Natural Ar- eas Journal 21: 368-377. BYERS, J. E., REICHARD, S., RANDALL, J. M., PARKER, I. M., SMITH, C. S., LONSDALE, W. M., ATKINSON, I. A. E., SEASTEDT, T. R., WILLIAMSON, M., CHORNESKY, E. & HAYES, D. (2002). Directing research to reduce the impacts of nonindignous species. Conservation Bi- ology 16: 630-640. CAB-CPC. Interactive Crop Protection Compendium [http://www.cabi.org/compendia/cpe/index.htm; visited August 2003]. CLARKE, A. R. & ZALUCKI, M. P. 2004. Monarchs in Aus- tralia: on the winds of a storm? Biological Invasions 6: 123-127. COLAUTTI, R. I. & MACISAAC, H. J. 2004. A neutral term to define ‘invasive’ species. Diversity and Distribu- tions 10: 135-141. Cox, M. L. 2000. Progress report on the Bruchi- dae/Chrysomelidae Recording Scheme. The Coleop- terist 9: 65-74, CsIKI, E. 1953. Uber neue und bekannte Coleopetren aus Ungarn und den angrenzenden Ländern. Annales his- torico — naturales Musei nationalis hungarici (N. ser.) 3: 115-135. CUDA, J. P., PARKER, P. E., COON, B. R., VASQUEZ, F. E. & HARRISON, J. M. 2002. Evaluation of exotic So- lanum spp. in Florida as host plants for the leaf beetles Leptinotarsa defecta and L. texana. Florida Entomolo- gist 85: 599-610. DHILEEPAN, K., SETTER, S. D., & MCFAYDEN, R. E. 2000. Response of the weed Parthenium hysterophorus (As- teraceae) to defoliation by the introduced biocontrol agent Zygogramma bicolorata (Coleoptera: Chry- somelidae). Biological Control 19: 9-16. DILLON, E. S. & DILLON, L. S. 1972. A manual of common beetles of Eastern North America. Corrected and re- published. Dover, New York. Volume II. Pp. 434-894. DOBERL, M. 1994a. Auffálige Ausbreitung einiger Al- ticinen-Arten in Westeuropa. Verhandlungen des 14. Internationalen Symposiums fiir Entomofaunistik in Mitteleuropa, SIEEC, in Miinchen: 276-281. DOBERL, M. 1994b. Bemerkenswerte Alticinenfunde aus Westeuropa. Entomologische Nachrichten und Be- richte 38: 179-182. DOWNIE, R. (ed.) 2001. Alien species friends or foes? The Glasgow Naturalist 23, Supplement: 1-113. ELTON, C. S. 2000. The ecology of invasions by animals and plants. With a new forword by D. Simberloff. University of Chiacago Press. i-xiv, 1-181. Originally published in 1958 by Methuen, London. FIELDS, P. G., ARNASON, J. T. & PHILOGENE, B. J. R. 1988. Distribution of Chrysolina spp. in Eastern Ontario, 18 years after their initial release. The Canadian Ento- mologist 120: 937-938. FRASER, B. & EMBERSON, R. 1987. Rediscovery of Chry- solina quadrigemina (Suffrian) in New Zealand. New Zealand Entomologist 9: 57-59, FRITZ-KOHLER, W. 1996. Blatt- und Rússelkáfer an Ack- erunkráutern. Okologie und Biogeographie in Mittel- europa und Untersuchungen an ungespritzten Acker- randstreifen. Agrarókologie 19: 1-138. FRITZLAR, F. 1998. Neozoen unter den Blattkäfern Thürin- gens. Mitteilungen des Thúringer Entomologenver- bandes 5: 61-63. GONZÁLES-MEGÍAS, A. & GOMEZ, J. M. 2003. Conse- quences of removing a keystone herbivore for the abundance and diversity of arthropods associated with a cruciferous shrub. Ecological Entomology 28: 299- 308. GRAY, M. E. & STEFFEY, K. L. 1999. Western Corn Root- worm adapt to crop rotation. Agrichemical and Envi- ronmental News 156: 12-14. HAMMOND, P. M. 1974. Changes in the British Coleopter- ous Fauna. Pp. 323-369 in: HAWKSWORTH, D. L. (ed.) The changing Flora and Fauna of Britain. HAWKES, R. B., ANDRES, L. A. & ANDERSON, W. H. 1967. Release and progress of an introduced flea beetle, Agasicles n. sp., to control alligatorweed. Journal of Economic Entomology 60: 1476-1477. HIGHT, S. D., BLOSSEY, B., LAING, J. & DECLERCK- FLOATE, R. 1995. Establishment of insect biological control agents from Europe agianst Lythrum salicariae in North America. Environmental Entomology 24: 967-977. HOFFMANN, J. H., MORAN, V. C. & IMPSON, F. A. C. 1998. Promising results from the biological control pro- gramme against a solanaceous weed (Solanum elaeag- nifolium). Agriculture, Ecosystems and Environment 70: 145-150. IUCN 1987. IUCN Position statement on translocation of living organisms: introductions, reintroductions and re-stocking. Prepared by the Species Survival Com- mission in collaboration with the Commission on Ecology, and the Commission on Environmental Pol- icy, Law and Administration, 13 pp. IUCN 2000. IUCN guidelines for the prevention of biodi- versity loss caused by alien invasive species. Prepared by the Species Survival Commission Invasive Species Specialist Group, 22 pp. IUCN/SSC. Global invasive species database. Invasive Species Specialist Group. [http://www.issg.org/ database/species; visited August 2004]. JOLIVET, P. 1967. Notes systematiques et ecologiques sur les Chrysomelides Marocains (2° note). Bulletin de la Société des Sciences Naturelles et Physiques du Maroc 46:305-393. JOLIVET, P. 1998. Les nouveaux envahisseurs ou les Chry- somelides voyageurs. L’Entomologiste 54: 33-44. JOLIVET, P. 2001. Vers la mondialisation des Chry- somelides? L’ Entomologiste 57: 123-141. JOLIVET, P. & HAWKESWOOD, T. J. 1995. Host-plants of Chrysomelidae of the world. An essay about the rela- Ron BEENEN: Translocation in Leaf Beetles 187 tionships between the leaf-beetles and their food- plants. Backhuys Publishers, Leiden. 281 pp. JOLIVET, P. & VERMA, K. K. 2002. Biology of leaf beetles. Intercept, Andover. i-xiv, 1-332. JULIEN, M. H. 1992. Biological Control of Weeds. A World Catalogue of Agents and their Target Weeds. Third Edition. CAB International in association with ACIAR. 186 pp. KIMOTO, S., 1988. Zoogeography of the Chrysomelidae. Pp. 107-114 in: JOLIVET, P., PETITPIERRE, E. & HSIAO, T. H. (eds.). Biology of Chrysomelidae. KOVALEV, O. V. 2002. The formation of the solitone-like waves under the invasions of the living organisms and in the evolution of the biosphere. Evolutsionnaia Biologiia 2: 65-81 [in Russian]. KOVALEV, O. V. 2004. The solitary population wave, a physical phenomenon accompanying the introduction of a chrysomelid. Pp. 591-601 in: JOLIVET, P., SANTI- AGO-BIAY, J. A. & SCHMITT, M. (eds.). New Devel- opments in the Biology of Chrysomelidae. KRAFSUR, E. S. & NARIBOLI, P. 1995. Elm leaf beetles have greatly reduced levels of gene diversity. Bio- chemical Genetics 33: 91-95. LESAGE, L. 1988. Notes on European Longitarsus species introduced in North America. The Canadian Ento- mologist 120: 1133-1145. LESAGE, L. 1990. Chaetocnema concinna (Marsham, 1802), a European flea beetle introduced in North America. The Canadian Entomologist 122: 647-650. LINDROTH, C. H. 1957. The Faunal Connections between Europe and North America. Almqvist & Wiksell, Stockholm / John Wiley & Sons, Inc., New York. 344 pp. MACK, R. N., SIMBERLOFF, D., LONSDALE, W. M., EVANS, H., CLOUT, M. & BAZZAZ, A. A. 2000. Biotic inva- sions: causes, epidemiology, global consequences, and control. Ecological Applications 10: 689-710. MOHAMEDSAID, M. S. 2001. Implications for mislabeling specimen. Serangga 6: 227-231. Morimoto, N & KIRITANI, K. 1995. Fauna of exotic in- sects in Japan. Bulletin of the National Institute of Agro- environmental sciences 12: 87-120. Opum, E. P., 1971. Fundamentals of ecology. Third edi- tion. W. B. Saunders Company. Philadelphia, London, Toronto. 574 pp. PEMBERTON, R. W. & PRATT, P. D. 2002. Skunk Vine. Pp. 343-351 in: VAN DRIESSCHE et al. Biological control of invasive plants in the Eastern Unites States. USDA Forest Service Publication FHTET-2002-04. PETIT, R. J. 2004. Biological invasions at the gene level. Diversity and Distributions 10: 159-165. Pic, M. 1931. Nouveautés diverses. Mélanges exotico- entomologiques 57: 1-36. PINHASI, R., FORT, J. & AMMERMAN, A. J. 2005. Tracing the origin and spread of agriculture in Europe. PLoS Biol 3: e410. PUTTLER, B. & BAILEY, W. C. 2003. Establishment of Oo- myzus gallerucae, an egg parasite of the elam leaf bee- tle, in Missouri and adjacent states. Biological Control 27: 20-24. SANNINO, L., BALBIANI, A. & BIONDI, M. 1985. Epithrix hirtipennis (Melsheimer, 1847): considerazioni tasso- nomiche, ecologiche ed etologiche. Atti XIV Congr. naz. ital. Ent., Palermo. Erice, Bagheria: 285-292. SCHOLLER, M. 1996. Ökologie mitteleuropäischer Blattkäfer, Samenkáfer und Breitrüssler. Die Käfer von Vorarlberg und Liechtenstein 11: 1-65. SCHROEDER, D. & GASSMANN, A. 1997The search for in- sect biocontrol agents to control a tough and variable target weed. Leafy Spurge News 19: 1-2. SEENO, T. N. & WILCOX, J. A. 1982. Leaf Beetle Genera. Entomography 1: 1-221. SHELDON, S. P. & CREED, R. P. 1995. Use of a native in- sect as a biological control for an introduced weed. Ecological Applications 5: 1122-1132. SILFVERBERG, H. 1978. The identity of Aulacophora pan- nonica Csiki. Folia Entomologica Hungarica 31: 205- 206. SILFVERBERG, H. 1995. Immigration and range expansion in Finnish insects. Entomologica Fennica 6: 163-167. SIMBERLOFF, D. & GIBBONS, L. 2004. Now you see them, now you don’t! — population crashes of established in- troduced species. Biological Invasions 6: 161-172. SMITH, K. G. V. 1990. Pyrrhalta luteola accidentally in- troduced into Britain. Entomologist’s Monthly Maga- zine 126: 190. SPRING, A. & Kok, L. T. 1999. Winter survival of Cassida rubiginosa, a biological control agent of Canada This- tle. Journal of Entomological Science 34: 489-493, STEWART, A. J.A. 1996. Interspecific competition rein- stated as an important force structuring insect herbi- vore communities. Trend in Ecology and Evolution 11: 233-234. STERRENBURG, F. C. F. 1989. Ergánzungen zur Kaferfauna der Niederlande. Entomologische Blatter 85: 81-92. SUZUKI, K. & NAKAMURA, H. 1999. Rapid invasion of Ophraella comuna LeSage, 1986 to Fukui and Ishi- kawa Prefectures, Central Honshu, Japan. Entomo- logical Journal of Fukui 25: 5-6. UNION OF CONCERNED SCIENTISTS 2001. The Science of invasive species. An information update by the Union of Concerned Scientists. 12 pp. UNIVERSITY OF RHODE ISLAND 2002. Lily Leaf Beetle. Uni- versity of Rhodes Island GreenShare Factsheets, 3 pp. VELU, S. 2001. Local Beetles’ battles. On: Asian’s review of biodiversity & environmental conservation website [www.arbec.com.my; visited April, 15, 2004]. VIRAKTAMATH, C. A., BHUMANNAVAR, B. S. & PATEL, V. N. 2004. Biology and ecology of Zygogramma bicol- orata Pallister, 1953. Pp. 767-777 in: JOLIVET, P., SANTIAGO-BLAY, J. A. & SCHMITT, M. (eds.). Devel- opments in the Biology of Chrysomelidae. WANG, C. & CHIANG, M. 1998. New record of a fastidious chrysomelid, Ophraella communa LeSage in Taiwan. Plant Protection Bulletin 40: 185-188. WATANABE, M. & HIRAI, Y. 2004. Host-use pattern of the ragweed beetle Ophraella communa LeSage for over- wintering and reproduction in Tsukuba. Applied En- tomology and Zoology 39: 249-254. WEEDA, E. J., WESTRA, R., WESTRA, Ch. & WESTRA, T. 1988. Nederlandse oecologische Flora. Wilde planten en hun relaties. 3: 1032. IVN, VARA, VEWIN. WILLIAMSON, M. H. & FITTER, A. 1996. The characters of successful invaders. Biological Conservation 78: 163- 170. ZHANG, L. YANG, X., LI, W, Cul, J. 2005. A new record of Ophraella communa of mainland China. Chinese Bul- letin of Entomology 42: 227-228 [in Chinese with English abstract]. Author’s address: Ron BEENEN, Martinus Nijhoffhove 51, NL — 3437 ZP Nieuwegein, The Netherlands. E-mail: r.beenen@wxs.nl Bonner zoologische Beitráge 54 (2005) 188 L661 NNVIASSVO) pnsa 7 'pip3.1aopnasd I 29 UAGAOUHIS +7661 Nalin x piqaoydng 7861 al (yoy) I 7661 Narıny sorssiuw9dAs viqsoydng Z861 A epeue) odoing ILIV oDISSIAOAA) puoyiydy L661 NNVWSSVD 29 (pruyosyna) I UACAOUHS : [007 LAATIOL pjnso piqsoydng 2661 el Vsn adoing ILIV sıjpunuopgp puoyiydy I Z66I NATIONS. 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epeue) pue ys. vsn OXOW OMXIA] INIIBI|P (A]e1]) ouezjog é adoing eleysny BIA BOTY BISOD AYHO AYHO AYHO DNIZ HTVO SNIOLIQR sıpeinyns eure 13ogAz sıadoy DIANASIP DULUDABOBAZ 19151118 d eJe10]091q BUILUVIBOSATZ ¡ UONEULIFUOI spoou uonesznusp]) URLYFNS s140]]93n98 pA0oydosnoz COTA) pjoaın] PINAD/PSOYIUPDX sindeyo pywaurig 1e9u ‘ds p30]d0.1/) I Bonner zoologische Beiträge Band 54 (2005) | Heft 4 | Seiten 201-208 Bonn, Oktober 2006 Functional Morphology of Copulation in Chrysomelidae-Criocerinae and Bruchidae (Insecta: Coleoptera)' / Susanne DUNGELHOEF & Michael SCHMITT Zoologisches Forschungsmuseum Alexander Koenig, Bonn, Germany Abstract. Until now, the genitalia of beetles have been investigated mainly under taxonomic aspects while there exist few studies on the functional morphology of genital structures. Many questions concerning the position of male genitalia inside the female during copulation, the functional role of parameres and endophallus, the mechanical coupling of the mates as well as sperm transfer remain unsolved for most species of Coleoptera. This article presents results gained from investigations of pairs of Acanthoscelides obtectus (Bruchidae), Oulema melanopus, Oulema duftschmidi and Lilioceris lilii (Criocerinae) fixed in copula. During copulation parameres remain outside the female in Acanthoscelides. In Crio- cerinae, the flagellum is placed near the entrance of the spermathecal duct. Campaniform sensilla were found on the sur- face of the endophallus in Lilioceris. Key words. endophallus, flagellum, aedeagus, sperm-transfer, parameres, bursa copulatrix, Acanthoscelides obtectus, Lilioceris lilii, Oulema duftschmidi, Oulema melanopus 1. INTRODUCTION Most evolutionary variations can be interpreted as a means to increase fitness. Anything connected with re- production is of central importance for this increase. The functional morphology of copulation, however, is unknown for most species of beetles. There exist studies on the functional morphology of genitalia in insects in general (HEBERDEY 1928, 1931) and in several beetle species. NYHOLM (1969) for example, analysed the structure and function of the copulatory organs of Cy- phon species (Coleoptera: Scirtidae). EBERHARD (1993) studied courtship and genital mechanics of three species of Macrodactylus (Coleoptera: Scarabaeidae). Sper- mathecal morphology and sperm transfer of the staphylinid beetle, Aleochara curtula (Coleoptera: Staphylinidae), are well understood (GACK & PESCHKE 1994; FORSTER et al. 1998). HAUBRUGE et al. 1999 found that in Tribolium castaneum (Coleoptera: Tene- brionidae) the male removes the sperm of previous males from the female tract by means of its median lobe. In Cicindela (Coleoptera: Cicindelidae) as well, the males seem to clear the spermatheca and the sper- mathecal duct with its flagellum before they place the spermatophore (FREITAG et al. 2001). Since all these groups are not closely related to the Chrysomelidae, the results may not be applicable to our group. ' Paper presented to the 6" International Symposium on the Chry- somelidae, Bonn, Germany, May 7, 2004. One of the first studies on the functional morphology of genitalia in Chrysomelidae is that of HARNISCH (1915) who described the copulatory apparatus of Chrysomela populi, Clytra quadripunctata and Plateumaris sericea. RODRIGUEZ et al. (2004) found that females of Chely- morpha alternans (Chrysomelidae: Cassidinae) prefer males with a longer flagellum. This result confirms the eryptic female choice hypothesis (THORNHILL 1983; EBERHARD 1985, 1997). CRUDGINGTON & SIVA JOTHY (2000) observed that the males of Callosobruchus maculatus (Coleoptera: Bruchidae) penetrate the wall of the female genital tract to prevent re-mating of the fe- male. (The “Bruchidae” are, in terms of phylogeny, simply a subtaxon of the Chrysomelidae, as discussed by SCHMITT 1996). The latter works show impressively that studies on func- tional morphology of genitalia can provide new insights on sexual selection and sexual conflict. Our aim is to investigate the functional role of genital structures in all subfamilies of Chrysomelidae to learn more about these phenomena in the whole family. Hopefully, the results will lead to new ideas on speciation processes in this large beetle family. In this paper we present some first results gained from three species. 202 Bonner zoologische Beitráge 54 (2005) 2. MATERIALS AND METHODS 2.1. Beetles Bruchidae Acanthoscelides obtectus (Say, 1831): Breeding of these beetles is very uncomplicated, in which a preserving glass was used, filled with beans and with a lid consisting of a membrane permeable to air. We received seed-beetles from the Institute for Plant Diseases, Bonn and used the bush beans Montano or Maja as substrate. Every three months we removed dead beetles and replaced a portion of the old beans with fresh ones. Chrysomelidae-Criocerinae Lilioceris lilii (Scopoh, 1763): These beetles were caught in private gardens around Bonn and Kevelaer (lower Rhine province near the Dutch border) in April and May 2003. We kept them in Drosophila-tubes, where they fed on leaves of Lilium sp. To achieve an improved climate, a layer of 1-2 cm of gypsum was poured into the bottom of the rearing tubes. Oulema melanopus (Linnaeus, 1758)/ Oulema duft- chmidi (Redtenbacher, 1874): Specimens of Oulema came from a wheat field near Bonn in May 2003. They were kept in Drosophila- glasses like L. /ilii and fed wheat plants from the native field. As the two species are only distinguishable by their genitalia and lack external differences, the first au- thor caught and worked with both forms. The results for each species were pooled and therefore are presented as a whole and not seperately. 2.2. Behaviour To initiate and observe copulations we put pairs or small groups of beetles together in a tube. The behaviour was visible with the naked eye. For some details we used a dissecting microscope. Ten copulations of different indi- viduals of A. obtectus were watched closely. In Crioceri- nae we observed fewer copulations accurately from be- ginning to end (n<3). Duration of copulation could be estimated by keeping an eye on the tubes while occupied with work elsewhere. We define copulation time as the time during which the aedeagus is inserted in the female. 2.3. Fixation in copula To fix the copulating beetles we used chloroethyl spray (Chloraethyl “Dr. Hennig”, Dr. Georg Friedrich Hennig, Chemische Fabrik Walldorf GmbH, D-69190 Walldorf, Germany). Shock freezing by this spray works well, but sometimes the, mating pairs separated partly or com- pletely by the pressure of spraying. In these cases, the mates had not been coupled sufficiently. We fixed 8 — 12 pairs of each species. Frozen beetles were dissected or stored in 70% ethanol at —12 °C for three weeks. 2.4. Histology After three weeks the entire beetles or their abdomina were transferred from 70% to 100% ethanol (30 min each in 80%, 90%, 95%, 100% ethanol). Each was im- mersed in a second soaking in 100% ethanol overnight. Resin was prepared with the “Low Viscositiy” set from Electron Microscopy Sciences (Hatfield, Pennsylvania). Objects were infiltrated by a further series (at intervals of 30 min aceton : resin 1:1, aceton : resin 1:3 and 60 min pure resin) and degased in an exsiccator. The Resin was put in silicon moulds (6x12x5 mm or 6x14x4 mm), the objects added and adjusted. The resin polymerised over night at 69 °C. The embedded objects (1-3 in copula fixed pairs of each species) were cut with a microtome in slices of 1.0 um or 1.5 um. Every fifth section was put on a slide, heat- dried and dyed with Richardson-dilution (BOCK 1989). To study the slices we used a Nikon Eclipse E600. 2.5. Scanning electron microscopy We used the SEM to study in detail the shape and sur- face of the endophallus. Three exemplars, one of each species, were prepared from in copula-fixed pairs. Therefore, the endophallus was everted in an authentic way in all preparations. Objects were transferred from 70% to 100% ethanol and dried using HMDS (Hexame- thyldisilazane) (BOCK 1987). After fixing them on a stub they were coated with a 35 nm gold-layer using a HUMMER VII sputter (ANATECH LTD). We used an HITACHI S-2460 N SEM. 3. RESULTS 3.1. Acanthoscelides obtectus Pairs of A. obtectus mate for about ten minutes. After this time, the male often leans backwards until it is al- most lying on its back, after which it separates from the female. Females do not seem to be willing to remate di- rectly after copulation but require a refractory period. It is striking that fixed pairs separate very easily during dissection. The linking between the dead male and fe- male is much less strong than in Criocerinae. As shown in Figure 1, the parameres in A. obtectus re- main outside the female and lay on the last female ster- nite during copulation while the median lobe is inserted. The photo is taken from a pair fixed in copula. One of us (SD) could observe this also while watching live in- Susanne DUNGELHOEF & Michael SCHMITT: Copulation in Chrysomelidae and Bruchidae 203 dividuals of A. obtectus through a dissecting micro- scope: The male moves the parameres up and down on the last female sternite during copulation and pushes only the tip of the median lobe into the female genital opening. Before inserting the median lobe, the male strikes the female genital opening with the setae of his parameres by moving them up and down. In Figure 2, the considerable size of the everted endo- phallus is visible. It is formed like a balloon and fills the whole bursa copulatrix during copulation. The close-up view (Fig. 3) shows that the parameres are nearly flush with the median lobe. That means that even during copulation, the median lobe and tegmen are hardly moved against each other. The surface of the endophallus bears teeth-like struc- tures on its central region that are obviously sclerotised and appear quite sharp (Fig. 4). These teeth are absent on the apical region; in the basal region they are longer, less sharp structures. 3.2. Oulema melanopus/duftschmidi In this species, pairing lasts longer than in 4. obtectus. The males insert their median lobes for about twenty minutes or even longer. After copulation, males con- tinue to sit on the females’ elytra, guarding their mates. Females try to get rid of them by kicking. Both sexes may remate directly after copulation. The endophallus of O. duftschmidi consists of a mem- branous and a sclerotised part, the so-called flagellum (Fig. 5). The flagellum is tubular, at least apically (Fig. 6). During copulation, the median lobe is inserted com- pletely in O. melanopus. The endophallus fills the whole bursa copulatrix (Fig. 7). A close-up view (Fig. 8) shows how well the walls of bursa and endophallus match. Every bulge of the bursa is filled by the endo- phallus. Figure 8 shows clearly that not only the distal part of the flagellum is tubular but also the whole struc- ture is tubular. An analysis of the neighbouring sections revealed that the flagellum ends at the entrance of the spermathecal duct. The surface of the endophallus of O. dufischmidi bears comb-like structures (Fig. 9) or something similar. They are not sclerotised, unlike the “teeth” of A. obtectus. The structures are directed towards the median lobe. 3.3. Lilioceris lilii As in O. melanopus, copulation in L. /ilii takes more than twenty minutes. The male clings to the female’s back for several hours while the male organ is intromit- ted. The everted endophallus of £. /i/ii is about three times the diameter of the median lobe (Fig. 10). This fact ob- viously facilitates a perfect mechanical coupling of Lilioceris-pairs. The fixed pairs could hardly be sepa- rated. As in Oulema, the endophallus is divided into a membranous and sclerotised part. The sclerotised pgrt includes the flagellum as one can see in the SEM photo (Fig. 11). The tip of the flagellum is also tubular (Fig. 12). The sagittal section of Lilioceris (Fig. 13) represents the mates during copulation. Even if one cannot see a connection between the male body and the median lobe in this figure, one can imagine that the median lobe is inserted. The endophallus fills the bursa. The sclerotised part of the endophallus is visible lying close to the basal part of the spermatheca. We can track this on the pre- ceding and following sections. Thus, in L. /ilii as well as in O. melanopus the tip of the flagellum is positioned very close to the entrance of the spermatheca. The surface of the endophallus of £. /i/ii bears the same comb-like structures as O. duftschmidi. But there are additional structures in L. /ilii (Fig. 14), closely resem- bling sensilla campaniformia, receptors responding to pressure. These receptors occur in rows on parts of the surface of the endophallus. After we had found these structures we searched for dendrites in the sections. And indeed, we found structures crossing the wall of the endophallus, which could be interpreted as dendrites. 4. DISCUSSION 4.1. General aspects We focused on three species that were easily available. Furthermore, we decided to work with one species pos- sessing parameres and two species without in order to obtain an idea on the functional role of these structures. We chose closely related species to enable a meaningful comparison. In future studies, the methods will be maintained but have to be expanded. Paraffin sections for example will be necessary to demonstrate sperm by giemsa-staining. In our sections we found structures inside the flagellum of Oulema that seem to be spermatozoa, however we are presently uncertain. Besides that, many more pairs of each species will have to be fixed for a comparison of different states of copulation. This will permit us to re- construct the process of copulation. 4.2. Sperm-transfer We assume that in L. /i/ii and O. melanopus/duftschmidi the flagellum is positioned near the spermatheca to guarantee a directed sperm transfer towards the female sperm-storage. The results show that the flagellum in O. 204 melanopus/duftschmidi as well as in L. lilii is tubular with a distal opening. Therefore, one could imagine that it is involved in sperm transfer. BERTI & RAPILLY (1976) studied the endophallus of L. /i/ii under a taxo- nomic aspect and figured the ejaculatory duct ending in the flagellum tube. That means that in L. /i/ii sperm transfer actually takes place through the tubular flagel- lum. It is self-evident that the same may be the case in O. melanopus/dufischmidi, though we have not found the ejaculatory duct in the sections. The idea of the fla- gellum as a sperm transferring structure is corroborated by the positioning of its tip: the males presumably ad- duct their flagellum close to the basal piece of the sper- matheca to increase the probability that their sperm ar- rives at the entrance. RODRIGUEZ et al. (2004) consider possible that the flagellum of Chelymorpha alternans (Chrysomelidae: Cassidinae) is pushed up into the spermathecal duct to release sperm there. Tactics of males to transport their sperm close to or even into the spermatheca have been investigated in other cases as well, for example in the staphylinid Aleochora curtula (GACK & PESCHKE 1994). It is conceivable that the long and thin flagellum of O. duftschmidi may reach even into the spermathecal duct during copulation. The sper- mathecal duct of this form is longer than in O. melanopus (BERT! 1989). One could suppose that fe- males have in the course of evolution lengthened the distance from the spermatheca to test the males quality. Males responded by extending their flagellum. In Che- lymorpha species (RODRIGUEZ et al. 2004) the length of the flagellum is actually adapted to the length of the spermathecal duct. Females of this species prefer males with a longer flagellum that reaches the spermatheca. The question if L. /ilii and O. melanopus/duftschmidi form spermatophores as well, is unsolved so far as for many other Chrysomelidae. Males of C. alternans transmit a spermatophore in addition to the pretended sperm transfer via the flagellum (RODRIGUEZ et al 2004). In the Criocerinae we have so far not found any trace of a spermatophore in the sections (which does not necessarıly mean that spermathophores do not exist). Males of Acanthoscelides obtectus are able to produce a spermatophore (HUIGNARD 1978, 1983), which we also could identify in the sections. It is an open question if Criocerinae males remove pre- vious males’ sperm by means of their genitalia, as males in many other groups do (HAUBRUGE et al. 1999; FREITAG et al. 2001). It can’t be excluded that the flagel- lum has this additional function. In Acanthoscelides, it is improbable that the males dispose other males’ sperm from the female genital tract because the median lobe is hardly inserted and no structures of the endophallus seem to be able to perform such a function. Bonner zoologische Beitráge 54 (2005) 4.3. Parameres HARNISCH (1915) has already observed that in Plateu- maris sericea (Chrysomelidae: Donaciinae) the pa- rameres remain outside the female genitalia. He sug- gests that they act as a grasping organ to connect the male more intensively to the female during copulation. Our results show that the parameres of 4. obtectus can’t fulfill this function because the lever formed by them is too short to stabilize. Another possible function of the parameres is to position the apical orifice of the median lobe opposite the opening of the female genital tract as KINGSOLVER (1970) suggests. SD observed in live A. obtectus individuals that the male touched the female genital area with the setae of the parameres before in- serting the median lobe. It is hard to decide if this touch- ing serves the male to orientate the position of the me- dian lobe or to stimulate the female. Such kind of paramere-movement has been observed in other groups as well: males of the carabid beetle Pasimachus punctu- latus tap their parameres rhythmically upon the apical edge of the female last abdominal sternite. After a while, the genital orifice of the female cpens and the male inserts his median lobe (ALEXANDER 1959). If these movements actually serve as stimulation, this would support the cryptic female choice hypothesis (EBERHARD 1985). 4.4. Surface of the Endophallus The comb-like structures on the surface of the endophal- lus in £. lilii and O. duftschmidi are directed towards the median lobe. We assume that the functional role of these structures 1s to support the mechanical coupling of the mates. They may make it more difficult to discon- nect the median lobe forcibly out of the bursa or without the agreement of the male, respectively. If the females want to stop copulation and get rid of the aedeagus, they use their legs to kick the male away as is common for many other species of Chrysomelidae as well (DICKIN- SON 1997; JOLIVET 1999; CRUDGINGTON & SIVA-JOTHY 2000). The everted matching endophallus with its mi- crostructures may prevent separation. Another possible function of the ultrastructures of the endophallus may be stimulatory. To verify this idea there shouid be recep- tors on the interior wall of the bursa copulatrix. The other structures on the endophallus of L. /ilii are without much doubt receptors. The existence of such re- ceptors means probably that the males are able to con- trol the eversion and positioning of the endophallus very well. The sensilla campaniformia may sense that correct position inside the bursa to provide the best opportunity for the flagellum to perform direct sperm transfer. Acanthoscelides obtectus males have teeth-like struc- tures on the surface of their endophallus. KINGSOLVER (1970) suggests that these structures or “endophallus Susanne DUNGELHOEF & Michael SCHMITT: Copulation in Chrysomelidae and Bruchidae 205 armature” serve as holding devices during copulation. This seems to be a good idea, especially because the shape of the endophallus of A. obtectus is not suitable to guarantee its foothold inside the female. But in contrast to the little combs in the Criocerinae, the structures in 4. obtectus are sclerotized and may have another function as well. CRUDGINGTON & SIVA JOTHY (2000) found that the armed endophallus of Callosobruchus maculatus penetrates the bursa wall during copulation. This dam- age of the genital tract is costly for the female because it has to repair the wall. The study even points out that copulation frequency has a life-history cost for females: doubly mated females died significantly younger than singly mated females. Hurting the female is thus a strat- egy of males to ensure that females will not remate. Maybe the same is the case in 4. obtectus. As our ob- servations of live individuals showed, females of 4. ob- tectus do not remate after copulation for a fairly long time (unlike females of O. melanopus). To reach clarity, we shall have to analyse the bursa wall after copulation. 4.5. Perspectives for evolutionary biology The results show that the males of A. obtectus on one side and the Criocerinae on the other embark on differ- ent strategies. In Criocerinae, the males try to prevent further copulations of the females by mate guarding. They obviously have no additional strategy because fe- males seem to be willing to remate just after copulation if only the males desist from them. In 4. obtectus, the males do not stay longer on the females than copulation or even the pure transfer of the spermatophore lasts. They do not spend as much time in pairing than the Criocerinae but have a different way to prohibit a re- mate of the females. As HUIGNARD (1983) described, the deposition of the spermatophore in A. obtectus 1s followed “by stimulation of oogenesis and egg-laying, as well as by a temporary inhibition of female receptiv- ity”. He ascertained that male secretions are transferred from the spermatophore into the female haemolymph through the wall of the bursa. It is uncleared if these se- cretions are the main cause of the refractory period of the females or if damage of the bursa wall like in C. maculatus (CRUDGINGTON & SIVA-JOTHY 2000) is an additional male strategy to inhibit a female remating. If there will be no damages in the bursa detectable after copulation, we have to ask for another function of the teeth on the endophallus surface. We then will agree with KINGSOLVER 1970 that they mainly serve as hold- ing devices. Males in Criocerinae obviously achieve the mechanical coupling with their voluminous inflated endophallus and its additional microstructures. So, if the mechanical coupling in Donaciinae is achieved by the parameres as grasping organs (HARNISCH 1915), it would be interest- ing to know if their endophallus is less voluminous and possesses no microstructures. An idea could be that all species without anchoring parameres offer a voluminous endophallus or at least an endophallus with holding mi- crostructures. One could speculate that there exist three types of Chry- somelidae — one without parameres, another with pa- rameres that serve mainly as grasping organs, and a third that uses the parameres for orientation or stimula- tion. Acknowledgements. We thank Dr. Bernhard Huber (Bonn) for help in all histological matters, Lasse Hubweber (Bonn) for stimulating discussions, Karin Ulmen (Bonn) for technical assistance at the SEM, Bradley J. Sinclair (Bonn) for linguistic assistance, Sandra Steiger (Freiburg im Breisgau) for her patience in searching for dendrite- like-structures in the sections and two anonymous referees for valuable input. Bonner zoologische Beitráge 54 (2005) A A nn 4 159] oO oO 2400 pm —4 ciel } Veer \ Susanne DUNGELHOEF & Michael SCHMITT: Copulation in Chrysomelidae and Bruchidae 207 H—— 500 um —— Fig. 1. Ventral view of a copulating pair of Acanthoscelides obtectus, fixed by chloroethyl spray. P: parameres, M: median lobe, ST: last female sternite. Stereomicrograph. — Fig. 2. 4. obtectus. Aedeagus with everted endophallus (E). M: median lobe, P: pa- rameres. Scanning electron micrograph. — Fig. 3. 4. obtectus. Close-up view of the aedeagus. E: endophallus, M: median lobe, P: parameres. Scanning electron micrograph. — Fig. 4. A. obtecuts. Surface of the endophallus. Scanning electron micrograph. — Fig. 5. Aedeagus of Oulema duftschmidi. E: endophallus, F: flagellum, M: median lobe. Scanning electron micrograph. — Fig. 6. O. dufischmidi. Tip of the flagellum. Scanning electron micrograph. — Fig. 7. Sagittal section of O. melanopus in copula. B: bursa copulatrix, E: endophallus, M: median lobe. Micrograph. — Fig. 8. Sagittal section of O. melanopus in copula. B: bursa copula- trix, F: flagellum, M: median lobe. Micrograph. — Fig. 9. O. dufischmidi. Surface of the endophallus. Scanning electron micro- graph. — Fig. 10. Dissected aedeagus of Lilioceris lilii. E: endophallus, M: median lobe. Stereomicrograph. — Fig. 11. L. /ilii. The sclerotised part of the endophallus (E) bears the flagellum (F). M: median lobe. Scanning electron micrograph. — Fig. 12. L. /ilii. Tip of the flagellum. Scanning electron micrograph. — Fig. 13. Sagittal section of £. /i/ii in copula. B: bursa copulatrix, E: endo- phallus, M: median lobe, S: spermatheca. Micrograph. — Fig. 14. Campaniform sensillum on the surface of the endophallus of L .lilii. Scanning electron micrograph. 208 Bonner zoologische Beitráge 54 (2005) REFERENCES ALEXANDER, R. D. 1959. The courtship and copulation of Pasimachus punctulatus Haldeman (Coleoptera: Carabidae). Annals of the Entomological Society of America 52: 485. BERTI, N 1989. Contribution á la Faune de France. L’identite d’Oulema (O.) melanopus (L.)[Col. Chry- somelidae Criocermae]. Bulletin de la Société entomo- logique de France 94(1-2): 47-57. BERTI, N. & RAPILLY, M. 1976. Faune d'iran. Liste d’especes et révision du genre Lilioceris Reitter (Col. Crysomelidae). Annales de la Société entomologique de France 12(1): 31-73. Bock, C. 1987. A quick and simple method for preparing soft insect tisues for scanning electron microscope us- ing Carnoy and Hexamethyldisilazane. Beitrage zur elektronenmikroskopischen Direktabbildung von Ober- flachen 20: 209-214. Bock, P. (ed.) 1989. Benno Romeis mikroskopische Tech- nik. Urban & Schwarzenberg, Miinchen. CRUDGINGTON, H. S. & SIVA-JOTHY, M. T. 2000. Genital damage, kicking and early death. Nature 407: 855-865. DICKINSON, J. L. 1997. Multiple mating, sperm competi- tion, and cryptic female choice in the leaf beetles (Coleoptera: Chrysomelidae). Pp. 164-183 in: CHOE, J. C. & CRESPI, B. J. (eds.) The Evolution of Mating Sys- tems in Insects and Arachnids. Cambridge University Press, Cambridge. EBERHARD, W. G. 1985. Sexual Selection and Animal Genitalia. Harvard University Press, Cambridge. EBERHARD, W. G. 1993. Copulatory courtship and genital mechanics of three species of Macrodactvlus (Coleop- tera Scarabaeidae Melolonthinae). Ethology, Ecology & Evolution 5: 19-63. EBERHARD, W. G. 1997. Sexual selection by cryptic female choice in insects and arachnids. Pp. 32-57 in: CHOE, J. C. & CRESPI, B. J. (eds.) The Evolution of Mating Sys- tems in Insects and Arachnids. Harvard University Press, Cambridge. FORSTER, M., GACK, C. & PESCHKE, K. 1998. Morphology and function of the spermatophore in the rove beetle, Aleochara curtula (Coleoptera: Staphylinidae). Zool- ogy 101: 34-44. FREITAG, R., HARTWICK, A. & SINGH, A. 2001. Flagellar microstructures of male tiger beetles (Coleoptera: Cicindelidae): implications for systematics and func- tional morphology. The Canadian Entomologist 133: 633-641. GACK, C. & PESCHKE, K. 1994. Spermathecal morphology, sperm transfer and a novel mechanism of sperm dis- placement in the rove beetle, Aleochara curtula (Col- eoptera, Staphylinidae). Zoomorphology 114: 227- 237. HARNISCH, W. 1915. Uber den mánnlichen Begattungsap- parat einiger Chrysomeliden. Ein Beitrag zur Phyloge- nie des Copulationsapparates der Káfer. Zeitschrift fúr wissenschaftliche Zoologie 114: 1-94. HAUBRUGE, E., ARNAUD, L., MIGNON, J. & GAGE, M. J. G. 1999. Fertilization by proxy: rival sperm removal and translocation in a beetle. Proceedings of the Royal So- ciety of London, B, Biological Science 266: 1183- 1187. HEBERDEY, R. F. 1928. Ein Beitrag zur Entwicklungsge- schichte des mánnlichen Geschlechtsapparates der Coleopteren. Zeitschrift fiir Morphologie und Okolo- gie der Tiere 10: 533-575. HEBERDEY, R. F. 1931. Zur Entwicklungsgeschichte, ver- gleichenden Anatomie und Physiologie der weiblichen Geschlechtsausfúhrwege der Insekten. Zeitschrift fiir Morphologie und Okologie der Tiere 22: 416-586. HUIGNARD, J. 1978. Tranfert de sécrétions males du sper- matophore vers l'hémolymphe chez Acanthoscelides obtectus Say (Coléoptere Bruchidae). Comptes rendus des seances de l’Acad&mie des Sciences Paris D 287: 1301-1304. HUIGNARD, J. 1983. Transfer and fate of male secretions deposited in the spermatophore of females of Acan- thoscelides obtectus Say (Coleoptera Bruchidae). Journal of Insect Physiology 29(1): 55-63. JOLIVET P. 2000. Courtship and Mating Behaviour Among Leaf-Beetles. Pp. 115-125 in: SOBTI, R. C. & YADAV, J. S. (eds.) Some Aspects on the Insight of Insect Biol- ogy. Narendra Publishing House, Dehli [1999]. KINGSOLVER, J. M. 1970. A study of the male genitalia in Bruchidae (Coleoptera). Proceedings of the Entomo- logical Society of Washington 72: 370-386. NYHOLM, T. 1969. Uber Bau und Funktion der Kopulation- sorgane bei den Cyphones (Col. Helodidae). Ento- mologisk Tidskrift 90: 233-271. RODRIGUEZ, V. , WINDSOR, D. M. & EBERHARD, W. G. 2004. Tortoise beetle genitalia and demonstrations of a sexually selected advantage for flagellum length in Chelymorpha alternans (Coleoptera, Cassidini, Sto- laini). Pp. 739-748 in: JOLIVET, P., SANTIAGO-BLAY, J. A. & SCHMITT, M. (eds.) New Developments in the Biology of Chrysomelidae. SPB Academic Publishing bv, The Hague. SCHMITT, M. 1996. The phylogentic system of the Chry- somelidae — history of ideas and present state of knowledge. Pp. 57-96 in: Jolivet, P. H. A. & Cox, M. L. (eds.) Chrysomelidae Biology, vol. 1: The Classifi- cation, Phylogeny and Genetics. SPB Academic Pub- lishing bv, Amsterdam. THORNHILL, R. 1983. Cryptic female choice and its impli- cation in the scorpionfly Harpobittacus nigriceps. The American Naturalist 122. 765-788. Authors’ address: Susanne DUNGELHOEF (correspond- ing author) and Michael SCHMITT, Zoologisches For- schungsmuseum Alexander Koenig, Adenauerallee 160, D-53113 Bonn, Germany; E-mail: s.duengelhoef.zfmk (@uni-bonn.de, m.schmitt@uni-bonn.de Bonner zoologische Beitráge Band 54 (2005) | Heft 4 | Seiten 209-237 Bonn, Oktober 2006 The Current Status of Knowledge of the Alticinae of Mexico (Coleoptera: Chrysomelidae)' David G. FURTH National Museum of Natural History, Smithsonian Institution, Washington, D.C., USA Abstract. The current state of knowledge of the diversity for Alticinae recorded from Mexico is taken from the histori- cal literature, determined specimens primarily in the U.S. National Museum, including material collected by the author, as well as from several other institutions. An Appendix is provided containing a list of generic and specific names re- corded from Mexico, including the known distribution for each species by Mexican state and the source for that infor- mation. There are 89 genera and 524 species currently known from Mexico. Although preliminary, this is a significant increase from the published literature. The Alticinae species diversity per Mexican state is also provided. Some discus- sion of the Mexican Alticinae biogeographic affinities is given. The following changes in status are made: Syphrea bibi- ana Bechyné = Nesaecrepida infuscata (Schaeffer); Palaeothona Jacoby is raised from synonymy with Lupraea Jacoby and considered valid; Varicoxa Bechyné = Luperaltica Crotch, Kuschelina gracilis Jacoby is raised from synonymy with K. /aeta (Perbosc) and considered valid; Aphthona dimidiaticornis Jacoby, A. fulvipennis Jacoby, A. maculipennis Jacoby, A. mexicana (Jacoby), A. pallipes (Jacoby), A. semipunctata Jacoby, and A. substriata Jacoby are all considered to be in Centralaphthona Bechyné; Aphthona amulensis Jacoby = Genaphthona Bechyné; Aphthona fulvitarsis Jacoby = Gioia Bechyné; A. championi Jacoby = Lupraea. Keywords. Alticinae, flea beetles, Chrysomelidae, Mexico, biogeography 1. INTRODUCTION Mexico is the largest part (ca. 50%) of the Meso- american biodiversity “hotspot” which is second in spe- cies diversity and endemism of the top 25 “hotspots” in the world (MITTERMEIER et al. 1999). It has been called one of the world’s “megadiversity” countries (MITTER- MEIER 1988) and ranks third in this respect according to some. Mexico is situated in a biogeographic transition zone between the Nearctic Region to the north and the Neotropical Region to the south. Generally in Mexico the Nearctic biotic elements are in the northern areas, the central arid plateau with some arid elements reach- ing as far south as Oaxaca. The Neotropical elements are in the south with some elements reaching along the coasts to Tamaulipas in the east and to southern Sinaloa in the west. Species diversity/richness has been docu- mented to be very high in many well-known groups, such as plants (RZEDOWSKI 1993, 1996), mammals (FA & MORALES 1993; MITTERMEIER et al. 1999), and but- terflies (LLORENTE & MARTINEZ 1993, LLORENTE et al. 1996). Species diversity also varies from group to group and from state to state. The highest diversity generally occurs in the more tropical states such as Chiapas, Oax- aca and Vera Cruz. Some examples of this from the in- sect world include Odonata (GONZALEZ & NOVELO 1996), Pscoptera (MOCKFORD & GARCIA ALDRETE 1996), Passalidae (REYES-CASTILLO 2002), Bruchidae with Jalisco and Morelos also high (NAPOLES 2002). Paper presented to the 6" International Symposium on the Chry- somelidae, Bonn, Germany, May 7, 2004 Because of this biogeographic transition zone between the Nearctic and Neotropical Regions, biogeographic af- finities may also vary greatly depending on the group considered. Examples of this in the Coleoptera are spe- cies of Curculionidae with 41% Neotropical and 14% Nearctic affinities (ANDERSON & O’BRIEN 1996) and Carabidae with 54% Nearctic and 14% Neotropical, but in generic affinities 40% are Neotropical and 20% are Nearctic (BALL & SHPELEY 2000). Levels of endemism vary greatly depending on the group considered and, of course, depending on the rela- tive knowledge of the group. As with any country some vertebrate and plant groups are well know, whereas most insect groups are not. In mammals about 32% of the species are endemic (FA & MORALES 1993) and in the phanerogamic plants 10% of the genera and 52% of the species are endemic (RZEDOWSKI 1993). Among the insects butterflies are well-known, yet in Mexico the Papilionidae have only 10% endemism there (LLOR- ENTE & MARTINEZ 1993). For native bees the level of endemism ranges from 25-60% depending on the region of Mexico, both the tropical state of Vera Cruz and the north of Baja California have 30%, whereas Guerrero has 50% and the Balsas (Guerrero-Oaxaca) has almost 60% (AYALA et al. 1993). Species endemism is gener- ally high, but may vary, for example, in Homoptera it varies from 4% in Membracidae to 63% in Aleyrodidae (O’BRIEN et al. 1996), Pscoptera 74% (MOCKFORD & GARCIA ALDRETE 1996), Carabidae 60% (Ball & Shpe- ley 2000), Tenebrionidae 52% (AALBU et al. 2002), Curculionidae 40.5% (ANDERSON & O’BRIEN 1996). 210 Bonner zoologische Beitráge 54 (2005) Nearctic Alticinae (ca. 40 genera) are reasonably well known at the genus and species level in comparison to the Neotropical taxa (over 200 genera); however, there are certainly many more new species to be discovered there. Recently there was an update of the treatment at the genus level (RILEY et al. 2002) including keys. RI- LEY et al. (2003) have published a detailed checklist to the Nearctic Chrysomelidae. Neotropical Alticinae have been recently treated in a number of references by J. and B. BECHYNE (see References), D. BLAKE (see Refer- ences), WILCOX (1975), SCHERER (1983), FURTH & SAVINI (1996, 1998), FURTH et al. (2003), and FURTH (2004). Almost half of the known Alticinae genera oc- cur in the Neotropics; about 230 generic names were listed in SCHERER (1983) about 90 of which were de- scribed by Jan and Bohumila Bechyné. Fig. 1. Asphaera abdominalis (Chevrolat) on Ludwigia peploides (Kunth) Raven, in copula, Zacatecas, July 1995. (Photo: D. Furth). In the historical literature (Tables 1 and 2), JACOBY in the Biologia Central: Americana (1884-1892) listed 334 species in 50 genera from Mexico, many of these genera and species were listed under old names that were later changed. HEIKERTINGER & CSIKI (1939-40) added 23 species and three genera Argopistes Motschulsky, Ble- pharonycha (Fall) and Heikertingerella (Csiki). BLACK- WELDER (1946) added 12 species (9 from Baja Califor- nia) and four genera: Distigmoptera (Blake); Glyptina (LeConte); Dysphenges (Horn); Euplectroscelis (Crotch) — the first to correctly clarify this genus. From more recent literature, FURTH & SAVINI (1996) listed 400 species in 74 genera. FURTH & SAVINI (1998) added 1 genus and 11 species. FURTH (1998) added 18 species of Blepharida Chevrolat and synonymized Ble- pharonycha as well as 3 species of Blepharida (s.s) (see Table 1). RILEY et al. (2001) placed Hornaltica atriven- tris (Melsheimer) in Margaridisa, thus eliminating the former as a separate genus known from Mexico and they raised Hemiphyrnus Horn out of synonymy with Phyrnocepha Baly, thus adding another genus name to the Mexican checklist. FURTH (2004) added 11 genera and 69 species to the Mexican fauna. Mexico is a very large country, 14" largest in the world, so one would expect a high diversity of species com- pared to smaller countries in Central America. There are about 350 species in 89 genera currently known from Costa Rica, particularly because of some recent inten- sive quantitative inventorying done at a single site (FURTH et al. 2003). In contrast, in the smaller and less surveyed Panama, there are only 270 species in 70 gen- era known (FURTH & SAVINI 1996, 1998). Fig. 2. Capraita conspurcata (Jacoby), Morelos, June 1993. (Photo: D. Furth). its dts OIE) RE RE TEE Fig. 3. Blepharida lineata Furth on Bursera trimera Bullock, Guerrero, June 1993. (Photo: D. Furth). David G. FURTH: Alticinae (Chrysomelidae) of Mexico Table 1. Numbers of historical and current Mexican Alticinae species Genus Species Source 74 400 FURTH & SAVINI (1996) | 11 (-1) FURTH & SAVINI (1998) -] 18 (-3) FURTH (1998) l 3 SAVINI et al. (2001) -1(1) 0 RILEY et al. (2001) 0 3 GILBERT & ANDREWS (2002) 9 32 FURTH (2004) — USNM 3 12 FURTH (2004) — USNM-DGF 0 25 FURTH (2004) — MCZC-FCB -] 4 (-3) Corrections to FURTH & SAVINI (1996) 4(-1) 21 NEW - this study 89 524 Total Table 2. Mexican Alticinae Genera — old and new records Genera General Distribution Source (Mexico) 13 Mexico, etc. FURTH & SAVINI (1996) “Crepidodera” Mexico (Baja California) FURTH & SAVINI (1998) - (Blepharonycha) Mexico FURTH (1998) Gioia Brazil, Venezuela, Colombia, Ecuador, Guade- SAVINI et al. (2001) loupe, Panama, Costa Rica Hemiphyrnus USA, Mexico RILEY et al. (2001) -/+ Hornaltica USA RILEY et al. (2001) Ayalaia El Salvador FURTH (2004) - USNM Chalatenanganya Venezuela, Trinidad, El Salvador, Guatemala, FURTH (2004) - USNM-DGF Costa Rica Egleraltica Brazil, Venezuela, Costa Rica FURTH (2004) -USNM Exoceras Brazil, Bolivia, Venezuela, Trinidad, St. Vincent, FURTH (2004) - USNM St. Lucia, Dominica, Guadeloupe, Puerto Rico, Dominican Republic, Haiti, Jamaica, Cuba, Pa- nama, Costa Rica Genaphthona Brazil, Paraguay, Bolivia, Venezuela, El Salva- FURTH (2004) - USNM dor, Costa Rica, Guatemala, Leptophvsa Brazil, Paraguay, Venezuela, Jamaica, Panama, FURTH (2004) — USNM Costa Rica Margaridisa Brazil, Venezuela, Costa Rica, Nicaragua FURTH (2004) - USNM Neothona Venezuela, Trinidad, El Salvador, Costa Rica FURTH (2004) - USNM Nesaecrepida USA, Jamaica, Cuba, Puerto Rico FURTH (2004) - USNM-DGF Terpnochlorus Venezuela, Madagascar FURTH (2004) -USNM Tetragonotes Brazil, Bolivia, Peru, Ecuador, Colombia, Vene- FURTH (2004) - USNM zuela, Panama, Costa Rica, Nicaragua, Guate- mala Deuteraltica Guatemala, El Salvador USNM — New Heikertingeria Guatemala,Panama, Venezuela, Bolivia, Peru, USNM — New Brazil, Trinidad Luperaltica USA, C. & S. America USNM - New Palaeothona C. & S. America New - Varicoxa C. & S. America New 89 Total Ashpaera abdominalis (Chevrolat) (Fig. 1) is one of the most common species in Mexico and, in fact in all of Central America. Another common species in Mexico 1s Capraita conspurcatus (Jacoby) (Fig. 2) that is often found in very large numbers and may completely destroy its host plants. New World Blepharida were monogra- phed by FURTH (1998) and of the 38 species, 31 are en- - demic to Mexico and all but 2 can be found in Mexico. Some species are apparently oliphagous and are recorded on several species of Bursera (FURTH 1998); however, most species are monophagous on species of Bursera (Burseraceae), such as B. lineata (Fig. 3). 2. METHODS Records of Mexican Alticinae were recovered in several ways for the current study. First a search of published literature, especially the references in FURTH & SAVINI (1996, 1998) as well as more recent publications, was made and the locality data for the Mexican states checked. The identified and unidentified collections of the National Museum of Natural History (Smithsonian Institution) (USNM) were checked for Mexican state records using a recent species inventory funded by the Smithsonian Women’s Committee. The author deter- mined many of the unidentified Mexican Alticinae specimens at the USNM. Additional Mexican state dis- tribution data was extracted from the F. C. Bowditch Collection (FCB) previously on long-term loan to the author from the Museum of Comparative Zoology (Harvard University - MCZC). Also some material col- lected and determined by the author in Mexico was in- cluded (USNM-DGF). Finally some records from a re- cent visit to the Naturhistorisches Museum Basel (Switzerland) (NHMB) and the Zoologisches Staats- sammlungen München (Germany) (ZSMC) were also incorporated. Some representative specimens will be deposited at the Colección Nacional de Insectos, Insti- tuto de Biología, Universidad Autonoma de Mexico (UNAM-IB). 3. RESULTS 3.1. General The Appendix contains the most up-to-date checklist of Mexican Alticinae with distributional information at the Mexican state level and a number of new synonymies, new combinations and other name changes (see below). Since FURTH (2004) the author has discovered three ad- ditional genera and 21 species to the Mexican fauna. Five species recorded in the historical literature only as “Mexico” have been more specifically recorded in par- ticular Mexican states. Thus, the total Alticinae diver- sity of Mexico consists of 89 genera and 524 species (Tables | & 2). Bonner zoologische Beitráge 54 (2005) 3.2. New synonymy and other nomenclatural changes During the course of this project the author has discov- ered several notable new synonymies and other name changes that should be clarified in order to facilitate the consistent usage of the names of Neotropical Alticinae, especially Mexican Alticinae. Syphrea bibiana Bechyné was described by BECHYNE (1955) from British Honduras. The author has examined the holotype and type series in the Frey Collection at NHMB and finds that it is conspecific with Nesae- crepida infuscata (Schaeffer) [combination established by RILEY et al. 2001]. SCHAEFFER (1906) described this species from a single specimen from Brownsville, Texas, deposited in the USNM and the author has also examined this holotype specimen. SCHAEFFER (1906) described this species as Lactica infuscatus and in- cluded a key to the three species of the genus, including L. brumeliae Schaeffer and L. tibialis Olivier. Lactica Erichson is currently considered to be a synonym of Monomacra Chevrolat. Lactica brumeliae is currently considered to be in the genus Monomacra and L. tibialis in the genus Parchicola Bechyné (RILEY et al. 2003). All of these species share a rather deeply impressed, transverse subbasal impression on the pronotum and this is no doubt why SCHAEFFER assumed they were con- generic. However, BLAKE (1964) accurately established the genus Nesaecrepida (type species N. asphaltina (Suffrian)) from Cuba and differentiated it from other “Crepidoderini” by having inconspicuous elytral punc- tation consisting of many fine striae. She also pointed out that the prebasal transverse impression extends completely across to the lateral margins of the pronotum without the usual limiting lines perpendicular to the base. This feature is also characteristic of Syphrea Baly and is probably the primary reason that Bechyné placed this species (as S. bibiana) in that genus. Nesaecrepida and Syphrea share some characteristics such as, base of the pronotum somewhat doubly sinuate, convex prono- tum. They differ in the shape of the epipleura (usually not straight, gently emarginate in Syphrea), antennal bossae of Nesaecrepida small round (usually larger and not round in Syphrea) divided by base of the long nar- row frontal ridge. Further characteristics of Nesae- crepida infuscata are: pronotum with anterolateral an- gles rounded, thickened and not distinctly angled, punctation dense, fine; dorsum light brown-yellow in color with elytral suture distinctly darkened, elytral punctures coarse, dense, not evidently striate; epipleura rather wide and parallel-sided, tapering only subapi- cally. This species has recently been studied as a potential bio- logical control agent for Mimosa pigra Linnaeus (Mi- David G. FURTH: Alticinae (Chrysomelidae) of Mexico 213 mosaceae) in Australia (W. Forno, Brisbane, pers. comm. 1992). BECHYNE & BECHYNE (1960:23-24) first designated Palaeothona Jacoby (1885:377) [type species: P. rugi- frons Jacoby, designated by BLAKE (1950:179)] as syn- onymous with Lupraea Jacoby (1885:359) [type spe- cies: L. longicornis Jacoby, designated by BECHYNÉ & BECHYNÉ (1960:24)]. The author has examined syn- types of Lupraea longicornis as well as of Palaeothona rugifrons and has determined that these two genera are in fact distinct as originally described by JACOBY (1885). BLAKE (1950) described four new species of Palaeothona from the USA, Mexico and Haiti and newly combined Trachymetopa (misspelled in BLAKE 1950 as Trachymetropa Weise) picta (Say) into Pa- laeothona. Palaeothona picta (Say) was originally de- scribed as Altica Geoffroy, then placed in Aphthona Chevrolat by CROTCH (1873), an then in Phyllotreta Chevrolat by HORN (1889). In fact in his original de- scription of the genus Jacoby compared Palaeothona to Aphthona and even to Phyllotreta, but acknowledged that characters to distinguish it as a separate genus were not obvious. While Palaeothona and Lupraea Jacoby have some similarities, the differences are more evident (Vilma P. Savini, Maracay, Venezuela, in litteris): Pa- laeothona has a more convex and robust body form; Lupraea has antennomere 2 almost moniliform, much less than half the length of segment 3, Palaeothona has a relatively longer second antennomere, more than half the length of segment 3; Lupraea epipleura usually nar- row throughout most of its length, not much wider than the first antennomere, gradually tapered apically, epipleura of Palaeothona wider throughout its length, tapered only subapically; eyes larger in most Lupraea; pronotum of Palaeothona with broad mid-basal depres- sion; pronotal shape more elongate and evidently nar- rowed basally with anterolateral angles evenly rounded in Lupraea opposed to often angulate in Palaeothona; elytral bossae usually more pronounced in Palaeothona; elytral punctation denser and coarser in most Lupraea; antennal bossae small narrow, rounded or vertically aligned in Palaeothona, but horizontal, transverse or rectangular in Lupraea; antennae long, often as long or longer than body in Lupraea, shorter in Palaeothona; dorsum rather flattened in Lupraea and more convex in Palaeothona; pronotum and vertex of Lupraea usually smooth and impunctate, but usually at least vertex dis- tinctly punctured or subrugose in Palaeothona; elytral punctation often coarser and denser in Lupraea. Most Mexican species were described as Palaeothona by Jacoby, but he also confused these two genera and so it is necessary to examine type specimens of each species in order to correctly place each in the correct genus. The author has been able to examine types of some, but not all of the Mexican species and, thus, verifies the place- ment of these as follows: Palaeothona discrepans (Schaeffer); Lupraea elongata (Jacoby); Lupraea fulvi- collis Jacoby; Lupraea godmani (Jacoby); Lupraea gua- temalensis (Jacoby); Lupraea imitans (Jacoby); Palaeothona melanocvanea Blake; Palaeothona rubro- viridis Blake; Palaeothona rugifrons Jacoby; Lupraga semifulva (Jacoby); Lupraea smithi (Jacoby); Pa- laeothona viridis Jacoby. Lupraea occipitalis Bechyné & Bechyné is problematic and does not seem to fit in el- ther genus well, but the author prefers to leave it in its current status until it can be studied in more detail. However, Palaeothona dilaticornis Jacoby and Pa- laeothona frontalis Jacoby are left in their originally de- scribed genus until their types can be studied. Therefore, the species here placed in Palaeothona are given a new status from recent publications (e.g., SAVINI & FURTH 1996, 1998; Riley et al. 2003; and FURTH 2004). CROTCH (1873:70) established Luperaltica [type spe- cies: L. senilis (Say) designated by WILCOX (1953:54)] for two species, including L. fuscula (LeConte) = L. ni- gripalpis (LeConte), from the USA. According to RILEY et al. (2003) this genus now includes two others from the USA (Z. nitida Wilcox and L. semiflava Fall). Upon close examination and comparison with many species of the genus Varicoxa Bechyné (1955:81) [type species: Systena ustulata Harold designated by BECHYNE (1955:81)], the author here designates Varicoxa Be- chyné to be a synonym of Luperaltica Crotch. The au- thor has examined many species of Varicoxa deter- mined by Bechyné (MIZA, NHMB), including Y. ustulata (Harold), and types of V. ustulata centralis Be- chyné, V. ustulata inconstans Bechyné, as well as other reliably determined species. In the course of describing L. nitida, WILCOX (1953) gives a very good account of the unusual history of the species of Luperaltica as well as of their morphology, including aedeagus figures. The original descriptions of both Luperaltica and Vari- coxa were very inadequate. CROTCH (1873) only said that Luperaltica looked like Luperus Mueller (Galeruci- nae), but had “incrassate femora, entire epipleurae, separated anterior coxae ...” making it an Alticinae. Also he stated that it was similar to Orthaltica Crotch, but the elytra more finely sculptured, not striate, first metatarsal segment elongate, anterior coxae closed (nar- rowly), base of prosternum dilated to meet epimera, an- tennae long with basal segment short. BECHYNE (1955) only said that Varicoxa had species with closed and open procoxal cavities, heart-shaped pronotum with a weak transverse, prebasal impression, long first metatar- sal segment, large head with projecting eyes wider than front margin of the pronotum. BECHYNE & BECHYNE (1967) give a checklist of the 26 described species of Varicoxa, 18 described by BECHYNE. Although the four North American species do not have the pronotum as evidently constricted basally, and L. senilis and L. ni- 214 Bonner zoologische Beitráge 54 (2005) gripalpis have most of the dorsum shagreened, other morphological characters, including those listed above, are the same. In addition, the males of all species have an unusually large, U-shaped opening at the apex of the strongly ventrally-projecting abdomen, apex of the aedeagus usually protruding, a thin, longitudinal, scle- rotized midline on the 7" ventrite, and similar form of ~ the aedeagus. In addition, comparing Varicoxa ustulata; Luperaltica senilis, L. nigripalpis in both: the meta- femoral spring is identical and of the Phyllotreta Mor- pho-Group; the aedeagus is very wide in ventral/dorsal view and strongly curved in lateral view, especially in apical 1/3, very broad basal and subapical/dorsal foram- ina; the spermatheca is C-shaped, forming % of a circle, receptacle and pump not differentiated in shape, equal width throughout, ductus a tight cluster of several coils. Therefore, the following species formerly considered as Varicoxa are newly combined in Luperaltica: L. longi- cornis (Jacoby); L. ustulata centralis (Bechyné); and L. viridipennis (Jacoby). Changes since FURTH (2004) include: Disonycha fumata labiata Jacoby, 1901 is synonymous with Disonycha fumata fumata LeConte, 1858, new synonym in BLAKE (1955: 23); Disonycha horni Jacoby, 1891 is synonymous with Disonycha fumata fumata Le- Conte, 1858, new synonym in BLAKE (1933b: 36). The “Oedionychina” genera Walterianella Bechyné, Alagoasa Bechyné, and Capraita Bechyné are often dif- ficult to distinguish using the traditional morphological characters as given in descriptions and keys like SCHERER (1962, 1983). There is increasing evidence that these genera and others are paraphyletic (DUCKETT & KJER 2003). Walterianella signata (Jacoby, 1886:431) was mistakenly listed under Alagoasa in FURTH & SAVINI (1996), but should have been listed as Walterianella as it was in WILCOX (1975), because it has the characteristics of that genus, especially the thin explanate sides of the pronotum and elytral epipleura (see SCHERER 1962, 1983); the author has examined syntypes in FCB. Also, there is increasing evidence that Walterianella and Capraita cannot reliably be separated and are possibly paraphyletic (DUCKETT & KJER 2003; C. Duckett, Washington, DC, pers. comm. 2004). The morphological characters given in SCHERER (1983), es- pecially the prebasal, transverse impression on the pronotum, is not very reliable. However, for the present study the author prefers to retain these genera as much as possible and considers Capraita durangoensis (Jacoby, 1892: 318) to be best placed in the genus Wal- terianella, and as such is a new combination; the author has examined syntypes in FCB. HORN (1889: 184) considered Oedionychis gracilis Jacoby as a synonym (variety) of O. interjectionis (Crotch 1873: 61). This was confirmed by BLAKE (1927: 15), HEIKERTINGER & CSIKI (1940), and BLACKWELDER (1946). Oedionychis interjectionis was apparently first considered a synonym of Kuschelina laeta (Perbosc 1839: 263) by WILCOX (1975) and later upheld by RILEY et al. (2003). Kuschelina gracilis seems to have the following morphological differences from K. /aeta: elytra green with strongly shagreened surface, opposed to black smooth surface in K. laeta; longitudinal, subsutural, light stripes straight and equal in width throughout, not apically enlarged or angled outward as in X. /aeta; pronotum distinctly punctate or shagreened, often densely, rather than smooth, mostly impunctate in K. /aeta. There are specimens of K. gracilis with green elytra, without any longitudinal sub- sutural light stripes. The author has examined syntypes of K. gracilis in FCB and many specimens of K. laeta in USNM. Therefore, for this study the author considers K. gracilis and K. laeta to be separate and valid taxa at the species level. Further detailed study of the genitalia and molecular study may better clarify the relationship be- tween these taxa. FURTH & SAVINI (1996) and SAVINI & FURTH (1998) pointed out that the genus Aphthona Chevrolat, 1836, in its true sense from the Old World, does not exist in the New World. Most of the Neotropical species previously listed in Aphthona have been reassigned to Brasilaph- thona Bechyné, 1956, Centralaphthona Bechyné, 1960, Genaphthona Bechyné, 1956. However, FURTH & SAV- INI (1996) provisionally listed as Aphthona some Neotropical species still in need of study in order to place them in correct genera. For Centralaphthona some significant characters are: convex dorsum; no pronotal transverse, subbasal im- pression; anterolateral angles distinctly beveled; elytra striate. After examination of specimens determined by Jacoby (FCB) the author has determined that the following spe- cies (formerly in Aphthona) are best placed in Centra- laphthona and, thus, constitute new combinations: C. dimidiaticornis Jacoby, 1891; C. fulvipennis Jacoby, 1885; C. maculipennis Jacoby, 1891; C. mexicana (Jacoby, 1885); C. pallipes (Jacoby, 1891); C. semi- punctata Jacoby, 1891 [syntypes examined]; C. substri- ata, Jacoby, 1891. However, there is some indication that Centralaphthona Bechyné, 1960 [type species: C. deyrollei, Baly, 1877:296] is synonymous with Glyptina LeConte, 1859:26 [type species: G. spuria LeConte, 1859:26]. This was indicated by RILEY et al. (2002) who state that “The Neotropical genus Centralaphthona Be- chyné, 1960, is probably a synonym [of Glyptina Le- Conte, 1859]”. The author tends to agree with this statement; however, more thorough study of these gen- era is needed and so the above new synonyms remain as Centralaphthona in this study. David G. FURTH: Alticinae (Chrysomelidae) of Mexico 21 Aphthona pilatei Baly, A. smithi Jacoby, and A. unicolor Jacoby will tentatively remain in Aphthona until they can be studied further, especially type specimens. Al- though syntypes of 4. unicolor (FCB) were examined by the author and it appears to belong in the “Monomacrini”, further study of other genera in that pu- tative tribe is needed. After examination of several species determined by BE- CHYNE to be in the genus Genaphthona, including syn- types (FCB) of G. transversicollis (Jacoby), the author places A. amulensis Jacoby, 1891, in Genaphthona. This species has the flattened dorsum with very fine puncta- tion, lateral margins of pronotum rounded, shallow sub- basal, transverse pronotal impression, and very faint an- tennal bossae typical of Genaphthona. Table 3. Mexican Alticinae by State State Abbreviation No. Species Aguascalientes AGS 7 Baja California BC 20 Baja California Sur BCS 13 Campeche CAMP 7 Chiapas CHIS 81 Chihuahua CHIH 43 Coahuila COAH 11 Colima EOL 21 Distrito Federal DF 41] Durango DGO 97 Guanahuato GTO 4] Guerrero GRO 141 Hidalgo HGO 29 Jalisco JAL 45 Mexico MEX 38 Michoacan MICH 47 Morelos MOR 84 Nayarit NAY 23 Nuevo Leon NL 12 Oaxaca OAX 122 Puebla PUE 48 Queretaro QRO 2 Quintana Roo QROO 8 San Luis Potosi SLP 3] Sinaloa SIN 17 Sonora SON 26 Tabasco TAB 81 Tamaulipas TAMPS 44 Tlaxcala TLAX 2 Veracruz VER 198 Yucatan YUC 23 Zacatecas ZAC 10 Species of the genus Gioia Bechyné, 1955, can also be confused with Aphthona (SAVINI & FURTH 1996); how- ever, Gioia has a very robust, convex body shape; dis- tinctly confused and strong elytral punctures; transverse in pronotum with strong punctures and thickened, broadly slanted anterolateral angles; usually evident transverse elytral impression. Considering these characters and af- ter examining a syntype (FCB) of A. fulvitarsis Jacoby, 1891:246, the author concludes this species is best placed in Gioia. / After examination of a syntype (FCB) of 4. championi Jacoby, 1885, and based on the characters given above, the author places this species in Lupraea. 4. DISCUSSION The author’s strategy for producing an inventory or fau- nal list consists of three phases. First, to document the historical record through the literature. Second, to sur- vey institutional collections; in this study including USNM, MCZC-FCB, NHMB, and ZSMC. The final phase is to conduct fieldwork. There have been very few Alticinae specialists that have collected in Mexico and none have ever lived there, so the above approach should prove very productive. Comparing the totals of genera and species recorded in the recent literature, the above-mentioned inventory method has significantly in- creased the totals, even at this preliminary stage. The number of recorded genera has risen from 74 to 85 (FURTH 2004), to 89, and the species from 431 to 50] (FURTH 2004) to 524. Some of the records need to be verified further, as well as some historical identifica- tions in the FCB. Even though this is a very preliminary list of Mexican Alticinae diversity, especially in comparison to the rela- tively more field-surveyed Costa Rican fauna (FURTH et al. 2003), still the Mexican fauna demonstrates high po- tential species richness. Comments on Biogeography Unlike many of the vertebrate, plant and even some in- sect groups mentioned above in the Introduction, spe- cies of Mexican Alticinae are poorly known. The cur- rent study is still a preliminary attempt to compile records from the literature and a few collections into a baseline from which to build in the future. Table 2 lists historical and some recently recorded (FURTH 2004) genera, including two actual newly recorded genera (Deuteraltica and Heikertingeria). Most of these genera have Neotropical affinity and in fact most are distrib- uted deep into South America. Gioia was discovered re- cently in Mexico (SAVINI et al. 2001), but has a similar distribution to the 12 genera recorded in FURTH (2004). Ayalaia Bechyné & Bechyné until now was only known from El Salvador. Deuteraltica Bechyné & Bechyné is monotypic and previously only known from Guatemala and El Salvador. Heikertingeria Csiki is South Ameri- can with only H. clarki extending into true Central America. Luperaltica is distinctly a Nearctic genus ex- 216 Bonner zoologische Beitráge 54 (2005) Division Politica Estatal 26 13 SIMBOLOGIA BE AGUASCALIENTE BE MORELOS BAJA CALIFORNIA NAYARIT BAJA CALIFORNIA SUR NUEVO LEON 0 CAMPECHE BEI OAXACA CHIAPAS PUEBLA | CHIHUAHUA I) QUERETARO DE ARTEAGA COAHUILA DE ZARAGOZA QUINTANA ROO 0 COLIMA SAN LUIS POTOSI EXI DISTRITO FEDERAL SINALOA BEE DURANGO ~~ SONORA EI GUANAJUATO TABASCO HA GUERRERO TAMAULIPAS = HIDALGO BEE TLAXCALA JALISCO VERACRUZ-LLAVE © MEXICO YUCATAN MICHOACAN DE OCAMPO ZACATECAS Map 1. Mexican Alticinae diversity per State tending into the Neotropical Region. Neothona Bechyné is clearly a Neotropical element. Paranaita Bechyné is a South American genus previously only known as far north as Panama and its distribution this far north is somewhat surprising, more detailed and verification of the Mexican distribution records would be useful. Nesaecrepida Blake has an apparent Caribbean (Neotropical) affinity and the record in this study of the type species N. asphaltina in the Yucatan demonstrates the Caribbean affinities of that part of Mexico. Exoceras Jacoby is particularly widely distributed even through- out many Caribbean islands in almost an are from the northern South American continent. Given the extent of the known distribution of all of the above genera it is not really surprising to find them also in Mexico. The most unusual distribution with no obvious explanation is Terpnochlorus Fairmaire represented in Mexico by 7. americana, also known from Venezuela, but the genus has a very strange disjunct distribution and is also in Madagascar, central and southern Africa. Among the other genera listed from Mexico in the Ap- pendix the majority are of primarily Neotropical affin- ity. Genera such as Altica Geoffroy, Chaetocnema 200 O 200 400 [me es | FUENTE: Instituto Nacional de Estadistica, Geografia e Informatica Stephens, Dibolia Latreille, Epitrix Foudras, Longitar- sus Berthold, Phyllotreta Chevrolat, Psylliodes Berthold are cosmopolitan and have no particular affinity to the New World. Some genera such as Disonycha Chevrolat, Kuschelina Bechyné, Monomacra Chevrolat, Systena Chevrolat although basically Neotropical also have sig- nificant penetration at the species level into the Nearctic Region. Crepidodera peninsularis Horn may be a true Crepidodera Chevrolat in which case this is a Nearctic affinity. However, it is apparent that about 90% of the biogeographic affinity of Mexican Alticinae at the ge- nus level is Neotropical. There are a few genera en- demic to Mexico such as Caloscelis Clark, Euplec- troscelis Crotch from Baja California, possibly Iphitroides Jacoby with four species in Mexico and one recorded species in Venezuela, but after careful study this latter species may be re-classified. A few other gen- era (e.g., Dysphenges Horn, Phydanis Horn, Pseudor- thygia Csiki) were previously known only to occur in Mexico, but recently RILEY et al. (2002) recorded them occurring in extreme southern USA. A few genera are not endemic to Mexico, but most of the species in them are; e.g., Blepharida Chevrolat, Chrysogramma Jacoby, Phyrnocepha Baly, Plectotetra Baly. David G. FURTH: Alticinae (Chrysomelidae) of Mexico 217 Blepharida has been well studied in Mexico (FURTH 1998) and there is an interesting eco-geographic pattern. New World Blepharida ıs distributed from the USA (one species) through Mexico (35 species) to Costa Rica and El Salvador (1 species). One species is also only found in the Caribbean. The genus Notozona Chevrolat is distributed from southern Mexico through the rest of Central America into northern South America. Table 3 and Map | indicate some preliminary general trends about the diversity and biogeographic affinities of the Mexican Alticinae species. As with many other groups of animals the highest diversity is in the southern states with strong Neotropical affinities such as Chiapas, Oaxaca, Guerrero, Vera Cruz and Tabasco. The latter two demonstrate the significant Neotropical affinity that extends northward along the Caribbean coast. The rather high diversity in Durango and Morelos is not easily ex- plainable at this time, in the Bruchidae NAPOLES (2002) also recorded a high species diversity in Morelos. How- ever, in the Alticinae possibly both the elevated num- bers in Durango and Morelos as well as the relatively low number of species in the states of the Yucatan Pen- insula, Aguascalientes, Queretaro, Tlaxcala, Zacatecas and others are probably primarily a factor of collecting bias; 1.e., better or poorly collected areas. Therefore, while it is more difficult to analyze the spe- cies level richness and biogeographic affinities at this very preliminary stage of knowledge of the Mexican Al- ticinae fauna, superficially it appears that the vast ma- jority of species have more Neotropical affinities than Nearctic affinities. Acknowledgements. | thank Michael Schmitt for organizing the Sixth International Symposium on the Chrysomelidae and editing its Proceedings. I am grateful to the following: E. O. Wilson and P. Perkins (MCZ, Harvard University, Cambridge) for the long-term use of the F. C. Bowditch Collection, Alexander Konstantinov (USNM, SEL, USDA) for some help with access to the collection at USNM, Santiago Nifio M. (Universidad Nacional Autonoma de Tamaulipas) for specimens and records from Tamaulipas, Eva Sprecher (Naturhistorisches Museum Basel), Martin Baehr (Zoologische Staatssammlung Miinchen) and Vilma Savini (Universidad Central de Venezuela, Museo del In- stituto de Zoologia Agricola, Maracay) for access to specimens at their respective museums. Support for the fieldwork by the author in Mexico (1991, 1993, 1995, 1997, 2002) was from personal funds. REFERENCES AALBU, R. L., FLORES, G. E. & TRIPLEHORN, C. A. 2002. Tenebrionidae. Pp. 499-512 in: LLORENTE B., J. & Morrone, J. J. (eds.) Biodiversidad, Taxonomía y Biogeografía de Artrópodos de México: Hacia una sín- tesis de su conocimiento. Vol. 3. UNAM, Mexico. ANDERSON, R. S. & O’BRIEN, C. W. 1996. Curculionidae (Coleoptera). Pp. 329-351 in: LLORENTE B., J., GAR- CÍA ALDRETE, A. N. & GONZÁLEZ S., E. (eds.) Biodi- versidad, Taxonomía y Biogeografía de Artrópodos de México: Hacia una síntesis de su conocimiento. Vol. 1. UNAM, Mexico. AYALA, R., GRISWOLD, T. L. & BULLOCK, S. H. 1993. The native bees of Mexico. Pp. 179-227 in: RAMAMOOR- THY, T. P., BYE, R., LOT, A, & FA, J. (eds.) Biological Diversity of Mexico: Origins and distribution. Oxford Univ. Press, New York. BALL, G. E. & SHPELEY, D. 2000. Carabidae (Coleoptera). Pp. 363-399 in: LLORENTE B. , J., GONZALEZ S., E. & PAPAVERO, N. (eds.) Biodiversidad, Taxonomía y Biogeografía de Artrópodos de México: Hacia una sín- tesis de su conocimiento. Vol. 2. UNAM, Mexico. BALSBAUGH, E. U. Jr. 1980. New collection records of three rare species of flea beetles: Distigmoptera foveo- lata Balsbaugh, D. texana Blake, and Altica nancyae Stirrett. Journal of the Kansas Entomological Society 53 (4): 833-836. BALY, J. S. 1877. Descriptions of new genera and of un- characterized species of Halticinae. Transactions of the Entomological Society of London 1877 (4): 283-323. BARBER, H. S. 1941. Some fleabeetles injurious to beans in tropical America (genus Diphaulaca, family Chry- somelidae). Proceedings of the Entomological Society of Washington 43 (3): 65-68. BECHYNE, J. 1955. Reise des Herrn Georg Frey in Südamerika: Alticinae (Col. Phytophaga). Entomolo- gische Arbeiten aus dem Museum G. Frey 6: 74-266. BECHYNÉ, J. 1956. Beitráge zur Kenntnis der neotro- pischen Alticiden und Galeruciden. Entomologische Arbeiten aus dem Museum G. Frey 7: 965-1071. BECHYNÉ , J. 1958a. Notizen zu den neotropischen Chry- someloidea (Col. Phytophaga). Entomologische Ar- beiten aus dem Museum G. Frey 9 (2): 478-706. BECHYNE, J. 1958b. Notes sur les Chrysomeloidea néotro- picaux des collections du Musée Zoologique de l’Université et de la ville de Strausbourg (Col. Phyto- phaga). Bulletin de la Société Entomologique de Mul- house 1958 (Mar.-Avr.): 23-24. BECHYNÉ, J. & BECHYNE, B. 1960. Beitráge zur Kenntnis der Salvadorenischen Chrysomeloidea (Col. Phytoph.). Pesquisas Zoologia 6: 5-73. BECHYNE, J. & BECHYNE, B. 1963. Beitráge zur Kenntnis der Salvadorenischen Chrysomeloidea (Col. Phytoph.). Iheringia, Zoologia No. 31: 1-79. BECHYNE, J. & BECHYNÉ, B. 1964. Notes sur quelques Chrysomeloidea Néotropicaux (Col. Phytoph.). Revi- sta Brasileira de Entomologia 11: 117-137. BECHYNE, J. & BECHYNE, B. 1967. Notes sur quelques Al- ticides Néotropicaux (Coleoopt. Phytophaga). Annali del Museo Civico dei Storia Naturale de Genova. 76: 172-195. BLACKWELDER, R. E. 1946. Checklist of the coleopterous insects of Mexico, Central America, the West Indies, and South America. Part 4. Bulletin of the United States National Museum. 185: 551-763. BLAKE, D. H. 1927. A revision of the beetles of the genus Oedionychis occurring in America north of Mexico. Proceedings of the United States National Museum 70 (2672): 1-44. BLAKE, D. H. 1933a. Two new species of Systena, with notes on differences in sexual coloration in the genus. 218 Bonner zoologische Beitráge 54 (2005) Proceedings of the Entomological Society of Washing- ton 32 (8): 180-183. BLAKE, D. H. 1933b. Revision of the beetles of the genus Disonycha occurring in America north of Mexico. Proceedings of the United States National Museum 82 (2969): 1-66. BLAKE, D. H. 1934. New West Indian and Central Ameri- can Chrysomelidae. Bulletin of the Brooklyn Entomo- logical Society 29 (2): 45-56. BLAKE, D. H. 1950. The generic status of Altica picta Say, with descriptions of four closely related species (Col- eoptera: Chrysomelidae). Proceedings of the Entomo- logical Society of Washington 52 (4): 178-183. BLAKE, D. H. 1951. Synonomies and new species of flea- beetles (Coleoptera, Chrysomelidae). Proceedings of the Entomological Society of Washington 53 (3): 138- 147. BLAKE, D. H. 1953. The chrysomelid beetles of the genus Strabala Chevrolat. Proceedings of the United States National Museum 103 (3319): 121-134. BLAKE, D. H. 1955. Revision of the vittate species of the chrysomelid beetle genus Disonvcha from the Ameri- cas south of the United States. Proceedings of the United States National Museum 104 (3338): 1-86. BLAKE, D. H. 1964. Notes on new and old species of Al- ticinae (Coleoptera) from the West Indies. Proceedings of the United States National Museum 115 (3477): 9- 29. CHITTENDEN, F. H. 1923. Notes on the distribution and habits of North American Phyllotreta (Coleopt.). Pro- ceedings of the Entomological Society of Washington 25 (5-6): 131-139. CROTCH, G. R. 1873. Check list of the Coleoptera of America, north of Mexico, Naturalist’s Agency, Sa- lem, Massachusetts, 136 pp. DOMINGUEZ, Y. & CARRILLO, J. L. 1976. Lista de insectos en la coleccion entomologica del Instituto Nacional de Investigaciones Agricolas. Segunda Suplemento. Insti- tuto Nacional Investigaciones Agricolas Folleto Mi- scelaneo No. 29: 1-242. DUCKETT, €. N. 1993. A revision of the genus Pedilia Clark and a cladistic analysis of series Paralactica and the Disonychini (Coleoptera: Chrysomelidae: Altici- nae). Unpublished Ph.D. dissertation. Cornell Univer- sity. 242 pp. DUCKETT, C. N. & KJER, K. M. 2003. Cladistic analysis of the Oedionychines of southern Brazil (Galerucinae: Alticini) based on two molecular markers. Pp. 117-132 in: FURTH, D. G. (ed.) Special Topics in Leaf Beetle Biology: Proceedings of the Fifth International Sym- posium on the Chrysomelidae. Pensoft Publishers, Sofia-Moscow, 339 pp. Fa, J. E. & MORALES, L. M. 1993. Patterns of mammalian diversity in Mexico, Pp. 319-361 in: RAMAMOORTHY, T. P., BYE, R., Lot, A. & FA, J. (eds.) Biological Di- versity of Mexico: Origins and distribution. Oxford Univ. Press, New York. FALL, H. C. 1927. XIII Expedition of the California Acad- emy of Sciences to the Gulf of California in 1921. The Chrysomelidae (Coleoptera). Proceedings of the Cali- fornia Academy of Sciences 16 (13): 381-395. FURTH, D. G. 1998. New World Blepharida Chevrolat 1836 (Coleoptera: Chrysomelidae: Alticinae). Mem- oirs of the Entomological Society of Washington 21: 1-109. FURTH, D. G. 2004. Alticinae (Coleoptera: Chrysomeli- dae). Ch. 29. Pp. 669-684 in: LLORENTE B., J. E., MORRONE, J. J., YÁNEZ, O. & VARGAS F., I. (eds.) Biodiversidad, Taxonomía y Biogeografia de Artrópo- dos de México: Hacia una síntesis de su conocimiento. Volumen IV. Facultad de Ciencias, Universidad Na- cional Autónoma de México, Mexico, D. F. FURTH, D. G., LONGINO, J. & PANIAGUA, M. 2003. Survey and quantitative assessment of flea beetle diversity in a Costa Rican rainforest (Coleoptera: Chrysomelidae: Alticinae). Pp. 1-23 in: FURTH, D. G. (ed.) Special Topics in Leaf Beetle Biology: Proceedings of the Fifth International Symposium on the Chrysomelidae. Pensoft Publishers, Sofia-Moscow, 339 pp. FURTH, D. G. & SAVINI, V. 1996. Checklist of the Altici- nae of Central America, including Mexico (Coleop- tera: Chrysomelidae). Insecta Mundi 10(1-4): 45-68. FURTH, D. G. & SAVINI, V. 1998. Corrections, clarifica- tions, and additions to the checklist of the Alticinae of Central America, including Mexico (Coleoptera: Chrysomelidae). Insecta Mundi 12(1-2): 133-138. GILBERT, A. J. & ANDREWS, F. G. 2002. Studies on the Chrysomelidae (Coleoptera) of the Baja California Peninsula: The genus Dysphenges Horn (Galerucinae: Alticin1). Pan-Pacific Entomologist 75(1): 8-12. GONZALES S., E. & NOVELO G., R. 1996. Odonata. Pp. 147-167. in: LLORENTE B., J., GARCÍA ALDRETE, A. N. & GONZÁLEZ S., E. (eds.) Biodiversidad, Taxonomía y Biogeografía de Artrópodos de México: Hacia una sín- tesis de su conocimiento. Vol. 1. UNAM, Mexico. HAROLD, E. 1876. Diagnosen neuer Arten. Coleopterolo- gischa Hefte 15: 118-124. HEIKERTINGER, F. & CSIKI, E. 1939-1940. Chrysomelidae, Halticinae. Coleopterorum Catalogus. Volume XXV, Pars 166: 1-336, Pars 169: 337-635. Uitgeverij Dr. W. Junk, Gravenhage. Horn, G. H. 1889. A synopsis of the Halticinae of boreal America. Transactions of the American Entomological Society 16: 163-320. Horn, G. H. 1895. Coleoptera of Baja California (Supple- ment 1). Proceedings of the California Academy of Sciences (2) 5: 225-259. JACOBY, M. 1884-1886. Insecta, Coleoptera, Galerucidae. Halticinae. Phytophaga. Biologia Centrali-Americana 6 (1): 263-625. JACOBY, M. 1891-1892. Insecta, Coleoptera, Galerucidae. Halticinae. Phytophaga. Supplement to Phytophaga. Biologia Centrali-Americana 6 (1): 1-374. JACOBY, M. 1902. Descriptions of new species of Coleop- tera of the family Halticidae from South and Central America. Proceedings of the Zoological Society of London 1: 171-204. JacoBy, M. 1904. Descriptions of thirty-two species of Halticinae from South and Central America. Proceed- ings of the Zoological Society of London 2: 396-413 JACOBY, M. 1905. Descriptions of new species of phyto- phagous Coleoptera of the genera Homophoeta, As- phaera, and Oedionychis. Proceedings of the Zoologi- cal Society of London 2: 398-460. LECONTE, J. L. 1859. The Coleoptera of Kansas and east- ern New Mexico. Smithsonian Contributions to Knowledge 11: 1-58. LLORENTE B., J. & LUIS MARTINEZ, A. 1993. Conserva- tion-oriented analysis of Mexican butterflies: Papili- onidae (Lepidoptera: Papilionoidea). Pp. 147-177 in: David G. FURTH: Alticinae (Chrysomelidae) of Mexico 219 RAMAMOORTHY, T. P., BYE, R., LoT, A, & FA, J. (eds.) Biological Diversity of Mexico: Origins and dis- tribution. Oxford Univ. Press, New York. LLORENTE B., J., LUIS MARTINEZ, A., VARGAS F., I. & SOBERON M., J. 1996. Papilionoidea (Lepidoptera). Pp. 531-548 in: LLORENTE B., J., GARCIA ALDRETE, A. N. & GONZALEZ S., E. (eds.) Biodiversidad, Taxono- mia y Biogeografía de Artrópodos de México: Hacia una síntesis de su conocimiento. Vol. 1. UNAM, Mex- ico. MAES, J.-M. & STAINES, C. L. 1991. Catalogo de los Chry- somelidae (Coleoptera) de Nicaragua. Revista Nicara- guense de Entomologia 18: 1-53. MITTERMEIER, R. A. 1988. Primate diversity and the tropi- cal forest: case studies from Brazil and Madagascar and the importance of megadiversity countries. Pp. 145-154 in: WILSON, E. O. (ed.) Biodiversity. Wash- ington, D. C. National Academic Press. MITTERMEIER, R. A., MYERS, N., MITTERMEIER, C. G. & GIL, P. R. 1999. Hotspots. CEMEX. 431 pp. MOCKFORD, E. L. & GARCIA ALDRETE, A. N. 1996. Pscop- tera. Pp. 175-205 in: LLORENTE B., J., GARCIA ALDRE- TE, A. N. & GONZÁLEZ S., E. (eds.) Biodiversidad, Ta- xonomía y Biogeografía de Artrópodos de México: Hacia una síntesis de su conocimiento. Vol. 1. UNAM, Mexico. NÁPOLES, J. R. 2002. Bruchidae. Pp. 513-534 in: LLOR- ENTE B., J. & MORRONE, J. J. (eds.) Biodiversidad, Taxonomia y Biogeografia de Artropodos de México: Hacia una sintesis de su conocimiento. Vol. 3. UNAM, Mexico. O’BRIEN, L. B., MILLER, D. R., MOORE, T. E., PENA M., R. & YANG, M.-M. 1996. Homoptera. Pp. 207-244 in: LLORENTE B., J., GARCÍA ALDRETE, A. N. & GON- ZALEZ S., E. (eds.) Biodiversidad, Taxonomía y Bio- geografía de Artrópodos de México: Hacia una síntesis de su conocimiento. Vol. 1. UNAM, Mexico. PALLISTER, J. C. 1953. The leaf beetles of North Central Mexico collected on the David Rockefeller Mexican Expedition (Coleoptera, Chrysomelidae). American Museum Novitates 1623: 1-95. PARRY, R. H. 1974. Revision of the genus Dibolia Latreille in America north of Mexico (Coleoptera: Chrysomeli- dae). Canadian Journal of Zoology 52 (11): 1317- 1354. PERBOSC, M. 1839. Insects nouveaux découverts au Mexi- que et descrits par M. Perbosc, chirurgien de la marine royale. Revue Zoologique 2: 261-264. REYES-CASTILLO, P. 2002. Passalidae. Pp. 467-483 in: LLORENTE B., J. & MORRONE, J. J. (eds.) Biodiversi- dad, Taxonomía y Biogeografía de Artrópodos de Mé- xico: Hacia una síntesis de su conocimiento. Vol. 3. UNAM, Mexico. RILEY, E. G., CLARK, S. M., FLOWERS, R. W. & GILBERT, A. J. 2002. 124. Chrysomelidae Latreille, 1802. Pp. 617-691 in: ARNETT, R. H. Jr., THOMAS, M. C., SKEL- LEY, P. E. & FRANK, J. H. (eds.) American Beetles, Vol. 2 Polyphaga: Scarabaeoidea through Curculion- oidea. CRC Press, Boca Raton. RILEY, E. G., CLARK, S. M. & GILBERT, A. J. 2001. New records, nomenclatural changes, and taxonomic notes for select North American leaf beetles (Coleoptera: Chrysomelidae). Insecta Mundi 15 (1): 1-17. RILEY, E. G., CLARK, S. M. & SEENO, T. N. 2003. Catalgg of the leaf beetles of America north of Mexico. Special Publication No. 1, The Coleopterists Society, Sacra- mento, 290 pp. RZEDOWSKI, J. 1993. Diversity and origins of the phan- erogamic flora of Mexico. Pp. 129-144 in: RAMA- MOORTHY, T. P., BYE, R., LoT, A, & FA, J. (eds.) Bio- logical Diversity of Mexico: Origins and distribution. Oxford Univ. Press, New York. RZEDOWSKI, J. 1996. Diversidad y orígenes de la flora fa- nerogamica de México. Pp. 27-40 in: LLORENTE B., J., GARCÍA ALDRETE, A. N. & GONZÁLEZ S., E. (eds.) Biodiversidad, Taxonomía y Biogeografia de Artrópo- dos de México: Hacia una síntesis de su conocimiento. Vol. 1. UNAM, Mexico. SAVINI, V. & FURTH, D. G. 1996. Nuevas combinaciones, designacion de lectotipos y redescripciones en Gioia Bechyne (Coleoptera, Chrysomelidae, Alticinae). Bo- letin Entomologia de Venezuela N. S. 11 (2):191-198. SAVINI, V., FURTH, D. G. & NINO-MALDONADO, S. 2001. Una nueva especie de Gioia (Coleoptera: Chrysomeli- dae: Alticinae). Folia entomologica Mexicana 40(1): 11-15 SCHAEFFER, C. 1906. On new and known genera and spe- cies of the family Chrysomelidae. Brooklyn Institute of Museum Sciences Bulletin 1: 221-253 SCHERER, G. 1960. Beitrag zur Kenntnis Alticidenfauna Brasiliens (Col. Phytoph.). Entomologische Arbeiten aus dem Museum G. Frey 11: 180-272. SCHERER, G. 1962. Bestimmungsschluessel der neotro- pischen Alticinen-Genera(Coleoptera: Chrysomelidae: Alticinae). Entomologische Arbeiten aus dem Museum G. Frey 13 (2): 497-607. SCHERER, G. 1983. Diagnostic key for the Neotropical Al- ticinae genera. Entomologische Arbeiten aus dem Mu- seum G. Frey 31/32:2-89. [English translation of Sche- rer, 1962] WILCOX, J. A. 1953. New species of Galerucinae and Al- ticinae with notes on other species (Coleoptera: Chry- somelidae). The Ohio Journal of Science 53 (1): 51- 58. WILCOX, J. A. 1975. Leaf Beetles (Red Version). Checklist of the beetles of Canada, United States, Mexico, Cen- tral America, and the West Indies. BRIA (Biological Research Institute of America), NABFP. 1 (7): 1-166. Author’s address: David G. FURTH, Department of En- tomology, MRC 165, National Museum of Natural His- tory, Smithsonian Institution, P. O. Box 37012, Wash- ington, D. C. 20013-7012, USA. E-mail: furthd@si.edu 220 Bonner zoologische Beitráge 54 (2005) Appendix List of genera and species names, species authors, Mexican states and the sources recorded in the literature, from the United States National Museum of Natural History (USNM) [USNM-DGEF are specimens collected by the author in the USNM collection], or from the F. C. Bowditch Collection at the Museum of Comparative Zoology, Harvard University (FCB). Species names with a “?” afterwards have some question as to the correct determination of this species from Mexico. Species with a “?” after the state acronym means that there is some question as to the certainty of the locality from that state. The state acronyms are standard abbreviations for the 32 Mexican states (see Table 3). When “Mexico ?” occurs in the state column, these are records from the literature for which there is some question about the specimen being from Mexico. Sources are listed only when they represent a new or different state record. Author names for genera can be found in FURTH & SAVINI (1996, 1998). Genus Species Author Distribution Source Acallepitrix castanea (Jacoby) TAB, TAMPS, VER JACOBY (1891) Acallepitrix clypeata (Jacoby) Mexico? FCB Acallepitrix fulvifrons (Jacoby) GRO, MOR FCB; FURTH & SAVINI (1996) Acanthonycha jacobyi Bechyné CHIS, VER BECHYNE € BECHYNE (1960); USNM Acrocyum dorsalis Jacoby COL, NL FURTH & SAVINI (1996); USNM Acrocyum maculicollis Jacoby DGO, MEX, VER FCB; JACOBY (1885); USNM Acrocyum sallei Jacoby OAX JACOBY (1885) Alagoasa acutangula (Jacoby) CHIS, COL, DGO, GRO, — JACOBY (1886); FCB; JAL, MEX, MOR, NL, USNM OAX, VER Alagoasa aemulae (Horn) SON JACOBY (1892) Alagoasa albomarginata (Latreille) GRO HEIKERTINGER & CSIKI (1940) Alagoasa arcuatofasciata (Jacoby) GRO JACOBY (1905) Alagoasa atroguttata (Jacoby) COL, DGO, GRO, JAL, JACOBY (1886); FCB; MOR, SLP, TAMPS, VER USNM Alagoasa bipunctata (Chevrolat) CHIS, DF, OAX, SLP, JACOBY (1886); FCB; D465VER, YUC USNM Alagoasa bipunctata boucardi (Harold) VER? NHMB Alagoasa bipunctata familiaris (Harold) VER? NHMB Alagoasa bipunctata salvini (Harold) CHIS NHMB Alagoasa cazieri Pallister CHIH PALLISTER (1953) Alagoasa ceracollis (Say) CHIS, DGO, MOR, OAX, = JACOBY (1885); VER USNM Alagoasa chevrolati (Baly) OAX, VER, YUC FCB; FURTH & SAVINI (1996) Alagoasa clypeata (Jacoby) CHIS, DGO, HGO, MICH, JAcoBy (1892); OAX, TAB, VER USNM Genus Alagoasa Alagoasa Alagoasa Alagoasa Alagoasa Alagoasa Alagoasa Alagoasa Alagoasa Alagoasa Alagoasa Alagoasa Alagoasa Alagoasa Alagoasa Alagoasa Alagoasa Alagoasa Alagoasa Alagoasa Alagoasa Alagoasa Alagoasa Alagoasa Alagoasa Alagoasa Alagoasa Alagoasa Alagoasa Alagoasa Allochroma Allochroma Allochroma Allochroma David G. FURTH: Alticinae (Chrysomelidae) of Mexico [55] Nw Species Author Distribution Source decemguttatus (Fabricius) CHIH, CHIS, COL, DGO, JACOBY (1886); FCB; GRO, JAL, MEX, MOR, PALLISTER (1953); NAY, OAX, QROO, SON, USNM TAB, VER , / defecta ? (Harold) VER? FCB donckieri (Jacoby) GRO JACOBY (1905) dugesi (Jacoby) GTO JACOBY (1886) duodecimmaculata ? (Jacoby) Mexico? FCB extrema (Harold) MOR, OAX, TAB, VER JACOBY (1886); FCB; USNM fimbriata (Forster) GRO, MICH, MOR, OAX JACOBY (1886); FCB; USNM forreri (Jacoby) DGO JACOBY (1886) frontalis (Jacoby) GRO, NAY JACOBY (1892); NHMB godmani (Jacoby) TAB, VER JACOBY (1886) haroldi (Jacoby) GRO, MOR, VER JACOBY (1892) hoegei (Jacoby) OAX, VER JACOBY (1886) inconspicua (Jacoby) DGO, JAL JACOBY (1886); FCB infirma (Jacoby) OAX, VER JACOBY (1886) jalapa Bechyné VER BECHYNE (1958a) lateralis (Jacoby) COL, GRO, JAL, MEX, JACOBY (1886); FCB; MICH, MOR, NAY, OAX USNM longicollis (Jacoby) OAX JACOBY (1886) millepora (Jacoby) Mexico JACOBY (1905) pavonina (Jacoby) CHIS, VER JACOBY (1892); USNM persimilis (Jacoby) GRO JACOBY (1905) quadrilineata (Harold) Mexico JACOBY (1886) semipurpurea (Jacoby) VER JACOBY (1886) seriata (Baly) GRO, MOR, OAX, PUE, — JACOBY (1886); FCB; VER NHMB tehuacana Bechyné JAL, PUE BECHYNE (1955); USNM tenuilineata (Horn) GRO JACOBY (1892) tridecimmaculata (Jacoby) GRO, MICH JACOBY (1886); JA- COBY (1892) trifasciata escuintla Bechyné Mexico BECHYNE (1958a) trilineata (Jacoby) DF ?, DGO JACOBY (1886) violaceomarginata (Jacoby) OAX JACOBY (1886) virgata (Harold) CHIH, CHIS, COL, DGO, JACOBY (1886); FCB: GRO, JAL, MEX, MOR, PALLISTER (1953): NAY, PUE, OAX, SIN, USNM; NHMB SLP, TAB, VER balyi Clark OAX JACOBY (1886) fasciatum Clark CHIS, VER JACOBY (1886); FCB festivus Clark DGO, VER FCB; USNM flohri Jacoby Mexico JACOBY (1892) Genus Allochroma Allochroma Allochroma Allochroma Allochroma Allochroma Allochroma Altica Altica Altica Altica Altica Altica Altica Altica Altica Altica Altica Altica Altica Altica Altica Altica Altica Altica Altica Altica Altica Altica Altica Altica Aphthona Aphthona Aphthona Argopistes Bonner zoologische Beitráge 54 (2005) Species Author Distribution Source fulvoplagiatum Jacoby VER JACOBY (1886) godmani Jacoby OAX, VER JACOBY (1886) guatemalensis Jacoby CHIS, DGO FCB hoegei Jacoby DGO, OAX, VER JACOBY (1886); FCB mexicanum Jacoby VER JACOBY (1886) semipunctatum Jacoby OAX JACOBY (1886) teapense Jacoby TAB JACOBY (1892) amicula (Jacoby) DGO JACOBY (1891) angulicollis (Jacoby) GRO JACOBY (1891) bimarginata (Say) DGO, GRO, OAX, VER FCB, JACOBY (1884) brisleyi Gentner SON USNM carinata ? Germar SON ? FCB complicata (Harold) DF, DGO, GRO, PUE, JACOBY (1884); FCB; VER USNM cupricollis (Jacoby) DF, DGO, ZAC JACOBY (1891); USNM-DGF flavicollis Jacoby MOR JACOBY (1884) foliacea (LeConte) CHIH, GTO, VER JACOBY (1891); FCB; PALLISTER (1953) forreri (Jacoby) DGO, VER? JACOBY (1884); FCB ignita (Mliger) CHIH, BC, VER JACOBY (1891); PALLISTER (1953) limitata (Jacoby) DGO? FCB litigata Fall CHIS? USNM marevagans ? Horn CHIH, DF, SON JACOBY (1891); FCB nitidiventris Fall BC BLACKWELDER (1946) obliterata LeConte CHIH, SON FCB; JACOBY (1884); PALLISTER (1953) patruelis Harold DF, GRO, GTO, MEX, JACOBY (1884); FCB; MICH, OAX?, PUE, TAB, USNM VER rugicollis Jacoby CHIH, OAX JACOBY (1884); FCB rugosa Jacoby CHIH, GTO, MICH JACOBY (1891) satellitia (Jacoby) DGO, GRO, VER JACOBY (1891); FCB schwarzi Blatchley Mexico? USNM (? Mexico, ex banana shipment) tincta LeConte BC BLACKWELDER (1946) tombacina Mannerheim CHIH, DGO PALLISTER (1953) torquata LeConte DF, SON JACOBY (1884); USNM pilatei Baly TAB JACOBY (1885) smithi Jacoby GRO JACOBY (1891) unicolor Jacoby TAB JACOBY (1891) rubicunda _ Blake VER BLAKE (1934) David G. FURTH: Alticinae (Chrysomelidae) of Mexico Genus Species Asphaera abdominalis Asphaera cyanopsis Asphaera icteridera Asphaera lustrans Asphaera mexicana Asphaera nigrofasciata Asphaera nobilitata Asphaera polita Asphaera reichei Asphaera xanthocephala Ayalaia minor Blepharida alternata Blepharida alticola Blepharida atripennis Blepharida balyi Blepharida bryanti Blepharida conspersa Blepharida flavocostata Blepharida flohri Blepharida gabrielae Blepharida godmani Blepharida hinchahuevosi Blepharida humeralis Blepharida jacobyana Blepharida Johngi Blepharida judithae Author (Chevrolat) Harold (Harold) (Crotch) (Harold) Jacoby (Fabricius) Jacoby (Harold) Harold Bechyné & Bechyné Jacoby Achard Horn Bryant Furth (Horn) Jacoby Jacoby Furth Jacoby Furth Furth Csiki Furth Furth Distribution Source AGS, CHIH, CHIS, JACOBY (1885 COAH, COL, DF, DGO, (1892)); FCB; GRO, GTO, HGO, JAL, PALLISTER (1953); MEX, MICH, MOR, NL, USNM OAX, SIN, SLP, TAMPS, VER, ZAC DF, DGO, OAX, SLP, TAB, VER CHIS, DGO, GRO, MOR, OAX, VER BC CHIS, DGO, GRO, MICH, MOR, NAY, OAX, VER VER CHIS OAX, TAB, VER CHIS, DF, OAX, SLP, VER VER GRO?, NAY CHIS, GRO, JAL, MOR, NAY, SLP, SIN, SON CHIS, DGO BCS GRO, MEX, MICH, MOR CHIS, OAX BCS GRO, MEX, MICH, MOR, OAX, PUE GRO, JAL, MEX, MICH, MOR, NAY COL, GRO, JAL, MEX, MICH. MOR, PUE CHIS, OAX, VER GRO, MOR, PUE GRO, MOR, PUE CHIS, QROO, VER, YUC DGO, GRO, MEX, MICH GRO JACOBY (1885); FCB; USNM JACOBY (1885); PALLISTER (1953); USNM BLACKWELDER (1946) JACOBY (1886); FCB; USNM USNM USNM JACOBY (1885); FCB JACOBY (1885); USNM FURTH & SAVINI (1996) USNM FURTH (1998) FURTH (1998) FURTH (1998) FURTH (1998) FURTH (1998) FURTH (1998) FURTH (1998) FURTH (1998) FURTH (1998) FURTH (1998) FURTH (1998) FURTH (1998) ) FURTH (1998 FURTH (1998) FURTH (1998) 224 Bonner zoologische Beitráge 54 (2005) Genus Species Author Distribution Blepharida lineata Furth GRO, MICH Blepharida maculicollis Jacoby MOR, PUE Blepharida melanoptera (Fall) MICH, OAX, SON Blepharida mexicana Jacoby OAX, VER Blepharida multimaculata Jacoby GRO, MEX, MOR, PUE, SON Blepharida notozonae Furth JAL, SIN, SON Blepharida pallida Blake GRO, JAL, MICH, SIN Blepharida parallela Furth MEX, MICH, PUE Blepharida punctatissima Jacoby CHIS, OAX, VER Blepharida quatuordecimpunctata Jacoby CHIS, OAX, VER Blepharida rhois (Forster) CHIH, COAH, DGO, GRO, HGO, NL, OAX, PUE, QRO, SLP, TAMPS Blepharida schlectendalii Furth PUE Blepharida singularis Jacoby DGO, GRO, MEX, MICH, MOR, PUE Blepharida sonorana Furth SIN, SON Blepharida sonorstriata Furth SON Blepharida sparsa (Clark) CHIS, GRO, MICH Blepharida trifasciata Jacoby OAX Blepharida unami Furth OAX, PUE Blepharida variegatus Furth PUE Blepharida verdea Furth GRO, MOR, OAX Blepharida xochipala Furth GRO Cacoscelis bicolorata Clark Mexico Cacoscelis coerulea Csiki TAB Cacoscelis flava Clark OAX, TAMPS Cacoscelis sallei Jacoby VER Cacoscelis varians (Jacoby) OAX, TAB, VER Caloscelis nigripennis (Jacoby) NAY Capraita conspurcata (Jacoby) CHIS, DF, DGO, GRO, GTO, HGO, MEX, MICH, MOR, OAX, PUE, VER Capraita maculata (Harold) CHIS, GRO, JAL, MEX, MOR, OAX, VER, YUC Capraita maculata (Jacoby) GRO, HGO, MEX tredecimmaculata Capraita nigrosignata (Schaeffer) Mexico ? Centralaphthona deyrollei (Baly) JAL, MOR, TAB Centralaphthona dimidiaticornis Jacoby GRO, TAB Source FURTH (1998) FURTH (1998) FURTH (1998) FURTH (1998) FURTH (1998) FURTH (1998) FURTH (1998) FURTH (1998) FURTH (1998) FURTH (1998) FURTH (1998) FURTH (1998) FURTH (1998) FURTH (1998) FURTH (1998) FURTH (1998) FURTH (1998) FURTH (1998) FURTH (1998) FURTH (1998) FURTH (1998) JACOBY (1884) JACOBY (1892) JACOBY (1884); USNM JACOBY (1884) JACOBY (1891) JACOBY (1891) JACOBY (1886); FCB; USNM JACOBY (1886); FCB USNM; NHMB USNM (? Mexico, ex snapdragon shipment) JACOBY (1885): USNM JACOBY (1891); FCB David G. FURTH: Alticinae (Chrysomelidae) of Mexico Genus Species Centralaphthona diversa Centralaphthona fulvipennis ? Centralaphthona fulvipennis near Centralaphthona maculipennis Centralaphthona mexicana Centralaphthona obscuripennis Centralaphthona pallipes Centralaphthona semicoerulea Centralaphthona semipuncata Centralaphthona substriata Chaetocnema balyi Chaetocnema capitata Chaetocnema cephalotes Chaetocnema confinis Chaetocnema costatipennis Chaetocnema cribrifrons Chaetocnema discoidalis Chaetocnema ectypa Chaetocnema frontalis ? Chaetocnema fulvicornis Chaetocnema fulvida Chaetocnema fulvilabris Chaetocnema gravida Chaetocnema horni Chaetocnema interstitialis Chaetocnema mexicana Chaetocnema parcepunctata Chaetocnema sallei Chaetocnema smithi Chalatenanganya frontalis Chrysogranma octomaculata Chrysogramma omiltenaia Chrysogramma pictipennis Chrysogramma septempunctata Chrysogramma trifasciata Cornulactica varicornis Crepidodera opulenta ? Author (Baly) Jacoby Jacoby Jacoby Jacoby (Jacoby) Jacoby (Jacoby) Jacoby Harold Jacoby Jacoby Jacoby Crotch Jacoby LeConte Jacoby Horn Jacoby Jacoby White Jacoby Baly Jacoby Jacoby Baly Crotch Baly Jacoby (Jacoby) Jacoby Jacoby Jacoby Jacoby Jacoby (Jacoby) (LeConte) Distribution Source CHIS, GRO, MOR, TAB, TAMPS? VER? TAMPS TAB COAH, DGO, GRO GRO, MOR CHIH, GRO, VER DGO JAL, VER GRO, TAB, VER COAH, DF DGO, GTO PUE, SIN DF GTO CHIH, DGO Mexico CHIH ? Mexico DGO, GRO, GTO BCS GRO, MOR, VER JAL, TAB TAB GRO CAMP, COAH, DF, DGO, MOR, TAB DF Mexico TAB CHIS CHIS, JAL, VER GRO DGO DGO, MOR, OAX, PUE OAX VER BC, MOR JACOBY (1885); FCB; USNM FCB, USNM-El Cielo JACOBY (1891) JACOBY (1885) (1891); FCB USNM JACOBY (1891); USNM-DGF JACOBY (1885) JACOBY (1891); FCB JACOBY (1891) JACOBY (1892); FCB JACOBY (1885); FCB NHMB USNM JACOBY (1892) PALLISTER (1953) JACOBY (1885) USNM (ex mustard shipment from Juarez) / FCB JACOBY (1885); FCB USNM JACOBY (1892); USNM JACOBY (1885): FCB JACOBY (1892) JACOBY (1892) JACOBY (1885); FCB; USNM JACOBY (1892) JACOBY (1885) JACOBY (1892) NEW — DGF: USNM DOMINGUEZ & CARRILLO (1976); USNM JACOBY (1891) JACOBY (1891) USNM; FURTH & SAVINI (1996) JACOBY (1891) JACOBY (1891) USNM 226 Bonner zoologische Beitráge 54 (2005) Genus Species Author Distribution Source Crepidodera peninsularis Horn BC FURTH & SAVINI (1998) Cyrsylus fulvipes Jacoby TAB JACOBY (1892) Cyrsylus recticollis Jacoby CHIS, TAB, VER JACOBY (1892); USNM Cyrsylus vittatus Jacoby CHIS, GRO, GTO, JAL JACOBY (1892); USNM Deuteraltica longicornis (Jacoby) CHIS USNM Dibolia championi Jacoby OAX, VER PARRY (1974); USNM Dibolia constricta Jacoby DGO, VER JACOBY (1891) Dibolia ovata LeConte DF, DGO, GRO, GTO, JACOBY (1891); FCB; MICH, VER USNM Dibolia violacea Jacoby GRO JACOBY (1891) Dinaltica chevrolati (Jacoby) VER JACOBY (1884) Diphaltica chiriquiensis ? (Jacoby) VER FCB; USNM Diphaltica columbica (Harold) VER JACOBY (1884) Diphaltica crassicornis ? (Jacoby) TAB, VER FCB Diphaltica fossifrons (Harold) VER JACOBY (1891) Diphaltica nitida (Jacoby) CHIS, DF, DGO, MICH, JACOBY (1884); FCB; OAX, TAB, VER USNM Diphaltica panamensis ? (Jacoby) DGO FCB Diphaltica sallei (Harold) Mexico HAROLD (1876) Diphaltica sobrina Jacoby VER JACOBY (1884) Diphaulaca aulica cordobae Barber CHIS, GRO, GTO, HGO, — JACOBY (1884); FCB; JAL, MEX, MICH, MOR, USNM NAY, OAX, PUE, QROO, SLP?, TAB, TAMPS, VER, YUC Diphaulaca intermedia Jacoby GRO JACOBY (1891) Diphaulaca wagneri Harold CHIS, GRO, OAX, YUC BARBER (1941); NHMB Disonycha discoidea abbreviata Melsheimer DGO, MEX, MOR, OAX JACOBY (1884); FCB Disonycha angulata Jacoby SLP, TAB, VER, YUC JACOBY (1891) Disonycha annulata Blake Mexico BLAKE (1955) Disonycha antennata Jacoby COL, DGO, GRO, JAL, JACOBY (1884); BLA- MEX, MICH, MOR, OAX, KE(1955); USNM VER Disonycha apicalis Jacoby VER JACOBY (1884) Disonycha arizonae Casey CHIH, DGO?, GRO, FURTH & SAVINI MOR? (1996); USNM Disonycha barberi Blake GRO, SIN, VER BLAKE (1955); USNM Disonycha brevicollis Jacoby DGO JACOBY (1902) Disonycha brevilineata Jacoby DGO, GRO, JAL, MOR, JACOBY (1884); JA- OAX COBY (1902); FCB; BLAKE (1955) David G. FURTH: Alticinae (Chrysomelidae) of Mexico 227 Genus Species Author Distribution Source Disonycha brunneofasciata Jacoby GRO}PUE,. SLP BLAKE (1955); USNM Disonycha caroliniana (Fabricius) DGO, NL, OAX, SIN, VER JACOBY (1884); USNM Disonycha collata (Fabricius) CHIH, COAH, DF, DGO, = JACOBY (1884); FCB: GTO, JAL, MEX, MICH, — PALLISTER (1953); MOR, OAX, PUE, TAB, USNM VER, YUC Disonycha crenicollis Say PUE, SON JACOBY (1884) Disonycha dorsata Harold MOR, OAX, TAB, VER, JACOBY (1884); FCB; YUC USNM Disonycha figurata Jacoby AGS, CHIA, CHIS, JACOBY (1884); FCB; COAH, COL, DF,DGO, — PALLISTER (1953); GRO, GTO, JAL, MEX, BLAKE (1955); MICH, MOR, NAY, OAX, USNM; NHMB SIN, TAB, VER, YUC Disonycha fumata fumata LeConte BC, CHIH, CHIS, DGO, BLAKE (1955); GRO, HGO, JAL, MEX, USNM MICH, MOR, NL, OAX, PUE, SLP, SON, TAB, VER, ZAC Disonycha fumata quinquerutata Schaeffer CHIH PALLISTER (1953) Disonycha glabrata (Fabricius) BC, BCS, CAMP, CHIS, JACOBY (1884); FCB; COL, DGO, GRO, JAL, BLAKE (1955); MOR, NAY, OAX, PUE, USNM SIN, SON, TAB, TAMPS, YUC, VER Disonycha gracilis Blake VER BLAKE (1955) Disonycha guatemalensis Jacoby CHIS, GRO, MOR, OAX, BLAKE(1955); VER? USNM Disonycha hoegei Jacoby VER, OAX JACOBY (1884) Disonycha jalapensis Blake VER BLAKE (1955) Disonycha knabi Blake GRO BLAKE (1955) Disonycha latifrons Schaeffer TAMPS? USNM Disonycha latiovittata Hatch & Beller BC BLAKE (1955) Disonycha leptolineata texana Schaeffer DGO, GRO, JAL, MOR, BLAKE (1955); NL, OAX, QROO, USNM TAMPS, YUC Disonycha limbata Jacoby DGO, MEX, MOR JACOBY (1891); USNM Disonycha maculipes Jacoby CHIS, VER JACOBY (1891); USNM Disonycha melanocephala Jacoby VER JACOBY (1884) 228 Genus Species Disonycha Disonycha Disonycha Disonycha Disonycha Disonycha Disonycha Disonycha Disonycha Disonycha Disonycha Disonycha Disonycha Disonycha Disonycha Disonycha Disonycha Disonycha Disonycha Distigmoptera Distigmoptera Distigmoptera Distigmoptera mexicana militaris nigripennis nigripes ? pluriligata politula procera quinquelineata recticollis sallei scriptipennis steinheili subaenea subcostata teapensis tenuicornis trifasciata trivittata varicornis chrysodaedala foveolata orchidophila suturalis Bonner zoologische Beitráge 54 (2005) Author Jacoby Jacoby Jacoby Jacoby LeConte Horn Casey (Latreille) (Jacoby) (Baly) (Jacoby) Harold Jacoby (Clark) Blake Horn Jacoby Blake Horn Blake Balsbaugh Blake (Jacoby) Distribution Source TAB, SLP, VER, YUC TAB, VER, YUC DGO, NAY CHIS BC, CHIH, DGO, JAL, NAY, SIN, SLP, SON, VER AGS, CAMP, CHIH, DF, DGO, GRO, GTO, HGO, JAL, MEX, MOR, OAX, PUE, QRO, SLP, SON, TAMPS, VER, ZAC NAY CHIS; COL, GRO, OAX, QROO, TAB, TAMPS, VER NAY, VER VER CHIS, COL, DGO, GRO, MOR, NAY, OAX, YUC Mexico DGO, GRO, MOR, OAX HGO, MEX, MICH OAX, SLP, TAB CHIH, DGO, HGO CHIS Mexico BC, BCS, DGO, TAMPS DF, NAY CHIS, PUE VER GRO, OAX FCB; JACOBY (1884) JACOBY (1884); USNM JACOBY (1884); USNM FCB FCB; FURTH & SAVINI (1996) FCB; JACOBY (1891); PALLISTER (1953): USNM BLAKE (1955) JACOBY (1884); FCB; BLAKE (1955); USNM JACOBY (1884); ZSMC JACOBY (1884) JACOBY (1891); USNM; NHMB ZSMC JACOBY (1884); FCB; USNM JACOBY (1884); USNM BLAKE (1955); NHMB PALLISTER (1953); BLAKE (1955) FCB BLAKE (1955) FURTH & SAVINI (1998); PALLISTER (1953); USNM BLAKE (1951); USNM BALSBAUGH (1980); USNM BLAKE (1951) JACOBY (1892); NHMB Genus Distigmoptera Dysphenges Dysphenges Dysphenges Dysphenges Egleraltica Epitrix Epitrix Epitrix Epitrix Epitrix Epitrix Epitrix Epitrix Epitrix Epitrix Epitrix Epitrix Epitrix Epitrix Epitrix Epitrix Epitrix Euphenges Euplectroscelis Exoceras Genaphthona Genaphthona Gioia Gioia Gioia Gioia David G. FURTH: Alticinae (Chrysomelidae) of Mexico 229 Species Author Distribution Source texana Blake GTO BLASBAUGH (1980); FURTH & SAVINI (1998) eichlini Gilbert & An- BCS GILBERT & ANDREWS drews (2002) f elongatulus Horn BCS, MICH”, PUE?, GILBERT & ANDREWS TAMPS? (2002); USNM lagunae Gilbert & An- BCS GILBERT & ANDREWS drews (2002) rileyi Gilbert & An- BCS GILBERT & ANDREWS drews (2002) sp. CHIS, GRO, MICH, MOR USNM aenicollis Jacoby GRO, TAB, VER JACOBY (1891) convexa Jacoby TAB, VER JACOBY (1891) cucumeris (Harris) DGO, GRO, MOR, PUE, JACOBY (1891); FCB: VER USNM; ZSMC fasciata Blatchley CHIH, DGO, NL, TAMPS MAES & STAINES (1991); USNM flavotestacea Horn BC BLACKWELDER (1946) fuscula Crotch GTO? JACOBY (1885) jacobyi Weise GTO, VER JACOBY (1885) metallica Jacoby GRO JACOBY (1885) minuta Jacoby VER, TAB FCB, JACOBY (1891) montana Jacoby VER, COAH JACOBY (1885) nigroaenea Harold VER JACOBY (1885) obliterata Jacoby TAB, VER JACOBY (1891); FCB parvula (Fabricius) CHIH, GTO, VER FCB, JACOBY (1891); USNM-DGF piceomarginata Jacoby TAB JACOBY (1891) pulchella Jacoby TAB, VER FCB, JACOBY (1885) robusta Jacoby GRO JACOBY (1891) rufula Weise DF, GRO, MOR JACOBY (1891); USNM fulginosus (Clark) Mexico JACOBY (1886) xanti Crotch BC; BCS BLACK WELDER (1946); USNM sp. CHIS USNM amulensis Jacoby GRO JACOBY (1891) transversicollis (Jacoby) CHIS, JAL, OAX, PUE USNM fulvitarsis Jacoby TAB JACOBY (1891) mexicana Savini, Furth, TAMPS SAVINI et al. (2001) Nino sp. 1 TAMPS SAVINI et al. (2001) sp. 2 TAMPS SAVINI et al. (2001) 230 Bonner zoologische Beiträge 54 (2005) Genus Species Author Glenidion flexicaulis Schaeffer Glenidion jacobyi (Bechyné) Glyptina atriventris Horn Glyptina cerina (LeConte) Glyptina nivialis Horn Heikertingerella fulvifrons (Jacoby) Heikertingerella pallida Jacoby Heikertingerella teapensis (Weise) Heikertingerella variabilis (Jacoby) Heikertingeria clarki (Jacoby) Hemiphvrnus intermedius (Jacoby) Homotyphus asper (Clark) Homotvphus maculicornis (Clark) Homotyphus squalidus ? Clark Hypolampsis elongatula (Jacoby) Hypolampsis inornata Jacoby Hypolampsis labialis (Clark) Hypolampsis pygmaea (Jacoby) Iphitroides nigrocinctus Jacoby Iphitroides quadrimaculatus Jacoby Iphitroides quadripunctatus Jacoby Iphitroides violaceipennis Jacoby Kuschelina gracilis (Jacoby) Kuschelina laeta (Perbosc) Kuschelina lugens (LeConte) Kuschelina miniata ? (Fabricius) Kuschelina modesta (Jacoby) Kuschelina petaurista ? (Fabricius) Kuschelina violascens (LeConte) Leptophysa hirtipennis (Jacoby) Longitarsus amulensis Jacoby Longitarsus antennatus Jacoby Longitarsus bicolor Horn Longitarsus buckleyi ? Baly Distribution Source TAMPS, YUC USNM CHIS, MICH, VER JACOBY (1885); USNM CHIH, MICH USNM BC BLACKWELDER (1946) MOR USNM GRO, VER JACOBY (1891) VER JACOBY (1891) TAB JACOBY (1892) GRO, GTO, TAB, TAMPS, JACOBY (1885); FCB; VER USNM VER USNM CHIS, DGO, SON FCB, JACOBY (1884) VER JACOBY (1886) VER USNM; FURTH & SAVINI (1996) VER JACOBY (1886) GRO JACOBY (1892) GRO JACOBY (1892) VER JACOBY (1886) TAB, VER JACOBY (1892) GRO JACOBY (1891) DGO, GRO JACOBY (1891): USNM GRO, MOR, NAY JACOBY (1891); USNM Mexico? JACOBY (1891) COL, JAL, MOR, SIN USNM; NHMB TAMPS, VER HEIKERTINGER & CSIKI (1940); FCB; USNM Mexico? USNM Mexico FCB CHIH, CHIS, DF, DGO, JACOBY (1886); FCB; GRO, GTO, HGO, MEX, — PALLISTER (1953); MOR, OAX, PUE, SLP, USNM TLAX, VER COAH USNM DGO FURTH & SAVINI (1998); PALLISTER (1953) OAX, VER USNM GRO JACOBY (1891) VER JACOBY (1891) BC BLACKWELDER (1946); FURTH & SAVINI (1998) TAB FCB Genus Longitarsus Longitarsus Longitarsus Longitarsus Longitarsus Longitarsus Longitarsus Longitarsus Longitarsus Luperaltica Luperaltica Luperaltica Lupraea Lupraea Lupraea Lupraea Lupraea Lupraea Lupraea Lupraea Lupraea Lupraea Lupraea Lysathia Lysathia Lysathia Lysathia Macrohaltica Macrohaltica Macrohaltica Macrohaltica Macrohaltica David G. FURTH: Alticinae (Chrysomelidae) of Mexico Species Author Distribution Source columbicus ? Harold GRO FCB concinnus Baly Mexico? JACOBY (1885) haroldi Jacoby TAB JACOBY (1891) livens LeConte BC BLACKWELDER (1946) mexicanus Csiki DF, DGO, GRO, GTO, JACOBY (1891); FCB; HGO, MEX, MICH, MOR, NHMB; USNM PUE ovipennis Jacoby GRO JACOBY (1891) repandus LeConte BC BLACKWELDER (1946) teapensis Jacoby TAB JACOBY (1891) varicornis Suffrian TAB, VER JACOBY (1885); JA- COBY (1891) longicornis (Jacoby) CHIS, COL?, MOR?, USNM OAX? ustulata centralis (Bechyné)+C254 NAY, TAMPS BECHYNE (1960); USNM viridipennis (Jacoby) OAX JACOBY (1884) championi Jacoby VER JACOBY (1885) dilaticornis (Jacoby) VER JACOBY (1891) elongata (Jacoby) GRO, MICH USNM frontalis (Jacoby) OAX JACOBY (1885); USNM fulvicollis ? Jacoby VER FCB godmani (Jacoby) TAB, VER? JACOBY (1891); USNM guatemalensis (Jacoby) CHIS, GRO, MOR, VER JACOBY (1891); USNM imitans (Jacoby) GRO, VER? JACOBY (1891); USNM occipitalis Bechyné & Mexico? BECHYNE & BECHYNE Bechyné (1964) semifulva (Jacoby) CHIS, OAX USNM-DGF smithi (Jacoby) GRO, MOR JACOBY (1891); USNM-DGF jacobyi (Csik1) DF, GTO, OAX, TAB, JACOBY (1891); ZAC USNM ludoviciana (Fall) Mexico? USNM (on banana: Galveston) occidentalis (Suffrian) YUC ZSMC rockefelleri (Pallister) CHIH, DGO PALLISTER (1953) amethystina (Olivier) PUE, VER JACOBY (1884) guatemalensis (Jacoby) DF, GRO, SON JACOBY (1891); FURTH & SAVINI (1998); USNM jamaicensis (Fabricius) TAMPS USNM mexicana mexicana (Jacoby) DF, DGO, MOR JACOBY (1884); FCB; USNM mexicana Bechyné CHIS, DF, DGO USNM salvadorensis 232 Genus Species Macrohaltica Margaridisa Margaridisa Mesodera Monomacra Monomacra Monomacra Monomacra Monomacra Monomacra Monomacra Monomacra Monomacra Monomacra Monomacra Monomacra Monomacra Monomacra Monomacra Monomacra Monomacra Neodiphaulaca Neothona Nesaecrepida Nesaecrepida Notozona Notozona Notozona Omophoita Omophoita Omophoita Omophoita Omophoita Omophoita patruelis atriventris managua ? brevicollis abdominalis binotata crassicornis cupreata elongata gracilicornis hidalgoensis hoegei inornata ? mexicana nicotinae pusilla salvini semiviolacea tibialis violacea violaceipennis elongatula sp. infuscata asphaltina elegans histrionica humilis aequatorialis aequinoctialis aequinoctialis albofasciata affinis ? championi cinctipennis Author Bonner zoologische Beiträge 54 (2005) Distribution (Harold) (Melsheimer) (Bechyné) Jacoby (Jacoby) (Baly) (Jacoby) (Jacoby) (Jacoby) (Olivier (Jacoby (Jacoby ( ) ) ) Harold) (Schaeffer) (Suffrian) Clark Baly Clark (Harold) (Linnaeus) (Jacoby) (Jacoby) (Jacoby) . (Chevrolat) DF, DGO, GTO, MEX, MICH, MOR, OAX, PUE, VER CHIS, VER DGO, SLP HGO CAMP, YUC TAB, VER CHIS, GRO OAX DGO Mexico HGO OAX, VER Mexico OAX, VER TAMPS, VER VER CHIH TAMPS OAX CHIS, VER Mexico ? DGO, SLP, VER JAL, MICH, OAX, VER CAMP, COL, GRO, JAL, MICH, OAX, TAB, TAMPS, VER YUC Mexico OAX, VER Mexico Mexico CHIS, HGO, MICH, OAX, QROO, SLP, TAB, TAMPS, VER TAB Mexico ? SON JAL, OAX, PUE, SLP, VER Source JACOBY (1884); USNM SNM SNM ACOBY (1892) SNM ACOBY (1884); SNM JACOBY (1891) JACOBY (1891) JACOBY (1884) JACOBY (1902) JACOBY (1892) JACOBY (1884) FCB JACOBY (1884): USNM JACOBY (1904); USNM JACOBY (1884) FCB JACOBY (1884); USNM USNM USNM-DGF JACOBY (1884) HEIKERTINGER & CSIKI (1939), FCB; USNM USNM Sc=-cc El USNM; USNM-DGF; ZSMC USNM JACOBY (1885) FCB; FURTH & SAVINI (1996) JACOBY (1885) JACOBY (1885) FURTH & SAVINI (1996): USNM FCB FCB DOMINGUEZ & CARRILLO (1976) JACOBY (1885): USNM Genus Omophoita Omophoita Omophoita Omophoita Omophoita Omophoita Omophoita Palaeothona Palaeothona Palaeothona Palaeothona Palaeothona Paranaita Parasyphraea Parchicola Parchicola Pedilia Phenrica Phenrica Phrynocepha Phrynocepha Phrynocepha Phrynocepha Phrynocepha Phrynocepha Phrynocepha Phydanis Phydanis Phyllotreta David G. FURTH: Alticinae (Chrysomelidae) of Mexico 233 Species Author Distribution Source clerica (Erichson) CHIS USNM octomaculata (Crotch) OAX, TAB, TAMPS, VER JACOBY (1886); ZSMC punctulata (Bechyné & Mexico BECHYNE & BECHYNE Bechyné) (1963) y quadrinotata Bechyné OAX, TAB, VER BECHYNE (1955); centraliamericana USNM recticollis (Baly) CHIS, HGO, OAX, TAB, JACOBY (1885) TAMPS, VER (1891); USNM simulans (Jacoby) CHIS, DGO JACOBY (1892) violacea Jacoby GRO JACOBY (1892) discrepans (Schaeffer) CHIH USNM-DGF melanocyanea (Blake) HGO, TAMPS BLAKE (1950); FURTH & SAVINI (1998): USNM rubroviridis Blake DGO BLAKE (1950) rugifrons (Jacoby) VER JACOBY (1885) viridis (near) (Jacoby) CHIH NEW — DGF (PDST) limbatipennis (Jacoby) GRO? ZSMC minuta (Jacoby) TAB, YUC JACOBY (1891); USNM uniformis ? (Jacoby) Mexico FCB variabilis (Jacoby) CHIS, VER JACOBY (1884); USNM inornata (Jacoby) OAX, VER DUCKETT (1993) in litt. cordovana (Jacoby) VER JACOBY (1884) sexmaculata (Jacoby) VER JACOBY (1884) capitata Jacoby CHIS?, GRO, JAL, OAX, JACOBY (1884); TAB? USNM deyrollei Baly AGS, CHIH, DGO, GRO, — JACOBY (1884); GTO, MICH, MOR, OAX, PALLISTER (1953); PUE, SLP ? USNM elongata Jacoby OAX, TLAX, VER JACOBY (1884) pulchella Baly CHIS, COL, DGO, GTO, — JACOBY (1884); JAL, MICH, MOR, OAX, USNM;NHMB VER punctulata Pallister CHIH, DGO PALLISTER (1953) sulcatipennis (Jacoby) GRO, MEX, OAX JACOBY (1891); NHMB tenuicornis (Jacoby) HGO, OAX FCB; JACOBY (1891) bicolor Horn OAX, TAMPS USNM nigriventris Jacoby GRO, OAX, SLP, SON JACOBY (1891); USNM albionica (LeConte) BC, CHIH, SON FALL (1927); FURTH & SAVINI (1998); USNM 234 Bonner zoologische Beitráge 54 (2005) Genus Species Author Distribution Phyllotreta crotchi Jacoby DF, DGO, HGO, MEX, ZAC Phyllotreta lativittata Jacoby DF, DGO, GTO: HGO, MEX, ZAC Phyllotreta mexicana Jacoby VER Phyllotreta pusilla Horn AGS, BC?, CHIH, DF, DGO, HGO, MOR, OAX, ZAC Phyllotreta subrugosa Jacoby DF Physimerus cordovensis (Jacoby) VER Physimerus femoralis (Jacoby) VER Physimerus scabrosus (Clark) DGO, OAX, VER Platiprosopus pallens (Fabricius) GRO, HGO, MOR, OAX, PUE, VER Plectrotetra clarki Baly DF, DGO, HGO, MOR, DAX, PUE; SIN, SLP; TAMPS, VER Plectrotetra dohrni Jacoby DE, DGO, EGO, PUE, VER, YUC Plectrotetra flohri Jacoby MICH Plectrotetra guatemalensis Jacoby MOR Plectrotetra inaequalis Jacoby OAX, TAMPS, VER Plectrotetra multipunctata Jacoby DGO?, MEX, MOR, OAX, VER Plectrotetra rugosa Jacoby GTO Plectrotetra sallei Jacoby CHIS, TAB? Plectrotetra submetallica Jacoby OAX, VER Prasona viridis Baly VER Propiasus fulvus (Jacoby) GRO Pseudodibolia picea Jacoby TAB Pseudorthygia n. sp. CHIS Pseudorthygia nigritarsis Jacoby GRO, OAX, TAMPS Pseudorthygia unifasciata Jacoby GRO Psylliodes capitata Jacoby GRO Psylliodes convexior LeConte BCS Psylliodes melanocephala Jacoby DF Psylliodes mexicana Jacoby DF Psylliodes sublaevis Horn GRO Resistenciana ornata (Jacoby) PUE, VER Rhinotmetus depressus Clark Mexico ? Rhinotmetus flavovittatus ? Jacoby Mexico ? Rhinotmetus minutus Jacoby VER Rhinotmetus modestus Jacoby GRO, MOR Source JACOBY (1885) (1891); USNM JACOBY (1891): FCB; USNM JACOBY (1885) CHITTENDEN (1923); USNM, USNM-DGF JACOBY (1891) JACOBY (1886) USNM JACOBY (1886); FCB FURTH & SAVINI (1996); USNM JACOBY (1884); FCB; USNM JACOBY (1884); FCB; USNM JACOBY (1884) JACOBY (1891) JACOBY (1884); USNM JACOBY (1891); FCB: USNM JACOBY (1884) JACOBY (1884); FCB; USNM JACOBY (1884) JACOBY (1886) JACOBY (1892) JACOBY (1891) USNM-DGF JACOBY (1891): USNM, USNM-DGF JACOBY (1891) JACOBY (1892) HORN (1895); FURTH & SAVINI (1998) JACOBY (1891) JACOBY (1891) JACOBY (1891) JACOBY (1884): FCB JACOBY (1886) FCB JACOBY (1892) JACOBY (1892); FCB David G. FURTH: Alticinae (Chrysomelidae) of Mexico Genus Species Author Distribution Source Scelidopsis rufofemorata Jacoby TAMPS, VER JACOBY (1888); USNM Scelidopsis violacea ? Jacoby GRO JACOBY (1892) Sphaeronychus fulvus (Baly) DGO, TAMPS JACOBY (1886): USNM : Sphaeronychus puncticollis (Jacoby) GRO JACOBY (1892) Stegnea amplicollis (Jacoby) GRO JACOBY (1891) Stegnea atra (Jacoby) GRO JACOBY (1891) Stegnea guatemalensis (Jacoby) GTO FCB Stegnea obliterata (Jacoby) TAB JACOBY (1891) Strabala acuminata teapensis Blake CHIS, TAB BLAKE (1953); USNM Strabala durangoensis Bechyné DGO BECHYNE (1955) Strabala rotunda Blake CHIS.COL, DE, GRO, BLAKE (1953); JAL, NAY., NL, SLP; USNM; NHMB, TAMPS, VER, YUC ZSMC Strabala rufa Illiger CHIS, COL, DGO, GRO, — JACOBY (1884, 1891) OAX, PUE, TAB, VER Strabala subcostata (Jacoby) VER JACOBY (1884) Suetes niger Jacoby GRO, MICH JACOBY (1891); USNM Syphrea abdominalis (Jacoby) TAB, VER JACOBY (1891) Syphrea aeneipennis (Jacoby) CHIS?, VER USNM-DGF Syphrea angustata Jacoby COAH, GRO, PUE, VER JACOBY (1891) Syphrea balnearia Bechyné & GRO, MEX USNM-DGF Bechyné Syphrea burgessi (Crotch) MOR, OAX, TAMPS USNM Syphrea cyaneipennis (Jacoby) GRO; HGO, JAL, SEP, JACOBY (1891); TAB, TAMPS USNM Syphrea flavicollis (Jacoby) BCS, GRO, GTO, JAL, FCB; JACOBY (1884); MOR, OAX, PUE RILEY, CLARK & GIL- BERT (2001); USNM Syphrea frigida Bechyné $ GRO, MEX USNM-DGF Bechyné Syphrea minuta (Jacoby) TAB, VER JACOBY (1884); FCB Syphrea parvula (Jacoby) JAL, TAB, VER, YUC JACOBY (1891); USNM Syphrea pretiosa Baly DGO, GRO, TAMPS JACOBY (1891); USNM Syphrea smithi (Jacoby) OAX, TAB, TAMPS JACOBY (1891); USNM Syphrea sublaevipennis (Jacoby) OAX, VER FCB; JACOBY (1891) Syphrea teapensis (Jacoby) OAX, SLP, TAB, VER JACOBY (1891); USNM-DGF; USNM Systena abbreviata Jacoby PUE JACOBY (1902) 236 Bonner zoologische Beitráge 54 (2005) Genus Species Author Distribution Source Systena basalis J. DuVal VER FURTH & SAVINI (1998) Systena bitaeniata LeConte CHIH USNM-DGF Systena blanda Melsheimer BC?, CHIH, JAL, MICH, — PALLISTER (1953); NL, SIN, SLP?, SON, USNM TAB, VER Systena capitata Jacoby DF, GTO,GRO, HGO?, JACOBY (1884); FCB; MOR?, SLP?, VER?, ZAC? USNM Systena championi Jacoby GRO, MOR, OAX, VER USNM Systena contigua Jacoby CHIS, GRO, GTO, HGO, — JACOoBY (1884); OAX, SON?, TAMPS, USNM VER? ZAC Systena discicollis Clark CAMP, CHIH, DF, DGO, FCB; JACOBY (1884); GTO, JAL, MEX, MICH, USNM TAB, TAMPS, VER?, ZAC Systena gracilenta Blake NL BLAKE (1933a); FURTH & SAVINI (1998) Systena laticollis Jacoby GTO? FCB? Systena marginata Jacoby PUE, VER JACOBY (1884); USNM Systena mexicana Jacoby VER JACOBY (1884) Systena nigroplagiata Jacoby AGS, CHIH, DF, DGO, FCB; JACOBY (1884); GTO, GRO, JAL, MICH, PALLISTER (1953); MOR, OAX, PUE, VER USNM; USNM-DGF Systena obliterata Pallister CHIH PALLISTER (1953) Systena palmeri Jacoby CHIS, COAH, DGO, GRO, JACOBY (1884); FCB; SIN, SON, ZAC USNM Systena pectoralis Clark CHIS, GTO, OAX, VER JACOBY (1884); FCB Systena posticata Jacoby VER JACOBY (1884) Systena puncticollis Jacoby OAX JACOBY (1884) Systena s-littera (Linnaeus) CHIS, GTO, TAB, VER FCB; JACOBY (1884); USNM-DGF Systena salvini Jacoby CHIS USNM Systena scutellaris Jacoby VER JACOBY (1884) Systena semivittata Jacoby BCS, GRO, GTO, HGO, JACOBY (1884); FCB; MEX, MOR, NL, OAX, USNM; NHMB SIN Systena subcostata Jacoby MICH, MOR, VER JACOBY (1884); USNM Systena subrugosa Jacoby GTO, MICH, MOR JACOBY (1884); USNM David G. FURTH: Alticinae (Chrysomelidae) of Mexico 237 Genus Species Author Distribution Source Systena sulphurea Jacoby CHIH, DGO, GRO, MOR, JACOBY (1891); FCB; OAX USNM Systena thoracica Jacoby CAMP, HGO, PUE, JACOBY (1884); FCB; QROO, TAB, VER USNM A Systena undulata Jacoby AGS?, CAMP?, GRO?, JACOBY (1884); GTO, MOR?, VER USNM Systena variabilis Jacoby CHIH, CHIS, COL, DGO, FCB; JACOBY (1884); GRO, GTO, MICH, MOR, PALLISTER (1953); NAY, OAX, VER USNM; ZSMC Systena viridilimbata Jacoby GRO JACOBY (1891) Terpnochlorus americanus Bechyne & TAMPS USNM Bechyné Tetragonotes vittatus (Clark) VER USNM Trichaltica semihirsuta (Jacoby) GRO, TAB, VER SCHERER (1960): FCB; USNM-DGF Trichaltica tibialis (Jacoby) Mexico WILCOX (1975) Walterianella biarcuata (Chevrolat) CHIS, VER JACOBY (1886); FCB Walterianella durangoénsis (Jacoby) CHIH, DGO JACOBY (1892); FCB; PALLISTER (1953) Walterianella gouini Bechyné Mexico BECHYNE (1958b) Walterianella humeralis ? (Fabricius) Mexico? FCB Walterianella inscripta (Jacoby) OAX, SLP, VER JACOBY (1886); FCB; USNM Walterianella oculata ? (Fabricius) VER FCB Walterianella signata (Jacoby) CHIS, JAL, TAB, TAMPS, JACOBY (1886); FCB; VER, YUC USNM; USNM-DGF Walterianella sublineata (Jacoby) OAX, TAB, VER, YUC JACOBY (1886); FCB; USNM Walterianella tenuicincta (Jacoby) SLP, TAB, VER JACOBY (1886); FCB; USNM Walterianella venustula (Schaufuss) CHIS, COL?, GRO, JAL, USNM MICH, MOR, NAY?, QROO, TAMPS?, VER, YUC Bonner zoologische Beitráge Band 54 (2005) | Heft 4 | Seiten 239-245 Frozen Chrysomelids from Alpine Glaciers — Witnesses of the Postglacial Resettlement’ Elisabeth GEISER Salzburg, Austria Abstract. In the last decades a significant retreat of alpine glaciers occurred. During these processes, some pieces of peat bogs melted out from below the glaciers. In this article, conclusions are discussed, which can be inferred from the Chrysomelidae fragments found in these pieces. The first site is situated at Unteraar glacier in the Central Alps of Switzerland. The pieces of peat bogs are 3600 to 3800 years old and contained more than 5000 insect fragments. The Donaciinae remains showed a typical assemblage of a Cyperaceae fen peat. They contributed to a detailed assessment of the ecological conditions at this site 3600 years ago. From below the Pasterze glacier, northeast of Grossglockner in Hohe Tauern, Austria, also pieces of peat bogs melted out. In the 8100 years old pieces, fragments of Chrysomelidae were found, which could be identified as Oreina cacaliae (Chrysomelinae). In the same pieces of peat bogs pollen of Senecio was found, too. With the Oreina fragments it was possible to narrow down the assignment to Senecio fuchsii, because Oreina cacaliae feeds only on this species of Se- necio. But also interesting conclusions can be inferred concerning the postglacial resettlement. Oreina cacaliae has low dispersal ability. It persisted during maximum glacial extension in refugia far from its actual habitats, but managed to reach such a far site as the Pasterze 8100 years ago. This observation is contradictive to the theory of the “unfinished postglacial resettlement” due to lack of time. Other reasons for the restriction of the range of some species, which did not resettle obviously adequate habitats in the postglacial period, have to be discussed. Key words. Holocene, beetle fragments, Donacia, Oreina, paleoecology, speed of dispersal, Pasterze, Unteraar glacier INTRODUCTION Bonn, Oktober 2006 1.1. Retreat of glaciers and insect fragment analysis In the last two decades a significant retreat of alpine glaciers could be observed. This retreat is not a new and singular event, it happened several times in the post- glacial period (HORMES et al. 2001). There were several intervals, lasting several hundred years each, with warmer climate compared to the average temperature of the 20th century (Fig. 1). During those intervals peat bogs established in the wetlands after the retreat of the glaciers. During the next period of lower temperature, these peat bogs were covered by the growing glaciers. Nowadays, in the relatively warm climatic period, pieces of peat bogs are melting out from below the gla- ciers. In peat bogs insect fragments can be found, which are preserved for several thousand to more than hundred thousand years. The most abundant sclerites are remains of beetles. In the last 30 years, the method of insect fragment analysis was established, mainly by G. R. Co- ope and his team, who published a lot of very interest- ing results from sites in southern Great Britain, Scandi- navia and Northern America (COOPE 1986). Almost all of these investigations were made from areas around the Paper presented to the 6" International Symposium on the Chry- somelidae, Bonn, Germany, May 7, 2004. southern fringe of the northern ice shield. In Central Europe there was also a large ice shield during glacia- tion, which covered large parts of the alps and partly the areas of the surrounding lowlands (Fig. 2). After the re- treat of the glaciers many lakes and peat bogs emerged. In some of these places insect fragments were found and investigated, but only selectively and not systematically. There are two main causes for this situation: Firstly, the insect fragment analysis is more difficult here, because in Central Europe the number of beetle species is three times the number of species in Great Britain or in the Scandinavian sites. Secondly, in Central Europe we had not such an outstanding person like G. R. Coope, who performed these investigations systematically and con- tinually for more than three decades, with a lot of ex- perience and many co-workers. Therefore, in most cases, insect fragments were found only occasionally by macrophytic remains screening and identified only oc- casionally. This article shows the results of two interesting sites in Central Europe, where Chrysomelid fragments were preserved in peat bogs and melted out from alpine gla- ciers. The first site is situated in the Central Alps of Switzerland and was studied by an interdisciplinary pro- ject at the Institute of Geology at the University of Bern. The other sample derived from the Pasterze glacier near Grossglockner, located in Hohe Tauern, Austria. From 240 Bonner zoologische Beitráge 54 (2005) sample sample Pasterze Unteraar glacier a= a PATOL Wal i rere eS | 10000 B.P. 9000 8000 7000 6000 5000 4000 3000 2000 1000 0 Fig. 1. Amplitude of the average summer temperature (May to September) in the Eastern Alps during the Holocene (graphics af- ter SLUPETZKY et al. 1998). 1 7 \ EEE ee ee fle Me re | \ . tx 1 Neon . YS en . . LK en en % 49 Se . ~ Sr”) Y 6 e que Boeguue E ’ . a Pr) .r “i 4 Ff v , ‘u N LR» eek =r?, \ a s a ® x a a a $ RO ze, Bie vu.» % mu? 4 Ha Fig. 2. Maximum extension of the Wiirm glaciers in Central Europe, 22000 B.P. U: Unteraar glacier, P: Pasterze glacier. soe” a Elisabeth GEISER: Frozen Chrysomelids from Alpine Glaciers 24] both sites interesting conclusions can be inferred from the Chrysomelidae fragments. 1.2. Coleoptera fragment analysis provides additional results to the method of pollen analysis The method of pollen analysis, palynology, is well es- tablished for more than 80 years und revealed many in- teresting results, especially on glacial, interglacial and postglacial ecology. Also almost all sediments with Coleoptera remains contain a variety of pollen. There are two aspects to why Coleoptera fragments provide additional knowledge. Firstly, with Coleoptera fragments it is possible to nar- row down the climatic conditions at a special site better than by palynology. The MCR (Mutual Climatic Range) method, developed by ATKINSON et al. (1986), demon- strated its usefulness in many cases. It was used also in the study of the fragments from Unteraar glacier (JOST- STAUFFER 2000). Secondly, Coleoptera fragments can provide also more precise information about the vegeta- tion, in addition to the results of palynology. Whereas the tree pollen looks very characteristic and can be as- signed to the tree species in most cases, the pollen of many herbs can be assigned only to genus level or even to genus group. Therefore, fragments of monophagous or oligophagous beetles, like Chrysomelids, support useful additional information and can narrow down the results of pollen analysis to species level. 2. DONACIINAE FROM THE WESTERN ALPS 2.1. Site, materials and methods From the Vorfeld of the Unteraar glacier, at 1920 m a.s.l., located in the Central Alps of Switzerland (Fig. 2), 3600 to 3800 years old pieces of peat bogs melted out. They contained about 5000 insect remains; most of them were beetle fragments. The items were assigned to the beetle families by the co-workers of a research pro- ject from the Institute of Geology at the University of Bern and identified as exact as possible. Then special- ists of the identified Coleoptera families were contacted and asked, to check the identification and to do it more precisely, if possible. I was asked to check the Chry- somelidae fragments, except the Alticinae (which were checked by Manfred Dóberl, Abensberg, Germany), and then 150 well prepared items were sent to me. 2.2. Results and discussion The Chrysomelidae (except Alticinae) species found at that site belonged to the subfamily Donaciinae. The fol- lowing species were determined (Table 1). Table 1. Donaciinae identified from the Vorfeld of the Unteraar glacier, Switzerland. Nomenclature according to KIPPENBERG (1994). Donacia clavipes Fabricius, 1792 Donacia semicuprea Panzer, 1796 Donacia aquatica (Linnaeus, 1758) Donacia impressa Paykull, 1790 Donacia brevitarsis Thomson, 1884 Donacia brevicornis Ahrens, 1810 Donacia marginata Hoppe, 1795 Donacia obscura Gyllenhal, 1813 Donacia thalassina Germar, 1811 Donacia vulgaris Zschach, 1788 Donacia simplex Fabricius, 1775 Plateumaris sericea (Linnaeus, 1758) Only a few fragments (most of them are elytra or the distal fragment of one elytron) could be assigned ex- actly to one species, but often it was possible to narrow down the identification to two or three species. Some- times it was possible to identify the genus only. Other fragments could be clearly assigned to the subfamily Donaciinae, but could not be identified more precisely. The species mentioned in Table | are distributed in the whole of Europe, or, at least, in large areas of Europe. Their habitats are in such a wide temperature range that these Donaciinae were not used for the MCR, like some other Coleoptera species found at the Unteraar glacier. But these Donaciinae fragments contributed to more precise information about that habitat at this site 3800 years ago. In combination with the other identified bee- tle species it can be inferred that that habitat contained a marshy wetland with different sedges and reeds with Phragmites communis, Glyceria maxima, Sparganium simplex, Sparganium ramosum, Scirpus lacustris, Scir- pus palustris, Carex rostrata and other Carex sp. (Bo- tanical nomenclature according to FISCHER et al. (2005)). 3. OREINA FROM THE EASTERN ALPS 3.1. Site, materials and methods The Pasterze glacier is the largest glacier in the Eastern Alps. It is situated northeast of Grossglockner in Carin- thia, Austria (Fig. 2). In 1990 to 1996 pieces of peat bogs and stems of coniferous trees (Pinus cembra) melted out. They were collected by G. Lieb and H. Slu- petzky during routine measurements of the glacier and then investigated thoroughly (SLUPETZKY et al. 1998: GEISER 1998). 242 Bonner zoologische Beitráge 54 (2005) Fig. 3. Sample of 8100 years old Oreina cacaliae fragments, found near Pasterze glacier (Photo: J. Burgstaller). By the screening for macrophytic remains some beetle fragments were found (Fig. 3). These pieces of peat bog also yielded a rich pollen assemblage. It was 8180 + 70 years old. The original site of the peat is still covered by the Pasterze glacier, about 0.5 to 4 km upwards of the actual glacier terminus at approximately 2100 to 2300 m a.s.l. The pollen analysis and the macrophytic remains analy- sis revealed that during a period from 9000 to 8100 years B.P. a mountain forest was established at that site. The main tree species was Pinus cembra, and the typical assemblage of vascular plants was found there, espe- cially the tall herbaceous vegetation with Senecio spe- cies. The forest also contained several peat bogs. This ecosystem is typical for the Hohe Tauern, but nowadays situated several hundred meters lower. This time period was the largest interval of relatively warm climate in postglacial times (Fig. 1). The beetle fragments are stored in the collection of the author, which is designated to the Biologiezentrum in Linz, Austria. 3.2. Results and Discussion 3.2.1. Result of the identification of these beetle frag- ments: Oreina cacaliae. The fragments found at that site could be identified as Oreina cacaliae (Schrank, 1785). They belong to a minimum of three individuals. Elisabeth GEISER: Frozen Chrysomelids from Alpine Glaciers 243 Subfossil fragments of that genus are very rare. The evidence that this species was established at this site 8100 years ago is a very interesting fact. 3.2.2. Biology and actual distribution area of Oreina cacaliae. The habitat of the leaf beetle O. cacaliae is constituted by tall herbaceous vegetation in mountain- ous areas in Europe at 800 to 2500 m altitude. On some sites with adequate microclimatic conditions it can also be found down to 600 m a.s.l. This species is ovovi- vipar, like other Chrysomelids at the same altitude. For pupal development and hibernation O. cacaliae stays in the soil. Imagines up to three years old are known (ROWELL-RAHIER 1992). Oreina cacaliae is oligophagous on Asteraceae, it feeds only on Senecio ovatus (syn. Senecio fuchsii) and Adenostyles alliariae (HAGELE et al. 1996; there exists a remarkable variation in the nomenclature of that spe- cies, which was called Senecio fuchsii for decades. Mostly it is named now Senecio nemorensis or Senecio ovatus or Senecio nemorensis ssp. fuchsii. There exist further names and combinations of names. But the plant species is always the same.). The preference of the alti- tudes mentioned above, and the restriction to these two host plants are the reason for the fragmentation of their distribution area. Oreina cacaliae shows an aposematic colour with high variation. They synthesise chemical defence secretions out of secondary plant compounds from their host plants (PASTEELS et al. 1994). Therefore, their restriction to their host plant species has evolved long ago. There is no indication to a host plant change in the last 10 000 years. 3.2.3. Supplementation of the pollen analysis. In the pieces of peat bogs, which contained the O. cacaliae fragments, pollen of Senecio could also be identified. No Adenostyles pollen was found. Therefore it can be inferred that the Senecio pollen identified at the Pasterze belong to Senecio fuchsii. 3.2.4. Glacial refugia of Oreina. The majority of to- day’s distribution area of O. cacaliae, especially in the Alps, was covered with glaciers until 13 000 years ago. In many other locations of the actual range it was also impossible for the host plants to settle there, due to cli- matic reasons. Therefore, during the periods of large glacier extension, the majority of the populations of O. cacaliae had to persist in refugia far from their actual habitats. Figure 2 shows the maximum of the glacier ex- tension in Central Europe at the period of Würm, 22 000 years ago. 3.2.5. Dispersal ability of Oreina cacaliae and its host plants. The host plants have very high dispersal ability, as usual in Asteraceae, which disperse easily with their pappi. Therefore they reached adequate habi- tats quickly during the periods of retreating glaciers and during their growing periods, also. On the other hand, O. cacaliae has relatively low dispersal ability. This is very difficult to observe directly, but there are some facts that support this assumption: Field studies in the Swiss Alps showed that the adults moved only a few meters during summer, and a high percentage of individuals were recaptured in the next year, less than 5 m from the places where they were marked (ROWELL-RAHIER 1992). Coloration of O. cacaliae is highly variable, even within one population. But the set of colour variations is very typical for different regions within the whole area of distribution. This indicates that the populations are iso- lated for a long time. 3.2.6. Dynamics of the postglacial resettlement. The majority of the animal and plant species of Central Europe have high dispersal ability. This is due to more than 50 periods of growing and retreating glaciers. The habitats shifted to the south and then to the north within a few thousand years. Therefore species with high dis- persal ability had more chance to survive the Quaternary Ice Age. Species with high dispersal ability will spread to their habitats in Europe within a few 100 years (GEISER 1997). But also some species with relatively low dispersal ability managed to survive the climatic and therefore the habitat fluctuations. The species that didn’t manage this became extinct in Central Europe. The retreat of the glaciers after their maximum exten- sion started about 20 000 B.P. This retreat was inter- rupted by some colder climate intervals. At 10 300 B.P. the glacier dimensions were similar to the dimensions in the 20th century. There were some small fluctuations in the Holocene (Fig. 1). The most recent retreat of the gla- ciers started in 1860, interrupted by a short growing pe- riod at 1920. Nowadays, a significant retreat can be ob- served since 1982. The longest warm period lasted from 9000 to 8000 B.P. At a site that is still covered by the Pasterze glacier, a mountain forest with Pinus cembra could establish. The fragments of Oreina cacaliae show that even a species with such low dispersal ability could reach this remote site within a few thousand years. 3.2.7. Is “not yet finished postglacial resettlement” a relevant explanation for restricted distribution ar- eas? There are several species known with a very patchy distribution area and adequate habitats in be- tween. Some of these gaps in distribution are explained by “not yet finished resettlement” (since HOLDHAUS 1954) that implies that this is due to the lack of time in 244 Bonner zoologische Beitráge 54 (2005) the postglacial period to reach that habitats. On the con- trary, the example of Oreina cacaliae shows that a spe- cies with low dispersal ability could reach remote habi- tats within a few thousand years. The extension of a definite distribution area of a definite species always changes dynamically, with lower or higher amplitude. Therefore, resettlement is never “fin- ished”. The restriction of distribution areas can be ex- plained more probably as a result of the interaction of the following reasons: ° low dispersal ability ° low competitive ability * highly specialised ecological demands + geographical barriers (from the point of view of the species) ° and last, but not least, our lack of knowledge about the detailed biology and ecological demands of the definite species. The dynamics of dispersal and the real process of the postglacial resettlement is inferred indirectly only, in most cases. Modern molecular methods, such as isoen- zyme electrophorese (SCHMITT 2000) or DNA analysis, are very useful, but indirect, too. Real items on a real site with a definite age have to match the theory, if not the theory has to be revised. Our knowledge about the causes of distribution, about postglacial resettlement and changes of climatic and other ecological conditions during the Quaternary, de- pends on pieces of a dazzling, but highly interesting puzzle, which is composed of many items and methods. The more such items we find and examine with differ- ent methods we have, the clearer our idea of the post- glacial ecology will be. Acknowledgements. All these interesting results derived from Chrysomelidae fragments, which were sampled and extracted by other scientists: the Doncaciinae fragments I received from Dr. Monika Jost-Stauffer were in excellent preparation and packing (that cannot be taken for granted!). Also, the cooperation with her and the Institute of Geology of the University of Bern was excellent. The pieces of peat bogs, which contained the Oreina frag- ments, were sampled by Univ. Prof. Dr. Gerhard Lieb (In- stitute of Geology, University of Graz). The fragments were detected by Univ. Prof. Dr. Robert Krisai (Institute for Botany, University of Salzburg). Univ. Prof. Dr. Heinz Slupetzky (Department of Glaciology at the Institute of Geography, University of Salzburg) coordinated these in- vestigations at the Pasterze glacier and supported me with useful information. He also initialized the cooperation with Mag. Johann Burgstaller (Salzburg), an experienced pho- tographer of crystals, who managed to photograph that shiny surface of the Oreina-fragments. My husband was, as always, a meticulous proof-reader and made a lot of critical comments. It is an inestimable advan- tage to be criticised by a real friend and expert Coleop- terologist. Cordial thanks to all these colleagues. ZUSAMMENFASSUNG Die Klimaerwármung der letzten beiden Jahrzehnte hat zu einem Rückzug der Alpengletscher geführt. Bei die- sen Schmelzprozessen wurden an mehreren Stellen der Alpen unter den Gletschern gelegene Torfstiicke her- ausgeschwemmt. Die vorliegende Arbeit berichtet über die Ergebnisse von zwei Fundstellen, an denen subfossi- le Chrysomelidenreste gefunden wurden. Die erste Fundstelle befindet sich am Unteraargletscher in den Schweizer Zentralalpen. Die ausgeschwemmten Torfstiicke waren 3600 bis 3800 Jahre alt und enthielten über 5000 Insektenreste, die im Rahmen eines Projektes des Geologischen Institutes der Universitat Bern unter- sucht wurden. Die Autorin tiberpriifte und bestimmte die Reste der Unterfamilie Donaciinae. Die Artzusam- mensetzung entsprach der typischen Donacien-Fauna eines Cyperaceen — Flachmoores. Diese Befunde trugen zusammen mit den anderen Káferresten zu einer sehr genauen Einschätzung der ökologischen Verhältnisse an dieser Stelle der Alpen im genannten Zeitraum bei. Die zweite Fundstelle befindet sich unter der Pasterze nordöstlich des Grossglockners (Hohe Tauern, Öster- reich). Die herausgeschwemmten Torfstücke enthielten Reste eines 8100 Jahre alten Bergblattkäfers: Oreina cacaliae. Dieser Fund einer oligophagen Chrysomeli- denart tragt zu einer Prazisierung der botanischen Be- funde durch Pollen- und Großrestanalyse bei. Dadurch ist die Aussage móglich, dass es sich bei den ebenfalls in dieser Probe nachgewiesenen Senecio-Pollen um die Art Senecio fuchsii handelt. Weiters liefert dieser Fund einer wenig ausbreitungsfreudigen Art an einer Stelle, die von den Glazialrefugien dieser Káferart weit ent- fernt ist, einen interessanten Hinweis tiber den Verlauf bzw. die Geschwindigkeit der postglazialen Wiederbe- siedlung. Die eingeschränkte Verbreitung mancher Arten, trotz Vorhandenseins zahlreicher (anscheinend!) geeigneter Habitate wird immer wieder mit einer „nicht abge- schlossenen nacheiszeitlichen Wiederbesieldung“ er- klärt, mit dem Argument, dass seit dem Ende der letzten maximalen Gletscherausdehnung im Hochwürm vor 20 000 Jahren die Zeit noch nicht ausreichte für die Are- alausweitung auch von wenig ausbreitungsfreudigen Ar- ten. Wenn aber Arten wie Oreina cacaliae bereits vor über 8000 Jahren die entlegensten Habitate in den Alpen besiedelt haben, dann erscheint diese Theorie wenig Elisabeth GEISER: Frozen Chrysomelids from Alpine Glaciers 245 stichhaltig und die Ursache der eingeschränkten Ver- breitungsgebiete muss neu diskutiert werden. REFERENCES ATKINSON, T. C., BRIFFA, K. R., COOPE, G. R., JOACHIM, M. J. & PERRY, D. W. 1986. Climatic calibration of Cole- opteran data. Pp. 851-858 in: BERGLUND, B. E. (ed.) Handbook of Holocene Palaeoecology and Palaeohy- drology. John Wiley & Sons (Chichester-New York). Coope, G. R. 1986. Coleoptera Analysis. Pp. 703-713 in: BERGLUND, B. E. (ed.) Handbook of Holocene Pa- laeoecology and Palaeohydrology. John Wiley & Sons (Chichester-New York). FISCHER, M, ADLER, W. & OSWALD, K 2005. Exkursions- flora für Osterreich, Liechtenstein und Südtirol. 2. Au- flage. Biologiezentrum der Oberósterreichischen Lan- desmuseen, Linz. 1374 pp. GEISER, E. 1997. Das Vorkommen der Wespenspinne Ar- giope bruennichi (Scopoli) (Chelicerata, Araneidae) in Salzburg. Zur Arealausweitung einer wármeliebenden Art. Entomologisches Nachrichtenblatt, Wien 4 (2-4): 17. GEISER, E. 1998. 8000 Jahre alte Reste des Bergblattkáfers Oreina cacaliae (Schrank) von der Pasterze. Wissen- schaftliche Mitteilungen aus dem Nationalpark Hohe Tauern 4: 41-46. HAGELE, B., HARMATHA, J., PAVLIK, M. & ROWELL- RAHIER, M.1996. Sesquiterpene from the Senecioneae and their effect on food choice of the specialised leaf beetles Oreina cacaliae, Oreina speciosissima and the generalist snail Arianta arbustorum. Entomologia Ex- perimentalis et Applicata 80: 169-172. HOLDHAUS, K. 1954. Die Spuren der Eiszeit in der Tier- welt Europas. Abhandlungen der Zoologisch-Botani- schen Gesellschaft, Wien 18: 493 pp. HORMES, A., MÜLLER, B. U. & SCHLÜCHTER, C. 2001. The Alps with little ice: evidence for eight Holocene phases of reduced glacier extent on the Central Swiss Alps. The Holocene 11, 3: 255-265. JOST-STAUFFER, M. 2000. A Holocene fossil coleopteran (beetle) assemblage from the Central Swiss Alps: cli- mate and ecology. Eclogae Geologicae Helveticae 93: 481-490. KIPPENBERG, H. 1994. Chrysomelidae. Pp. 9-92 in: LOHSE, G. A. & LUCHT, W. H. (eds.) Die Käfer Mitteleuropas, Band 14, 3.Supplementband. Goecke & Evers, Kre- feld. PASTEELS, J. M., ROWELL-RAHIER, M., BRACKMAN, J.-C. 8 DALOZE, D. 1994. Chemical defense in adult leaf bee- tles updated. Pp. 289-310 in: JOLIVET, P. H., Cox, M. L. & PETITPIERRE, E. (eds.) Novel aspects of the biol- ogy of Chrysomelidae. Dordrecht: Kluwer Academic Publishers. ROWELL-RAHIER, M. 1992. Genetic structure of leaf-beetle populations: microgeographic and sexual differentia- tion in Oreina cacaliae and Oreina speciosissima. En- tomologia Experimentalis et Applicata 65: 247-257. SCHMITT, T. 2000. Glaziale Refugien und postglaziale Arealausweitung von Polyommatus coridon (Lepidop- tera: Lycaenidae). Verhandlungen der Westdeutschen Entomologentagung 1999, Diisseldorf: 65-79. SLUPETZKY, H., KRISAI, R. & LIEB, G. K. 1998. Hinweise auf kleinere Gletscherstánde der Pasterze (National- park Hohe Tauern, Kárnten) im Postglazial. Ergeb- nisse von '*C-Datierungen und Pollenanalysen. Wis- senschaftliche Mitteilungen aus dem Nationalpark Hohe Tauern 4: 225-240. Author’s address: Dr. Elısabeth GEISER, Saint-Julien- Straße 2/ 314, 5020 Salzburg, Austria. E-mail: elisa- beth.geiser@gmx.at f Bonner zoologische Beitráge Band 54 (2005) | Heft 4 | Seiten 247-252 Bonn, Oktober 2006 Striking Differences in Behaviour and Ecology Between Populations of Chrysomela lapponica' j Jürgen GROSS!’ & Nina E. FATOUROS” "Federal Biological Research Centre for Agriculture and Forestry, Institute for Plant Protection in Fruit Crops, R Dossenheim, Germany "Institut für Biologie, Freie Universität Berlin, Germany Abstract. Willows and poplars have been suggested as ancestral host plants of Chrysomela species (Coleoptera, Chry- somelidae). In the leaf beetle species C. lapponica, some populations are specialized on willow species, but others on birch. In this paper, we review results from previous studies on ecological interactions between C. /apponica and its bi- otic and abiotic environment. We found striking differences between individuals of different populations in morphologi- cal traits, chemical composition and origin of larval glandular secretions, feeding habits, host preferences and feeding stimulants of larvae and adults. Furthermore, we discovered significant between-population differences in bottom-up, host plant depending performance parameters such as developmental time, body mass, and mortality as well as differ- ences in top-down effects such as defensive secretions against predators, parasitoids and pathogens. We review results of hybridization experiments between different populations and then address the question whether the different host- specific populations of C. /apponica represent more than one species. Key words. Speciation, post zygotic isolation, host shift, larval defence, top-down effect, bottom-up effect, willow, birch, Chrysomela lapponica 1. INTRODUCTION Leaf beetle species belonging to the Chrysomela inter- rupta-group, a subtaxon of the Chrysomelina, show variation in host plant specialization. Some species are specialized to feed and oviposit on willows (Salicaceae), while others live only on birch or alder (Betulaceae) (BROWN 1956; GROSS 1997; GROSS et al. 2004b; GROSS & HILKER 1995; TERMONIA et al. 2001; TERMONIA & PASTEELS 1999). Among the C. interrupta-group, popu- lations of C. lapponica are known to be exclusively spe- cialized on either birch or on willow. Chrysomela lap- ponica is distributed almost continuously in Northern Europe (e.g., Finland, Sweden, Norway, Russia, Baltic States) (GROSS 2001; GROSS et al. 2004b; HILKER & SCHULZ 1994; MACHKOUR M'RABET 1996; ZVEREVA et al. 1995) and is divided by the boreal coniferous forest belt from the populations in Central Europe, which are distributed patchily in mostly higher or colder regions such as Germany, Czech Republic, Poland, Northern It- aly and France (GROSS 1997; GROSS et al. 2004c; HILKER & SCHULZ 1994). The larvae and adults from Northern Europe feed and oviposit only on some willow species, while most of the Central European populations are monophagous on birch (GROSS 1997; GROSS & HILKER 1995). The aim of this review is to summarize information about the ecological factors, which may have driven the divergent evolution of host plant spe- ' Paper presented to the 6" International Symposium on the Chry- somelidae, Bonn, Germany, May 7, 2004. cialization in the Northern and Central European popu- lations of C. lapponica. To elucidate this matter, we compared two populations of C. /apponica, which are specialized on different host plants. One of the popula- tions occurs in Finnish Lapland and feeds mainly on the willow Salix borealis. The other population, feeding ex- clusively upon the birch Betula pubescens, was ob- served in the Czech Republic, close to the city of Mari- anske Lazne. Both, birch and willow hosts occur sympatrically in the habitats of the Finnish and Czech beetles. We conducted field and laboratory investiga- tions over nine years, using morphological, chemical, physiological and microbiological analyses to determine whether or not these host specialized populations were one and the same species. 2. OVERVIEW OF PRESENT RESULTS 2.1. Morphological differences The adults of birch-feeding populations of C. lapponica differ in the colour patterns of their red and black elytra compared with their willow-feeding counterparts from Northern Europe and Central Europe. While the ratio of black and red colour is about fifty-fifty in the Czech population, the relative size of the black marks of the Finnish beetles’ elytra is significantly higher (GROSS et al. 2004c). The tibiae of the adults’ first leg pair of the Czech population are predominantly black coloured, whereas the tibiae of the Finnish beetles are mainly red. Sometimes complete blue morphs occur spontaneously 248 Bonner zoologische Beitráge 54 (2005) within the populations of Central Europe, but never in the populations from Northern Europe. The larvae of the Chrysomelina and Phyllodectina pos- sess nine pairs of dorsolateral defensive glands located on thorax and abdomen. When disturbed, the glandular reservoirs can be everted, and subsequently be inverted with the help of specialized retractor muscles. The res- ervoirs are filled with volatile compounds (GARB 1915; HINTON 1951). However, there are striking chemical differences between the Czech (birch-feeding) and Fin- nish (willow-feeding) populations of C. lapponica in the chemical composition of these larval secretions. Gas chromatography coupled with mass spectrometry (GC- MS) was used to identify the composition of the larval secretions and to quantify their amounts. The secretion of the birch-feeding larvae from the Czech population contained carboxylic acids and carbonic acid esters (Fig. 1A) which were synthesized de novo and esterified with alcohols taken from the plant (SCHULZ et al. 1997). However, like in all other willow-feeding Chrysomela- species, the secretion of the willow-feeding larvae from Finland contained salicylaldehyde as the major compo- nent (Fig. IB). In all systems studied to date, salicylal- dehyde is derived from salicin uptaken from willow leaves by the larvae (GROSS 1997; GROSS et al. 2002; GROSS & HILKER 1995; PASTEELS et al. 1988). Interest- ingly, the two C. lapponica secretion types can easily be distinguished in the field according to their different odours. The secretion of the willow-feeders smells like fruit salad, while the birch-feeder’s secretion smells like almond liqueur. 2.2. Ecological interactions First, we investigated the bottom-up effects, which are influenced by the first trophic level, the host plants. To assess these effects, we used different parameters for measuring the performance of the beetles on different plants. These were mortality, developmental time, vol- ume of the larval secretion, fecundity and body weight of the neonate larvae and adults. Secondly, we investi- gated the top-down effects, that is, those influenced by the third trophic level, like predators, parasitoids and pathogens. An interspecific competition (within the 2nd trophic level) could not be detected (GROSS 2001). Bottom-up effects. The adults of both Finnish and Czech populations prefer their natural host plant for feeding and oviposition over that of the host plant of the other population (GROSS & HILKER 1995). While there was neither a difference in fecundity nor in body weight of neonate larvae, we observed significant differences in mortality when the beetles were reared on their natural host plant versus the host plant of the other population without enemies in the laboratory (GROSS et al. 2004a). While a few individuals of the birch-specialized popula- tion could be reared on willow, all neonate larvae of the willow-specialized population died after having been transferred to birch (GROSS et al. 2004a). When indi- viduals of the two C. /apponica populations were reared on their natural host plants, the birch specialists suffered higher mortality than the willow specialists (GROSS et al. 2004a). Probably as a consequence of host plant spe- cialization in C. lapponica, there was a significantly shorter developmental period for the Finnish beetles on willow compared to that of the Czech beetles feeding upon birch (GROSS et al. 2004a). If the “slow-growth, high-mortality” hypothesis (WILLIAMS 1999; DENNO et al. 1990) applies to C. lapponica, the prolonged devel- opmental time might be detrimental to the birch-feeding larvae, because they would remain in the more vulner- able larval stages for a longer time period. Higher mor- tality could result from higher predation, parasitism or higher microbial infection rates. Fecundity does not dif- fer between individuals of both populations. However, a smaller volume of larval secretion is produced by the birch-feeding larvae (GROSS et al. 2004a). Top-down effects. Laboratory experiments showed that both types of larval secretions (above-mentioned) of C. lapponica are highly effective against generalist preda- tors with chewing-biting mouthparts like coccinellids, but ineffective against generalists with sucking-piercing mouthparts (GROSS 2001). However, significant differ- ences exist between both secretion types in the effec- tiveness towards pathogens. The secretion of the wil- low-feeding larvae was highly effective against various microorganisms, especially against the entomopathoge- nous fungus Metarhizium anisopliae. When applied di- rectly onto the blastospores, fungal germination and growth was greatly inhibited by the main component of the secretion, salicylaldehyde. Additionally, physiologi- cal quantities of volatile salicylaldehyde were shown to reduce germination and growth of M. anisopliae up to 45 mm around the application site (GROSS et al. 2002). In contrast, no antimicrobial activity was found in the larval secretion of the birch-feeding population. Labora- tory results supported field observations because no in- fected ontogenetic stages of C. lapponica were found in Finland, while infected larvae and pupae were collected in the Czech Republic (GROSS 2001). It was previously known that salicylaldehyde, the main component of the secretion of willow-feeding larvae, did not repel but instead attracted a specialized enemy, the predaceous hoverfly Parasyrphus nigritarsis (KOPF et al. 1997). Therefore, laboratory studies were con- ducted to evaluate the response of P. nigritarsis larvae towards the salicylaldehyde-free secretion of the birch- feeding larvae. Parasyrphus nigritarsis hunted for lar- vae of both populations, but the secretion of the birch- feeding larvae was significantly less attractive than that of the willow-feeding larvae. Faeces of the willow- Jürgen Gross & Nina E. FATOUROS: Behaviour and Ecology of Chrysomela lapponica 249 feeding larvae attracted this predator, as well. However, P. nigritarsis did not respond to faeces of larvae fed with birch leaves. In the field, P. nigritarsis is common in both habitats, but infested egg batches were found only on willows. The hover fly was never observed preying on individuals of the birch-feeding population (GROSS et al. 2004b). In general, a higher percentage of the willow-specializ- ed C. lapponica were found to be parasitized compared with the birch specialists. Both, in 1999 and in 2000, the percentage of parasitation by the phorid fly Megaselia opacicornis of the willow-specialized C. lapponica specimens was significantly higher than of the birch specialists. This parasitoid cannot be repelled by larval secretions. Parasitization by other species was low in each year both for birch and willow specialists (GROSS et al. 2004b). The parasitoids that were reared from parasitized C. lapponica larvae or pupae are all known to only develop in Chrysomela species (COX 1994; DIS- NEY et al. 2001; ZVEREVA & KOZLOV 2000). The influ- ence of host plants or larval secretions on the host- finding behaviour of parasitoids was tested in field ex- periments in Finland. These experiments showed that parasitoids use both chemical signals of the willows (GROSS et al. 2004b) and volatile secretions of the wil- low-feeding larvae as kairomones for host localization (ZVEREVA & RANK 2004). Abiotic factors (e.g., temperature, radiation). The adaptive value of the elytral colouring, one of the most conspicuous differences between the adults of both populations of C. lapponica, was examined. Adults of the Finnish and other populations from Northern Europe were found to be significantly darker in colour than their Czech counterparts (GROSS 1997; GROSS et al. 2004c; ZVEREVA et al. 2003). As a result, Finnish bee- tles heated up more quickly when exposed to insolation. The dark colouring of the Finnish beetles may therefore be an adaptation to their colder environment with shorter seasonal vegetation periods (GROSS et al. 2004c). Also, some endogenous factors were found to differ between the populations. Batch calorimetry ex- periments gave evidence that the specific metabolic rate during hibernation at low temperatures was lower in the Czech beetles. Individuals of the Czech population may have to cope with higher and fluctuating winter tem- peratures. Additionally, the walking speed and the feed- ing activities of the Finnish beetles were higher at low temperatures when compared to the Czech beetles (GROSS et al. 2004c). Crossing experiments. Very few examples of host race formations are known in chrysomelids. Among them are the Gallerucella nymphaeae species complex (HIPPA & KOPONEN 1986; NOKKALA & NOKKALA 1994; NOK- KALA 8 NOKKALA 1998; PAPPERS et al. 2002b; PAPPERS et al. 2002a) and the Lochmaea caprea (Kreslavskiy & Mikheyev 1994). However, unconfined hybrid forma- tion was possible between the host races in these exam- ples. Fatouros ef al. (unpublished results) tried to inter- breed individuals of C. lapponica from Finland (willow- feeders) and the Czech Republic (birch-feeders) in the laboratory. Only a unidirectional hybrid formation be- tween females from the Czech Republic and males from Finland succeeded, whereas the reciprocal pairings pro- duced no offspring. But the resulting hybrid larvae could not survive. By keeping the ability to feed on only willow, they were doomed to death on birch, as their birch-feeding mothers laid the eggs exclusively on birch leaves. This represents possibly a new postzygotic 1sola- tion mechanism (Fatouros et al. in press). 3. DISCUSSION This review focuses on the ecological factors which se- lectively influence phenotypically different individuals of C. lapponica. But natural selection is only one of several factors driving the evolution of organisms (MAYR 1997). Other random evolutionary factors like genetic drift and mutation were not investigated. Influence of biotic selection factors. The influence of biotic selection factors on the two investigated popula- tions of C. lapponica is summarized in Table 1. No competition between C. lapponica and other herbivores could be detected in both habitats and upon both host plants (GROSS 2001). The respective host plant species had an effect on the performance of the leaf beetle. Birch-feeding larvae feeding under standardized condi- tions on leaves of their natural host plant, B. pubescens, developed more slowly than willow-feeding larvae reared on leaves of their natural host plant, S. borealis (GROSS et al. 2004a). This laboratory result, however, may not be of ecological significance, because devel- opmental time increases with higher temperatures (HONEK & KOCOUREK 1988; HOWE 1967). Therefore the adverse effect of the birch on the beetles’ develop- mental time could be compensated for, since the mean temperatures during the vegetation period in the Czech Republic were significantly higher than in Finland (GROSS et al. 2004c). If this is not the case, the pro- longed developmental time might be detrimental to the birch-feeding larvae, because they remain in vulnerable immature stages for a longer period of time. Higher mortality could result from enhanced predation, parasi- tation or microbial infection (HAGGSTROM & LARSSON 1995; WILLIAMS 1999). Birch-feeding larvae were also found to have reduced volumes of secretions. This had obviously no effect on the larvae, because even little quantities of secretion were highly effective against ladybird beetles (GROSS et al. 2004a). Apart from directly affecting the beetles de- 250 velopment and larval secretion quantity, the host plant may also have an indirect influence on top-down effects by determining the chemical composition of the larval defence secretions. A laboratory experiment showed that willow-feeding larvae were better defended against pathogens than birch-feeding larvae due to the salicylal- dehyde in the secretion of the willow-feeding larvae (GROSS et al. 2002). Although birch-feeding larvae seem to face a number of disadvantages as a result of feeding on an apparently suboptimal host, there is evidence for a trade-off: the secretion of birch feeding larvae was shown to be less attractive to the specialist predator, P. nigritarsis, than the salicylaldehyde-containing secretion of larvae of the willow feeding populations (GROSS et al. 2004b). Con- sequently, our data suggest that the secretion of the birch-feeding larvae provides a selective advantage, due to a decreased predation pressure by specialists on eggs and larvae. Influence of abiotic selection factors. Individuals of C. lapponica in Finland and the Czech Republic live in en- vironments, which differ in their temperature conditions and in the length of their vegetation period. Those adult beetles living in colder environments (Finland) benefit from their darker elytral colouring by heating up more rapidly. The higher body temperatures attained posi- tively affect their walking and feeding behaviour (GROSS et al. 2004c). Additionally, their temperature range for feeding and walking activity is lower than in their counterparts living in warmer environments (Czech Republic). In contrast, the lower specific meta- bolic rates of the Czech beetles at low temperatures are favourable during hibernation, because their energy re- sources decrease more slowly than the energy resources of the Finnish beetles. The elytral colouring as well as internal physiological factors lead to an optimal adapta- tion to temperature conditions in the different habitats of C. lapponica in Central and Northern Europe (GROSS et al. 2004c). Summarizing the effects of the three trophic levels on C. lapponica, plant (1.e., host plant species), herbivorous insect (1.e., competitors), and antagonists (1.e., predators, parasitoids, and pathogens), the following evolutionary scenario arises. Since larvae adapted to willow have a shorter developmental period if reared on willow than birch-feeders reared on birch, willow is obviously a bet- ter food plant than birch. Competition between C. lap- ponica and other herbivores could not be detected. Con- sequently food does not seem to be limiting. Feeding on willow benefits larvae due to the antimicrobial activity of their willow-derived salicylaldehyde-rich secretion. The secretion protects willow-feeding larvae against fungal infection. Birch-feeding larvae aré more vulner- able to fungal infections since their secretion lacks sig- Bonner zoologische Beitrage 54 (2005) nificant antifungal activity. Nevertheless, feeding on birch has a major advantage as it allows those larvae to produce a secretion which is significantly less attractive to specialist predators like the hoverfly, P. nigritarsis, than the secretion of the willow-feeding larvae. The dif- ferent elytral colouring and temperature optima traits of the imagoes from both populations appear to be highly adaptive to the extant environmental conditions in their current ranges. Is the willow the ancestral host plant? The results re- viewed here suggest that the ancestral host plant of C. lapponica belonged to the Salicaceae: While individuals of the birch-feeding population of C. lapponica can ob- viously cope with the secondary metabolites of the wil- low, as they are still able to develop from larvae to adults when fed with willow leaves, individuals of the wiliow-feeding population cannot survive when fed with birch leaves (GROSS et al. 2004a). When reared on Salix fragilis in the laboratory, birch-feeding larvae pro- duce trace amounts of salicylaldehyde in their secretion (GROSS 1997; HILKER & SCHULZ 1994), suggesting that this ability could be a vestige of the ancestral pathway. These findings indicate that the birch-feeding popula- tion has adapted very well to its new host-plant in the interim, because its performance is negatively affected when reared on willows (GROSS et al. 2004a). Further studies of the natural enemies of the Nearctic species within the C. interrupta-group feeding upon alder have to be conducted in order to elucidate the convergent or reversal host shifts between Betulaceae and Salicaceae, as it was suggested by TERMONIA et al. (2001). Do the birch feeding populations represent a host race or a new species? According to our findings ob- tained over the past nine years of field and laboratory experimentation, we conclude that the pronounced and consistent differences in morphology, ecology and ethology between most birch-feeding and willow- feeding populations of C. lapponica, the species C. lap- ponica should be divided in two distinct species. With regard to ecologically important traits like feeding spe- cialization and oviposition behaviour, one may distin- guish between a birch-feeding species, to which most of the investigated populations from Central Europe be- long (Germany, Czech Republic, Poland and France (Massif Central)), and a willow-feeding species, which includes all populations from Northern Europe (Finland, Norway, Russia) and one population from France (Queyras). Acknowledgements. The authors greatly appreciate Monika Hilker (Berlin), Horst Kippenberg (Herzogenau- rach), Seppo Neuvonen (Kevo Subarctic Research Station, Finland), Jacques Pasteels (Belgium), Andreas Vilcinskas (GieBen), and Andrzej Warchalowski (Poland). We thank Jürgen GROSS & Nina E. FATOUROS: Behaviour and Ecology of Chrysomela lapponica 251 Meike Brick (Walldorf) and two anonymous reviewers for important remarks and linguistic improvements. REFERENCES BROWN, W. M. 1956. The new world species of Chry- somela L. (Coleoptera, Chrysomelidae). The Canadian Entomologist 88: 3-54. Cox, M. L. 1994 The Hymenoptera and Diptera parasitoids of Chrysomelinae. Pp 419-467 in: JOLIVET, P. H., Cox, M. L. & PETITPIERRE, E. (eds) Novel aspects of the biology of Chrysomelidae. Kluwer Academic Pub- lisher, Dordrecht. DENNO, R. F., LARSSON, S. & OLMSTEAD, K. L. 1990. Role of enemy-free space and plant quality in host-plant se- lection by willow beetles. Ecology 71: 124-137. DISNEY, R. H., ZVEREVA, E. L. & MOSTOVSKI, M. B. 2001. A scuttle fly (Diptera: Phoridae) parasitizing a beetle (Coleoptera: Chrysomelidae) in Russia. Entomologica Fennica 12: 59-63. FATOUROS, N. E., HILKER, M. & Gross, J. in press. Re- productive isolation between populations from North- ern and Central Europe of the leaf beetle Chrysomela lapponica L. Chemoecology. GARB, G. 1915. The eversible glands of a chrysomelid larva, Melasoma lapponica. Journal of Entomology & Zoology 7: 87-97. Gross, J. 1997. Speziation beim Blattkáfer Chrysomela lapponica L.: Vergleichende Studien zu Chemischer Okologie, Verhalten und Morphologie verschiedener Ontogenesestadien aus zwei allopatrischen Popula- tionen. [Speciation in the leaf beetle Chrysomela lap- ponica L.: Comparative studies in chemoecology, ethology and morphology of different stages of two al- lopatric populations]. Sitzungsberichte der Gesell- schaft naturforschender Freunde zu Berlin N.F. 36: 125-142. Gross, J. 2001. On the evolution of host plant specializa- tion in leaf beetles (Coleoptera: Chrysomelina). 137 pp., Logos Verlag, Berlin Gross, J. & HILKER, M. 1995. Chemoecological studies of the exocrine glandular larval secretions of two chry- somelid species (Coleoptera): Phaedon cochleariae and Chrysomela lapponica. Chemoecology 5/6: 185- 189. Gross, J., FATOUROS, N. E. & HILKER, M. 2004a. The significance of bottom-up effects for host plant spe- cialization in Chrysomela leaf beetles. Oikos 105: 368- 376: Gross, J., FATOUROS, N. E., NEUVONEN, S. & HILKER, M. 2004b. The importance of specialist natural enemies for Chrysomela lapponica in pioneering a new host plant. Ecological Entomology 29: 584-593. Gross, J., PODSIADLOWSKI, L. & HILKER, M. 2002. An- timicrobial activity of exocrine glandular secretion of Chrysomela larvae. Journal of chemical Ecology 28: 317-331. Gross, J., SCHMOLZ, E. & HILKER, M. (2004c) Thermal adaptations of the leaf beetle Chrysomela lapponica L. (Coleoptera: Chrysomelidae) to different climes of Central and Northern Europe. Environmental Ento- mology 33: 799-806. HAGGSTROM, H. & LARSSON, S. 1995. Slow larval growth on a suboptimal willow results in high predation mor- tality in the leaf beetle Galerucella lineola. Oecologia 104: 308-315. HILKER, M. & SCHULZ, S. 1994. Composition of larval se- cretion of Chrysomela lapponica (Coleoptera, Chry- somelidae) and its dependence on host plant. Journal of chemical Ecology 20: 1075-1093. HINTON, H. E. 1951. On a little-known protective device of some chrysomelid pupae (Coleoptera). Proceedings of the Royal entomological Society of London 26: 67-73. HIPPA, H. & KOPONEN, S. 1986. Morphological, cytologi- cal, ecological and ethiological evidence of reproduc- tive isolation between Galerucella nvmphaeae (L.) and G. sagittariae (Gyll.) (Coleoptera, Chrysomelidae) in Fennoscandia. Annales entomologici Fennici 52: 49- 62. HONEK, A. & KOCOUREK, F. 1988. Thermal requirements for development of aphidophagous Coccinellidae (Coleoptera), Chrysopidae, Hemerobiidae (Neurop- tera), and Syrphidae (Diptera): some general trends. Oecologia 76: 455-460. Howe, R. W. (1967) Temperature effects on embryonic development in insects. Annual Review of Entomol- ogy 12: 15-42. KOPF, A., RANK, N. E., ROININEN, H. & TAHVANAINEN, J. 1997. Defensive larval secretions of leaf beetles attract a specialist predator Parasvrphus nigritarsis. Ecologi- cal Entomology 22: 176-183. KRESLAVSKIY, A. G. & MIKHEYEV, A. V. 1994. Gene ge- ography of racial differences in Lochmaea capreae L. (Coleoptera, Chrysomelidae), and the problem of sympatric speciation. Entomological Review 73: 85- 9) MACHKOUR M'RABET, S. 1996. Differenciation genetique de Chrysomela lapponica (L.) en Europe et evolution des secretions defensives: comparaison des popula- tions de Finlande et de France (Queyras). Universite Libre de Bruxelles, Section Interfacultaire d'Agrono- mie, Laboratoire de Biologie Animale et Cellulaire, Bruxelles MAYR, E. 1997. The objects of selection. Proceedings of the National Academy of Sciences of the USA 94: 2091-2094. NOKKALA, C. & NOKKALA, S. 1994. Sympatric speciation in Galerucella. Pp. 259-262 in: JOLIVET, P. H., Cox, M. L. & PETITPIERRE, E. (eds) Novel aspects of the bi- ology of Chrysomelidae. Kluwer Academic Publisher, Dordrecht. NOKKALA, C. & NOKKALA, S. 1998. Species and habitat races in the chrysomelid Galerucella nymphaeae spe- cies complex in Northern Europe. Entomologia ex- perimentalis et applicata 89: 1-13. PAPPERS, S. M., VAN DER VELDE, G. & OUBORG, N. J. 2002a. Host preference and larval performance suggest host race formation in Gaterucella nymphaeae. Oeco- logia 130: 433-440. PAPPERS, S. M., VAN DER VELDE, G., OUBORG, N. J. & VAN GROENENDAEL, J. M. 2002b. Genetically based polymorphisms in morphology and life history associ- ated with putative host races on the water lily leaf bee- tle, Galerucella nymphaeae. Evolution 56: 1610-1621. PASTEELS, J. M., ROWELL-RAHIER, M. & RAUPP, M. J. 1988. Plant-derived defense in chrysomelid beetles. Pp. 235-272 in: BARBOSA, P. & LETOURNEAU, D. K. (eds) Novel aspects of insect-plant interactions. John Wiley & Sons, New York. [597 Nn m SCHULZ, S., GROSS, J. & HILKER, M. 1997. Origin of the defensive secretion of the leaf beetle Chrysomela lap- ponica. Tetrahedron 53: 9203-9212. TERMONIA, A. & PASTEELS, J. M. 1999. Larval chemical defence and evolution of host shifts in Chrysomela leaf beetles. Chemoecology 9: 13-23. TERMONIA, A., HSIAO, T. H., PASTEELS, J. M. & MIL- INKOVITCH, M. C. 2001. Feeding specialization and host-derived chemical defense in Chrysomeline leaf beetles did not lead to an evolutionary dead end Pro- ceedings of the National Academy of Sciences of the USA 98: 3909-3914. WILLIAMS, I. S. (1999) Slow-growth, high-mortality — a general hypothesis, or is it? Ecological Entomology 24: 490-495. ZVEREVA, E. L. & KOZLOV, M. V. 2000. Effects of air pol- lution on natural enemies of the leaf beetle Me/asoma lapponica. Journal of applied Ecology 37: 298-308. ZVEREVA, E. L. & RANK, N. E. 2004. Fly parasitoid Megaselia opacicornis uses defensive secretions of the Bonner zoologische Beitráge 54 (2005) leaf beetle Chrysomela lapponica to locate its host. Oecologia 140: 516-522. ZVEREVA, E. L., KOZLOV, M. V. & KRUGLOVA, O. Y. 2003. Colour polymorphism in relation to population dynamics of the leaf beetle, Chrysomela lapponica. Evolutionary Ecology 16: 523-539. ZVEREVA, E. L., KOZLOV, M. V. & NEUVONEN, S. 1995. Decrease in feeding niche breadth of Melasoma lap- ponica (Coleoptera: Chrysomelidae) with increase in pollution. Oecologia 104: 323-329. Authors’ addresses: Dr. Jiirgen GROSS (corresponding author), Biologische Bundesanstalt fiir Land- und Forstwirtschaft, Institut fiir Pflanzenschutz im Obstbau, Schwabenheimer Str. 101, D-69221 Dossenheim, Ger- many, E-mail: j.gross@bba.de; Nina E. FATOUROS, In- stitut für Biologie, Freie Universität Berlin, Had- erslebener Str. 9, 12163 Berlin, Germany. Bonner zoologische Beitráge Band 54 (2005) | Heft 4 | Seiten 253-259 Bonn, Oktober 2006 Parameres — Similarities and Differences in Chrysomelidae and Cerambycidae (Coleoptera)' P Lasse HUBWEBER & Michael SCHMITT Zoologisches Forschungsmuseum Alexander Koenig (ZFMK), Bonn, Germany Abstract. Parameres, or lateral lobes, of the male copulatory organ are present in many insect groups, while in others they are reduced or completely lacking. Nearly all long-horned beetles have parameres, while only the leaf beetle sub- taxa Megalopodinae, Zeugophorinae, Orsodacninae, Bruchidae, Sagrinae, Donaciinae and the genus Timarcha possess them. In these taxa, parameres can consist of one or two lobes, can be very long or quite short, and the number and den- sity of their setae can vary. Based on this wide morphological variability, we draw conclusions on the phylogeny of Chrysomeloidea and then formulate a hypothesis accounting for the probably diverse functions parameres might have in both the long-horns and in the leaf beetles. One recent discovery is the existence of minute openings in the surface of the parameres, which may indicate the presence of glands. These structures are illustrated and their possible functions dis- cussed. Key words. aedeagus, male genitalia, morphology, phylogenetics 1. INTRODUCTION 1.1. General Surprisingly there are only few papers on parameres, or lateral lobes, of the male aedeagus found in many bee- tles. Although parameres are quite complex and species- specific, their functional role is largely unknown, even though these structures have at least one known func- tion. Hubweber (unpubl. data) used genital characters, including those of the aedeagal parameres in an analysis of the phylogenetic relationships among the subfamilies of the Cerambycidae. Because some Chrysomelidae groups also possess parameres, we expanded this initial survey to include both the Cerambycidae and Chry- somelidae. We then compared character states of pa- rameres between the two groups. In addition, we briefly examined the function of the parameres and then went on to discuss the discovery and possible significance of minute holes indicative of glandular openings on the surface of the parameres. 1.2. Terminology Over the decades many genitalic structures in beetles have been described, mostly for taxonomic purposes. But not all entomologists have followed the same terms in the description of these different structures. The term “aedeagus” for example, is often used synonymically with “median lobe”, but we use it to describe the me- dian lobe and the tegmen together. In our opinion the latter structure, the tegmen, consists of the parameres ' Paper presented to the 6" International Symposium on the Chry- somelidae, Bonn, Germany, May 7, 2004 (or lateral lobes) and the basal piece or tegminal struts, connected by the tegminal ring. 1.3. Literature Review VERMA (1996) and CHEN (1985) illustrate the loss of the parameres within the Chrysomelidae. While Megalopo- dinae, Zeugophorinae, Orsodacninae, Bruchidae, Sagri- nae, Donaciinae and Timarcha show a typical tegmen, the tegminal ring and therefore also the parameres, are reduced in all other subfamilies. Stenhomalus is the sin- gle cerambycid genus known lacking parameres (WU & JIANG 1993). Another question concerning the homol- ogy of genitalia in chrysomelids is the single genus within Chrysomelinae, Timarcha, showing a complete tegminal ring with parameres. HARNISCH (1915) examined the male genital system of some chrysomelids including Plateumaris sericea. Har- nisch observed that in this species the tegmen remains outside the female genitalia during copulation, acting as a grasping organ and lever. This case is quite similar to Bruchidae as DUNGELHOEF & SCHMITT (2006) show, but they observed slight movements in Acanthoscelides- obtectus-parameres during copulation. Similar move- ments of the parameres have been observed in the carabid beetle Pasimachus punctulatus (ALEXANDER 1959) and in the coccinellid beetle Cycloneda san- guinea (EBERHARD 1985). These movements could serve as a kind of stimulation, supporting the “cryptic- female-choice”-hypothesis (EBERHARD 1985, 1996). KINGSOLVER (1970) observed that in Mimosestes sallaei (Bruchidae) the parameres are known to enter the fe- male vaginal tract during copulation. In all other 254 bruchids investigated, the parameres obviously “serve as guides to position the median lobe opposite the vagi- nal opening.” EBERHARD (1993) examined the functional role of pa- rameres in the scarabaeid species, Macrodactylus costu- latus. He described a “foot-in-door-sac” that is inflated right beneath the apex of the parameres and thus occu- pies the entrance to the female genitalia and can trigger the vulva to open. 2. MATERIALS AND METHODS We examined dry museum specimens (from the ZFMK collection) and some field-collected fresh specimens. These latter specimens were collected mostly in summer 2002 near Dahlem and Blankenheim in the Eifel moun- tain region (about 70 km SW of Cologne). The structures of the aedeagi of 1-5 specimens of the following species have been studied: Cerambycidae Cerambycinae: Stenopterus rufus (Linnaeus, 1767), Obrium brunneum (Fabricius, 1792), Pyrrhidium san- guineum (Linnaeus, 1758), Plagionotus detritus (Lin- naeus, 1758) Lamiinae: Dorcadion pedestre (Poda, 1761), Acantho- cinus aedilis (Linnaeus, 1758), Agapanthia villosoviri- descens (Degeer, 1775) Lepturinae: Dinoptera collaris (Linnaeus, 1758), Carilia virginea (Linnaeus, 1758), Grammoptera ruficornis (Fab- ricius, 1781), Alosterna tabacicolor (Degeer, 1775), An- astrangalia dubia (Scopoli, 1763), Stictoleptura scutellata (Fabricius, 1781), Stictoleptura maculicornis (Degeer, 1775), Stictoleptura rubra (Linnaeus, 1758), Rutpela maculata (Poda, 1761), Leptura quadrifasciata Linnaeus, 1758, Stenurella melanura (Linnaeus, 1758) Prioninae: Prionus coriarius (Linnaeus, 1758) Spondylidinae + Aseminae: Spondylis buprestoides (Linnaeus, 1758), Asemum striatum (Linnaeus, 1758), Arhopalus rusticus (Linnaeus, 1758), Tetropium casta- neum (Linnaeus, 1758) Chrysomelidae Chrysomelinae: Timarcha tenebricosa (Fabricius, 1775) Donaciinae: Donacia vulgaris Zschach, 1788 Orsodacninae: Orsodacne cerasi (Linnaeus, 1758) In addition, many drawings from the literature were ex- amined in order to reach a wider basis for comparison. Illustrations of male genitalia of the following taxa were included in our analysis: Bonner zoologische Beitráge 54 (2005) Cerambycidae Anoplodermatinae — 5 taxa (DIAS 1984, 1986, 1987) Cerambycinae — 27 taxa (BAHILLO DE LA PUEBLA & ITURRONDOBEITIA 1996; FRAGOSO 1985a; FRANCE- SCHINI 2002; IUGA & ROSCA 1961; KUMAR & VERMA 1980; MONNÉ & NAPP 2000; MOURA & GALILEO 1992; NHSATO 1987; SHARP € MUIR 1912; WANG 1995; WU & JIANG 1993; ZIA 1936) Lamiinae — 53 taxa (BAHILLO DE LA PUEBLA & ITUR- RONDOBEITIA 1996; BENSE 1995; BREUNING & TEOCCHI 1977; IUGA & ROSCA 1961; JULIO 1999; MARTINS & GALILEO 1998; SAMA 1988; SHARP & Muir 1912: TSHERNYSHEV & DUBATOLOV 2000; TYSON 1973; VER- DUGO PAEZ 1994; WU & JIANG 1989; ZIA 1936) Lepturinae — 22 taxa (BAHILLO DE LA PUEBLA & ITUR- RONDOBEITIA 1996; BENSE 1995; DEVESA REGUEIRO & BAHILLO DE LA PUEBLA 2003; IUGA & ROSCA 1961; KI- NEL 1931; OHBAYASHI 1970; VILLIERS 1982) Oxypeltinae — 2 taxa (FRAGOSO 1985b) Parandrinae — | taxon (GILBERT 1952) Philinae — 6 taxa (WU & JIANG 2000) Prioninae — 73 taxa (IUGA & ROSCA 1961; QUENTIN & SIMONETTA 1992; QUENTIN & VILLIERS 1977, 1978, 1981, 1983; ZIA 1936) Spondylidinae + Aseminae — 2 taxa (BAHILLO DE LA PUEBLA 1991) Vesperinae — 2 taxa (BENSE 1995) Chrysomelidae Aulacoscelidinae — 4 taxa (JOLIVET 1957; SUZUKI 1994) Bruchidae — 10 taxa (KINGSOLVER 1970; SCHMITT 1985a; TERAN & MURUA DE L’ARGENTIER 1981; ZIA 1936) Chrysomelinae (Timarcha) — 4 taxa (IABLOKOFF- KHNZORIAN 1966; JOLIVET 1957; POWELL 1941; SHARP & MUIR 1912) Donaciinae — 8 taxa (HARNISCH 1915; IABLOKOFF- KHNZORIAN 1966; POWELL 1941; SCHMITT 1985a, 1985b; SHARP & MUIR 1912; ZIA 1936) Megalopodinae — 3 taxa (JOLIVET 1957; ZIA 1936) Orsodacninae — | taxon (POWELL 1941) Palophaginae — 2 taxa (KUSCHEL & MAY 1990) Sagrinae — 18 taxa (JOLIVET 1957; MANN & CROWSON 1991; SCHMITT 1985b; ZIA 1936) Zeugophorinae — 2 taxa (CHUJO 1952; IABLOKOFF- KHNZORIAN 1966) Lasse HUBWEBER & Michael SCHMITT: Parameres in Chrysomeloidea In total the parameres of 216 taxa of Cerambycidae and 55 taxa of Chrysomelidae have been compared. Genitalia were dissected using forceps and insect pins under a WILD M3 Z stereoscope. The genitalia were placed in 80% ethanol for 20 min and then in 100% ethanol until dissection. Finally, they were treated with ultrasound to complete the cleaning process. In preparation for scanning electron microscopy, the air- dried genitalia were coated with gold in an ANATECH HUMMER VII and examined with an HITACHI S- 2460 N (SEM). tu Un Nn 3. RESULTS 3.1. Characters of the parameres The first character we examined was the degree of scle- rotization of the aedeagus, including the tegmen which varies from extreme, with nearly black pigmentation ip lepturine beetles of the genera Leptura, Stictoleptura, Rutpela and Stenurella and in Agapanthia (Laminae) to weak with light yellow in other lepturine genera (Alo- sterna, Carilia, Dinoptera, Grammoptera). Most of the long-horned species studied have reddish-brown aedeagi, representing a medium sclerotized state. In chrysomelids, the species studied have poor to medium degrees of sclerotization of their aedeagi with light yel- low to brown colours. Fig. 1-4. Parameres of Rutpela maculata (1), Leptura quadrifasciata (2), Spondylis buprestoides (3), and Obrium brunneum (A). The second character we measured was the relative pa- ramere length, which is herein referred to only as the non-fused portion of each paramere or lobe. The pa- ramere width was measured at the mid point of the non- fused portion to its lateral extension and then was re- lated to the overall length of the lobes. Sometimes there is no division of the parameres if they are fused together (Fig. 4). In other cases the parameres are divided, very long and slender (Fig. 2). Within some subfamilies of Cerambycidae, it is possible to separate some genera from others with the help of this character. There is only one known genus in the Cerambycidae (Stenhomalus) without parameres (WU & JIANG 1993). Within the taxa of this family studied, only some genera of Cerambyci- nae (Callimellum, Hybodera, Obrium, Skeletodes and Stenopterus) and the Oxypeltinae — altogether only 5 % of the cerambycid taxa studied so far — show totally fused parameres. Within Chrysomelidae, the parameres in Donaciinae, Timarcha, Zeugophorinae and some Bruchidae are fused, but in the other groups, like Orso- dacninae, Sagrinae and most Bruchidae, they are di- vided. Parameres are fused in 34 % of the paramere- possessing chrysomelid taxa we studied. However, even in the chrysomelid groups with non-fused parameres, these structures are comparatively short. The mean rela- tive length of all taxa with non-fused parameres is 3.2 in 256 Bonner zoologische Beitráge 54 (2005) Cerambycidae (standard deviation =1.4; n=194) and 1.6 in Chrysomelidae (standard deviation = 0.9; n=34). Only two of the chrysomelids (both from Aulacoscelid- inae) have parameres with a relative length of 4, all other taxa | to 3. In both families, Cerambycidae and Chrysomelidae, there are taxa with exceptionally long and slender parameres, which were not considered in calculating the mean values, because they would have disturbed the other data: The Madagascan Lepturinae of the genus Mastododera and related genera (VILLIERS 1982) with a relative length of 13 (other cerambycids not higher as 7) and an undetermined Bruchus (ZIA 1936) with a relative length of 32 (other chrysomelids not higher as 4). The third character we considered was the orientation of the lobes, or the direction of the apex. They can be shaped like an arrow (Fig. 3), V-shaped (Fig. 2) or cling tightly next to each other (Fig. 1). Most cerambycids have converging lateral lobes, but most subfamilies show two or three conditions. Among chrysomelids with non-fused parameres, only most Sagrinae and some Bruchidae have divergent parameres. Only one species of Sagrinae has parameres which stretch out parallel. All Megalopodinae, Palophaginae, Aulacoscelidinae and Orsodacninae studied have converging parameres. Additionally, the apex of the parameric lobes can look ragged or cut (Fig. 1), pointed (Fig. 3) or broadly rounded (Fig. 2). Most cerambycid parameres are pointed, but at least in Lepturinae, there are also some species with ragged and broadly rounded lateral lobes. Chrysomelid beetles show in addition to pointed apices of lateral lobes, protracted parameres (in Orsodacninae). Hubweber (unpubl. data) divided the number and den- sity of setae on the parameres into three different char- acters, according to their position (apical, ventral and dorsa-lateral). Probably the best character of these is the number and density of dorsa-lateral setae. All studied == 30m === species of Lepturinae (Cerambycidae) lack setae at this region (Fig. 2), except one species with a few sparse se- tae (Fig. 1), while all species of the other long-horn sub- families studied possess setae dorsalaterally (Fig. 3). On the other hand, ventral setae are widely variable, even between very closely related groups. The leaf beetle species examined possess long setae at the apex of the parameres, but only a few taxa are known with dorsa- lateral or ventral setae. The tegminal ring can either be narrow (Fig. 2) or wide (Fig. 3). “Wide” means that the lateral gap between the ring and median lobe is at least as wide as the ring itself. In cerambycid beetles, most lepturine species have a narrow tegminal ring, while most (but not all) species of the other subfamilies have a broad ring. In Ceramby- cids, 39 % of the taxa studied (n=37) have a wide teg- minal ring, while only 19 % of chrysomelid taxa studied (n=26) show this condition. Additionally, the connection of between the parameres and the basal piece is either angular (Fig. 3) or rounded (Fig. 2). It is not easy to distinguish between “angled” or “rounded”, consequently if there is a sudden bend we call it “angled”. In the longicorn beetles studied (n=68), 44 % show the angled condition, while only 7 % or two species, Amblycerus robiniae (KINGSOLVER 1970) and Orsodacne cerasi (own observation), of the leaf beetles studied (n=26) show this condition. 3.2. Presumed glandular openings A recent discovery is tiny holes near the apex of the pa- rameres of Donacia (Fig. 5). We suggest that these are most probably not sensilla, because they do not contain traces of setae. Similar, but not so many holes are found on the parameres of Prionus (Fig. 6), Stenopterus and Zeugophora. In Alosterna none such holes are found on the parameres. — 8 um —— Fig. 5-6. (5) Apex of the paramere of Donacia vulgaris with tiny holes, (6) Tiny hole on paramere of Prionus coriarius. Lasse HUBWEBER & Michael SCHMITT: Parameres in Chrysomeloidea 257 — Mi pan — Fig. 7-8. (7) Sensilla on the median lobe of Donacia vulgaris, (8) Sensilla on the apex of the median lobe of Orsodacne cerasi. HAMMOND (1972) described true, peg-liked setae of un- known function on the parameres of Staphylinidae. In contrast to the holes on the parameres, sensilla with a minute seta can be found along the inner margin of the median lobe in Donacia and Orsodacne (Figs. 7 and 8). These campaniform sensilla look similar as those ob- served on the median lobe of Platypodidae (EPILA- OTARA & TRIPLEHORN 1990). 4. DISCUSSION We suggest that the sclerotization in Cerambycidae is thicker than in Chrysomelidae generally, because of thicker sclerotization of the female vaginal tract. Many species of long-horned beetles lay their eggs just be- neath the bark of trees and presumably a more stout or heavily sclerotized ovipositor is required and — possibly —amore thickly sclerotized aedeagus, too. The mean value of the relative length of the lateral lobes through 194 cerambycid species is 3.2. We know only a few chrysomelid species from our own investigations or from drawings in the literature with a relative length greater than 3. Of course, there is one subfamily among Chrysomelidae with long and narrow parameres, the Donaciinae, but these parameres are fused. We can also summarise that the parameres of Cerambycidae are gen- erally more deeply divided than those of Chrysomeli- dae. Obviously, the parameres of Chrysomelidae bear less dorsolateral and ventral setae than Cerambycidae. We have no proper explanation for this observation. How- ever, if KINGSOLVER’s (1970) hypothesis holds true — that parameres outside the female genital tract are used to position the male genitalia — then it may be that the setae on the parameres perform a sensorial function. For example they could sense the position of the median lobe inside the female or press on the sternites of the female to facilitate the release of sperm to the proper re- gions inside the female. What is the function of the tiny holes on the parameres? One possibility is that they serve as glandular openings, but what could they secrete? Striking is the fact that they are more numerous on the parameres of Donacia, a beetle with a semi-aquatic mode of life. Therefore it may be they secrete a hydrophobic substance. One question remains: Why are parameres divided in most of the groups studied, but undivided or totally re- duced in some chrysomelids and in some Cerambycinae like Obrium and Stenhomalus? Possibly the function of these two states is quite different, for example the undi- vided condition serves like a lever in Donaciinae, as de- scribed by HARNISCH (1915) and in divided parameres they are sensory, as suggested by DUNGELHOEF & SCHMITT (2006), or stimulatory, as assumed by EBER- HARD (1985), and others. CROWSON & CROWSON (1996) suggested that the parameres have been reduced in some groups because of the problems of copulation on an un- stable leaf surface. However, nearly all cerambycid bee- tles have parameres and many of them copulate on an unstable flower or leaf surface. Future studies should include a comprehensive compari- son of paramere morphology in more closely related cerambycids and chrysomelids. Because the taxa stud- ied may not be representative, or taxa with especially noticeable parameres could have been overlooked in this study, we recommend selecting paired groups of sister taxon. In addition, live beetles in copula should be observed to solve the riddle of the parameres’ function. In particular, the function of the tegmen in the closely related genera of Obrium and Stenhomalus could prove very interesting because of the obvious reduction and loss of parameres in these groups. The Madagascan spe- cies of Lepturinae and the bruchid species also have in- 258 Bonner zoologische Beitráge 54 (2005) teresting morphologies with very long and narrow pa- rameres. Histological and TEM-studies should clarify the structure of the tiny holes found on the apex of the parameres. Acknowledgements. We thank Susanne Düngelhoef (ZFMK, Bonn) for discussion on the question of functional morphology of the parameres and two anonymous referees for helping to improve our manuscript. Bradley Sinclair (ZFMK, Bonn) kindly checked the manuscript for lan- guage corrections. Lasse Hubweber is grateful for funding of his doctoral studies by the Cusanuswerk. REFERENCES ALEXANDER, R. D. 1959. The courtship and copulation of Pasimachus punctulatus Haldeman (Coleoptera: Carabidae). Annals of the Entomological Society of America 52: 485. BAHILLO DE LA PUEBLA, P. 1991. La armadura genital mas- culina en la diagnosis especifica de los Arhopalus Ibericos (Coleoptera, Cerambycidae). Estudios del Museo de Ciencias Naturales de Alava 6: 115-119. BAHILLO DE LA PUEBLA, P. & ITURRONDOBEITIA, J.C. 1996. Cerambicidos (Coleoptera, Cerambycidae) del País Vasco. Cuadernos de Investigacion Biologica (Bilbao) 19: 1-244. BENSE, U. 1995: Bockkáfer: illustrierter Schliissel zu den Cerambyciden und Vesperiden Europas. Margraf, Weikersheim. BREUNING, S. & TEOCCHI, P. 1977. Création de la tribu des Aderpasini, nov. Révision et bionomie des especes des genres Aderpas Thoms. et Ancylonotopsis Br. (Co- leoptera Cerambycidae Lamiinae). Bulletin de l'Institut Fondamental d'Afrique Noire, Série A 39: 142-168. CHEN, S. 1985. Phylogeny and classification of the Chry- someloidea. Entomography 3: 465-475. CROWSON, R. A. & CROWSON, E. A. 1996. The phyloge- netic relations of Galerucinae — Alticinae. Pp. 97-118 in: JOLIVET, P. H. A. & Cox, M. L. (eds.) Chrysomeli- dae Biology, Vol. 1: The Classification, Phylogeny and Genetics. SPB Academic Publishing, Amsterdam. DEVESA REGUEIRO, S. & BAHILLO DE LA PUEBLA, P. 2003. Primer registro de Paracorymbia maculicornis (De- geer, 1775) en Galicia (N.W. de la Península Ibérica) con datos sobre su armadura genital (Coleoptera, Cerambycidae). Boletin de la SEA 32: 83-86. Dias, M. M. 1984. Revisäo da subfamilia Anoplodermat- inae. Parte I. Tribo Anoplodermatini. Género Migdolus Westwood, 1863 (Coleoptera, Cerambycidae). Revista Brasileira de Entomologia 28: 507-535. Dias, M. M. 1986. Revisäo da subfamilia Anoplodermat- inae. Parte II. Tribo Anoplodermatini. Géneros: Acan- thomigdolus Bruch, 1941, Paramigdolus, gen. n., Anoploderma Guérin-Méneville, 1840, Sypilus Guerin-Meneville, 1840 (Coleoptera, Cerambycidae). Revista Brasileira de Entomologia 30: 115-139. Dias, M. M. 1987. Revisäo da subfamilia Anoplodermat- inae. Parte III. Tribo Anoplodermatini. Géneros: Cher- rocrius Berg, 1898, Hypocephalus Desmarest, 1832 (Coleoptera, Cerambycidae). Revista Brasileira de En- tomologia 31: 101-112. DUNGELHOEF, S. & SCHMITT, M. 2006. Functional mor- phology of copulation in Chrysomelidae — Criocerinae and Bruchidae (Insecta: Coleoptera). Bonner zoolo- gische Beitráge 54: 201-208. EBERHARD, W. G. 1985. Sexual Selection and Animal Genitalia. Harvard University Press, Cambridge Mas- sachusetts & London. EBERHARD, W. G. 1993. Copulatory courtship and genital mechanics of three species of Macrodactylus (Coleop- tera Scarabaeidae Melolonthinae). Ethology, Ecology and Evolution 5: 19-63. EBERHARD, W. G. 1996. Female control: Sexual selection by cryptic female choice. Princeton University Press, Princeton. EPILA-OTARA, J. S. & TRIPLEHORN, C. A. 1990. Mechano- receptors on the male genitalia of Platypodidae (Co- leoptera): a pregamous isolation mechanism? Journal of African Zoology 104: 345-350. FRAGOSO, S. A. 1985a. The terminalia as a basis for the classification of Cerambycidae (Coleoptera) subfami- lies. Part I. Terminology and genital morphology of Cerambyx cerdo L. Revista Brasileira de Entomologia 29: 125-134. FRAGOSO, S. A. 1985b. The terminalia as a basis for the classification of Cerambycidae (Coleoptera) subfami- lies. Part II, Oxypeltinae. Revista Brasileira de Ento- mologia 29: 165-168. FRANCESCHINI, A. F. 2002. Revisáo do género Appula (Coleoptera, Cerambycidae, Cerambycinae, Elaphi- diini). Iheringia, Série Zoologia 92: 5-40. GILBERT, E. E. 1952. The homologies of the male genitalia of Rhynchophora and allied Coleoptera. Annals of the Entomological Society of America 45: 633-637. HAMMOND, P. M. 1972. The Micro-Structure, Distribution and Possible Function of Peg-Like Setae in Male Co- leoptera. Entomologica Scandinavica 3: 40-54. HARNISCH, W. 1915. Uber den mánnlichen Begattungsap- parat einiger Chrysomeliden. Ein Beitrag zur Phy- logenie des Copulationsapparates der Käfer. Zeit- schrift fiir wissenschaftliche Zoologie 114: 1-94. IABLOKOFF-KHNZORIAN, S. M. 1966. Considérations sur l’edeage des Chrysomelidae et son importance phy- logénique. L'Entomologiste 22: 115-137. IUGA, V. G. & ROSCA, A. 1961. La morphologie de apex abdominal chez les Cerambycidae (Col.), comparé a celui des Hymenopteres. Travaux du Museum d’his- toire naturelle «Grigore Antipa» 3: 101-156. JOLIVET, P. 1957. Recherches sur I'aile des Chry- someloidea (Coleoptera). Mémoires Institut Royal des Sciences Naturelles de Belgique, 2. Série 57: 1-180. JULIO, C. E. de A. 1999. Revisáo do género Scleronotus White, 1855 (Coleoptera, Cerambycidae, Lamiinae, Acanthoderini). Boletim do Museu Nacional do Rio de Janeiro, N.S., Zoologia 406: 1-35. KINEL, J. 1931. Leptura inexspectata Janss. & Sjöb. na ziemiach polskich oraz pozycja systematyczna tej formy. Bulletin entomologique de Pologne 10: 189- 196. KINGSOLVER, J. M. 1970: A study of the male genitalia in Bruchidae (Coleoptera). Proceedings of the Entomo- logical Society of Washington 72: 370-386. KUMAR, D. & VERMA, K. K. 1980. Aedeagal musculature in Phytophaga (Coleoptera). Journal of Natural History 14: 237-270. Lasse HUBWEBER & Michael SCHMITT: Parameres in Chrysomeloidea 259 KUSCHEL, G. & MAY, B. M. 1990: Palophaginae, a New Subfamily for Leaf-Beetles, Feeding as Adult and Larva on Araucarian Pollen in Australia (Coleoptera: Megalopodidae). Invertebrate Taxonomy 3: 697-719. MANN, J. S. & CROWSON, R. A. 1991. Some observations on the genitalia of Sagrinae (Coleoptera: Chrysomeli- dae). Pp. 35-60 in: ZUNINO, M., BELLES, X. & BLAS, M. (eds.) Advances in Coleopterology. AEC, Barce- lona. MARTINS, U. R. & GALILEO, M. H. M. 1998. Revisáo da tribo Aerenicini Lacordaire, 1872 (Coleoptera, Ceram- bycidae, Lamiinae). Arquivos de Zoologia 35: 1-133. MONNÉ, M. L. & NAPP, D. S. 2000. Revisäo do género Ceralocyna (Coleoptera, Cerambycidae, Cerambyci- nae, Trachyderini, Ancylocerina). Iheringia, Série Zo- ologia 88: 103-137. Moura, L. de A. & GALILEO, M. H. M. 1992. Genitalia masculina e feminina de Dorcacerus barbatus (Oliv- ier, 1790) (Coleoptera, Cerambycidae, Cerambycinae, Trachyderini). Iheringia, Série Zoologia 72: 135-139. NIISATO, T. 1987. A new Cyrtoclytus (Coleoptera: Ceram- bycidae) from Northern Thailand. Coleopterists’ Bul- letin 41: 297-301. OHBAYASHI, N. 1970. On some cerambycid-beetles from the Ryukyu Islands. Bulletin of the Japan Entomologi- cal Academy 5: 1-4. POWELL, E. F. 1941. Relationships within the Family Chrysomelidae (Coleoptera) as Indicated by the Male Genitalia of Certain Species. American Midland Natu- ralist 25: 148-195. QUENTIN, R. M. & SIMONETTA, J. 1992. Révision du genre Cantharocnemis Serville (Coleoptera: Cerambycidae). Les sous-genres Cantharocnemis sensu stricto et Can- tharofoedus Gilmour. Annales de la Société Ento- mologique de France N.S. 28: 83-100. QUENTIN, R. M. & VILLIERS, A. 1977. Genera et catalogue raisonné des Prioninae africains. I.Ergatini (Col. Cerambycidae). Annales de la Société Entomologique de France N.S. 13: 509-523. QUENTIN, R. M. & VILLIERS, A. 1978. Genera et catalogue raisonné des Prioninae africains. II. Macrotomini (Col. Cerambycidae). Annales de la Société Entomologique de France N.S. 14: 207-274. QUENTIN, R. M. & VILLIERS, A. 1981. Les Macrotomini de l’ancien monde (région Ethiopienne exclue). Genera et catalogue raisonné (Col. Cerambycidae Prioninae). Annales de la Société Entomologique de France N.S. 17: 359-393. QUENTIN, R. M. & VILLIERS, A. 1983. Genera et catalogue raisonné des Prioninae africains. III. Acanthophorini (Col. Cerambycidae). Annales de la Société Ento- mologique de France N.S. 19: 79-100. SAMA, G. 1988. Phytoecia flavipes (Fabricius, 1801) a valid species, not a synonym of P. rufipes (Olivier, 1795) (Insecta, Coleoptera, Cerambycidae). Steen- strupia 14: 121-125. SCHMITT, M. 1985a. Versuch einer phylogenetisch- systematischen Analyse der Criocerinae (Coleoptera, Chrysomelidae). Zoologische Beitráge N.F. 29: 35-85. SCHMITT, M. 1985b. On the Phylogeny of the Criocerinae (Coleoptera, Chrysomelidae). Entomography 3: 393-401. SHARP, D. & Muir, F. 1912. The comparative anatomy of the male genital tube in Coleoptera. Transactions of the Royal Entomological Society of London 60: 477- 642. SUZUKI, K. 1994. The systematic Position of the Subfamily Aulacoscelinae (Coleoptera: Chrysomelidae). Pp. 45- 59 in: FURTH, D.G. (eds.) Proceedings of the Third Iy- ternational Symposium on the Chrysomelidae, Beijing, 1992. Backhuys, Leiden. TERAN, A. L. & MURUAGA D'ARGENTIER, S. 1981. Obser- vaciones sobre Bruchidae (Coleoptera) del noroeste Argentino. IV. Estudios morfologicos y biologicos de Amblycerus hoffmannseggi (Gyll.), Acanthoscelides comptus Kingsolver y Merobruchus bicoloripes (Pic). Acta Zoologica Lilloana 36/2: 61-84. TSHERNYSHEV, S. E. & DUBATOLOV, V. V. 2000. A new species of longhorn-beetle from East Siberia (Insecta: Coleoptera: Cerambycidae). Reichenbachia 33: 385- 389. Tyson, W. H. 1973. The Spalacopsis of the West Indies and America north of Mexico (Coleoptera: Ceramby- cidae). Coleopterists’ Bulletin 27: 117-137. VERDUGO PAEZ, A. 1994: Los /berodorcadion (Breuning, 1943) des suroeste ibérico. Anatomía des las fases in- maduras, ciclo vital, ecología y distribución (Coleop- tera : Cerambycidae). Zapateri 4: 87-103. VERMA, K. K. 1996. Inter-subfamily relations among Chrysomelidae (Coleoptera) as suggested by organiza- tion of the male genital system. Pp. 317-351 in: JOLI- VET, P. H. A. £ COX, M.L. (eds.) Chrysomelidae Biol- ogy, Vol. 1: The Classification, Phylogeny and Genetics. SPB Academic Publishing, Amsterdam. VILLIERS, A. 1982. Démembrement du genre Mastododera Thomson (Coleoptera, Cerambycidae, Lepturinae). Annales de la Société Entomologique de France N.S. 18: 129-162. WANG, Q. 1995. The Australian longicorn beetle genus Skeletodes Newman (Coleoptera: Cerambycidae). Co- leopterists’ Bulletin 49: 109-118. Wu, W.-W. & JIANG, S.-N. 1989. A taxonomic study of the male genitalia of the genus Anoplophora Hope (Col. Cerambycidae). Acta Entomologica Sinica 32: 211- 220. Wu, W.-W. & JIANG, S.-N. 1993. On the male genitalia of the genus Stenhomalus (Coleoptera: Cerambycidae). Acta Entomologica Sinica 36: 81-84. Wu, W.-W. & JIANG, S.-N. 2000. A taxonomic study of the male genitalia of some philid beetles with one new species in China (Coleoptera: Cerambycoidea). Acta Entomologica Sinica 43: 78-87. ZIA, Y. 1936. Comparative studies of the male genital tube in Coleoptera Phytophaga. Sinensia 7: 319-352. Authors’ address: Lasse HUBWEBER (corresponding author), Michael SCHMITT, Zoologisches Forschungs- museum Alexander Koenig, Adenauerallee 160, D-53113 Bonn, Germany. E-mail: 1 hubweber.zfmk(Vuni- bonn.de, m.schmitt@uni-bonn.de Bonner zoologische Beitráge Band 54 (2005) | Heft 4 | Seiten 261—269 Bonn, Oktober 2006 Leaf Beetles (Insecta: Coleoptera: Chrysomelidae) Suffer From Feeding on Fern Leaves’ / Michael SCHMITT? & Sigrun BOPP”? 2) E Zoologisches Forschungsmuseum Alexander Koenig, Bonn, Germany ~ Universität Ulm, Abteilung Systematische Botanik und Ökologie, Ulm, Germany Abstract. Two age groups of larvae of Agelastica alni (Linnaeus, 1758) and Phratora vitellinae (Linnaeus, 1758) were provided with treated or untreated leaves of their actual host plant (4/nus glutinosa or Salix spp., respectively) or of the ferns Athyrium filix-femina, Dryopteris austriaca and D. filix-mas. “Treated” leaves were covered with an aqueous ex- tract of host plant or fern leaves. Mortality of young larvae (one-day old) of A. a/ni was significantly higher when pro- vided with treated or untreated fern leaves and treated food plant leaves compared to untreated food plant leaves. Mor- tality of older larvae (six-day old) was significantly lower than that of young larvae. Young larvae did not feed on fern leaves at all, while older larvae ingested material from treated fern leaves, however at low rates. Phratora vitellinae in- dividuals showed abnormal elytra and irregular melanisation patterns when they had taken up fern substances. In both species, the larvae produced less defensive secretion if they had ingested an extract of Athyrium filix-femina. Key words. Agelastica alni, Phratora vitellinae, Filicatae: Aspidiaceae, phytophagy, larval development 1. INTRODUCTION There are 12,500 species of ferns and 230,500 species of seed plants (SPECK 1992). Some 140,000 species of Phytophaga (= Pseudotetramera = Curculionoidea + Cerambycidae + Bruchidae + Chrysomelidae; Lexikon der Biologie, Urania-Tierreich) mean a proportion of one species of pseudotetramerous beetles per 1.7 species of spermatophyte plant, but only 105 species of Phyto- phaga are known to feed on ferns (BALICK et al. 1978), producing a proportion of 1:119.0. According to HENDRIX (1980) of 726,000 species of insects known at that time, 338,000 are phytophagous (1:0.7), but only 465 feed on ferns (1:26.9). Of the 413 arthropod species reported as found on ferns, 93 attack bracken (Preridium aquilinum) exclusively, making the proportion 1:33.3. Even if these numbers must be corrected for uneven numbers of individuals or/and different availability of biomass, it is clear that there are much less leaf beetles (Chrysomelidae) feeding on ferns than could be ex- pected according to species numbers. There might be regional exceptions (e.g., Veracruz, Mexico: BALICK et el. 1978; Hawaii: SWEZEY 1922). Especially arthropods with chewing mouthparts are remarkably underrepre- sented on ferns as compared to the large percentage of insects with sucking mouthparts (Thysanoptera, Heter- optera, Homoptera; HENDRIX 1980; HILL 1998; D'Ro- ZARIO & BERA 2003). ' Paper presented to the 6" International Symposium on the Chry- somelidae, Bonn, Germany, May 7, 2004. Data from bracken (e.g., MARTINS et al. 1995; GILMAN & COOPER-DRIVER 1998) can be generalised only with care since this fern species is (1) certainly the most abundant fern species on earth, and (2) is regarded a pest since it causes illness to cattle when fed. Thus, bracken is definitively the most intensively studied fern species. In other taxa, for example, Lithinine moths, there might have been a phylogenetic adaptation towards fern feed- ing (WEINTRAUB et al. 1995). The sawfly Strongy- logaster osmundae (Takeuchi, 1941) (Hymenoptera: Tenthredinidae) develops obligatorily on the osmund fern, Osmunda japonica. In leaf beetles, however, there is no indication that any species is especially adapted to the exploitation of this food resource. For example, WINTERBOURN (1987) found beetles of seven families on bracken, amongst them no leaf beetles, a result also supported by MESIBOV (2001). MUKHOPADHYAY & THAPA (1994) list 19 beetle species, among them seven species of Chrysomelidae as “associated with ferns”, however without providing any information as to the in- tensity of this “association”. Since among the seven leaf beetles listed by MUKHOPADHYAY & THAPA there is also Aspidimorpha sanctaecrucis (Fabricius, 1792), which is definitely restricted to Convulvulaceae (GHATE et al. 2003), one should draw conclusions very cau- tiously. Possible reasons for the strikingly lower number of phytophagous beetles on ferns than on angiosperms could be that ferns are either toxic, or have repellents, or lack attractants. 262 In order to shed some light on these questions for three species of Central European ferns (Athyrium filix-femina (Lady Fern), Dryopteris austriaca (Wood Fern) and D. filix-mas (Male Fern), we conducted feeding experi- ments with two species of leaf beetles, Agelastica alni (Linnaeus, 1758) and Phratora vitellinae (Linnaeus, 1758). 2. MATERIALS AND METHODS Fifteen adult individuals of Agelastica alni were col- lected from Common Alder (4/nus glutinosa) in the sur- roundings of Freiburg im Breisgau, Germany. They were kept in plastic containers on a layer of gypsum and fed with fresh alder leaves. Their eggs were collected and the hatched larvae were fed on fresh leaves of their original food plant until they were used in the tests. We conducted two series of tests: (1) with “young” — 1.e., freshly hatched — larvae, and (2) with “old” — 1.e., six- day old — larvae. In both age groups feeding traces and Table 1. Experimental Settings Test Number Kind of Food Offered Bonner zoologische Beitráge 54 (2005) mortality were registered on day 3, 8, and 10 after the beginning of the experiment. Adults of Phratora vitellinae were collected from wil- lows and poplars growing in a park area on the outskirts of Freiburg im Breisgau. They were kept and treated as the individuals of Age/astica alni. However, since con- siderably less larvae of Phratora vitellinae could be ob- tained, meaningful statistical analysis was not possible. Leaves of Athyrium filix-femina, Dryopteris austriaca and D. filix-mas were collected in a forest close to the city of Freiburg im Breisgau (“Sternwald”), always from the same area. Aqueous extracts were made from fresh leaves and from those dried for four weeks at room temperature. The leaves were put into tab water for 30 minutes and then squeezed with broad forceps in order to gain not only water-soluble but also lipophilous substances. The test leaves were soaked with the respec- tive extracts for 30 minutes in order to mask their con- tact chemical properties. Normal food plant (f.p.) masked with extract from dried Drvopteris filix-mas (D.f.) Normal food plant (f.p.) masked with extract from dried Dryopteris ausriaca (D.a.) Dried D. filix-mas masked with extract from the normal food plant | 2 5 Normal food plant (f.p.) masked with extract from dried Athyrium filix-femina (A.f.) 4 5 Dried D. austriaca masked with extract from the normal food plant 6 Dried A. filix-femina masked with extract from the normal food plant 7 Dried D. filix-mas, swelled in tab water 8 Dried D. austriaca, swelled in tab water 9 Dried A. filix-femina, swelled in tab water 10 Dried normal food plant, swelled in tab water 11 Normal food plant masked with extract from fresh Dryopteris filix-mas [2 Normal food plant masked with extract from fresh Dryopteris ausriaca 13 Normal food plant masked with extract from fresh Athyrium filix-femina 14 Fresh D. filix-mas masked with extract from the normal food plant 15 Fresh D. austriaca masked with extract from the normal food plant 16 Fresh A. filix-femina masked with extract from the normal food plant 7 Fresh D. filix-mas, swelled in tab water 18 Fresh D. austriaca, swelled in tab water 19 Fresh A. filix-femina, swelled in tab water 20 Fresh normal food plant, swelled in tab water 21 Fresh D. filix-mas, untreated 22 Fresh D. austriaca, untreated 23 Fresh A. filix-femina, untreated 24 Fresh normal food plant, untreated For each of the two test series 24 setups were designed. The details are given in Table 1. The test larvae were kept under the described conditions until either all were dead or the first individuals tried to pupate (which can easily be recognised as Agelastica alni beetles pupate in the soil. Thus, larvae ready to pupate attempt at digging holes in the gypsum). Phratora vitellinae pupate on the leaves of their food plant, the initiation of their pupation can be observed directly. We used the Statistical Package for Social Sciences (SPSS) to calculate a logistic regression. Probabilities of error were calculated by Chi? or Fisher’s exact test (the latter in cases where n was too low for Chi’). 63 5) Michael SCHMITT & Sigrun Bopp: Leaf Beetles (Chrysomelidae) Feeding Fern “y Aep Joye IBALB] DAI] DOU! OU “SuIpady JO saov.y E pur Z SÁRp uO Aug : “y Aep JOY JeAle] DAI] SOUL OU “SII IIOUI OU 19)/89194) ‘| Aep UO SIDRA) BUIPII} MI : OVAIR] DAT] DOW! OU € ABP JOR ‘sdov1] SuIpoay Moy ¢ Avp 01 dy] : "S39P.1) DOW OU 19IJ89194) *] Aep UO SIDRI Suipsoy MO : “$ Aep WO IBAN] DAI] OU “SIDRA DIOW OU € Aep Jaye ‘Aep tod sa9en Surpsay Moy Alaa € Aep o] dy] : “QBAIL] SAT] QIOW OU y Aep Jaye ‘Aep tad so9e1] Suipsoy sso] ¢ Áep 01 df) : [Je Je sae. SuIpsoay o10ur ou QT Aep Ysnosy} / Aep woy ‘Aep 19d saov.y Surpsay ssa] / Avep 0] dy] : S/bL S/bL 0/6L 0/6L 0/08 0/€8 payeo.nun “dy ysaıy) v7 ST/OL 0/71 (payeayun / y Yysorq) €7 8/L ST/E (payeayun 97 Ys94) TE wel €7/0 r1/6 (a (payonun fq YS914) le 0/07 0/07 0/07 = €/SI WEI 0/61 a (parrems “dy ysor) 07 1/61 O 71/0 (parroms SP ysalg) 61 0/07 O 07/0 O (pay]ams D'( Ysa.) 81 SI/ZI O 81/0 (parroms fa YS914) Él L1/0 O/LI Mo 91/p O Cd yım poyseur fp Usa) 91 61/0 U/LI qa 07/0 (dy qua poyseut 2° YSa1) SI 0€/0 61/11 O) 07/0 Cdy yy poyseut GQ 48914) pl ULl ULl 0/61 al C/LI GL 0/07 = (Sr Ysoy YIM poyseur *d 1) El E/LI E/LI E/LI = p/SI €/91 1/61 a (DG ysoy YIM poyseut “d’ 4) ZI E/LI E/LI 1/61 = 0/61 0/61 0/07 a (fa Ysoy yım poyseut “d° 4) II S/LI Y/81 0/77 = 61/0 EME 1/61 (8 (parroms “dy potiq) Ol 0/07 Mo) 8/6 (pay[ams SF paid) 6 1/61 Mo) Lp (POII PAS pq pag) 8 7/81 Mo) 61/0 Ll xO) (params fq petiq) L 81/0 0/81 a b/p1 Cd qua payseu Sp pong) 9 07/0 1/61 a 7/61 (dy qua poyseu 2° paid) S 17/0 1/07 Mo) 91/1 Cd yim poyseut fq paid) r viel €/p1 1/91 = €1/S 21/9 ULl = CS Y pauıp yyım poyseut 'd’y) € viel viel ULT = S/vI S/pl GLI = (eq palip YIM poyseut 'd’y) (6 UTI Z/EI 01/91 a 6/01 LIT 1/81 B (Sa pauıp yyım payseut “d° 4) [ peap/aare peop/aye = peap/aarye peop/aare peap/aarye peap/aare "ON 01 Ata 8 ded ¢ Avg 01 Ata 8 Áta ¢ Ata 39 L IVAIV] PIO IVAIV] SUNOA (psouynj3 snujy) yuejd pooy ¡eurou dy “spw-xıpf stiajdodig fa vooi4sno staajdodig 0 punwmaf-sapf unidyp y :(99139p SS9] A[qesopisuod e 0] pue) ÁJUO SÁRBP 39) JULOS UO 3UIP99] JO SICA : @ ‘Aep 159) Aue uo Burpaay Jo saov. ou : O “SABp 189] [je UO SIALA]| UO JUIPIAJ JO SIDRA] : m “SUONIPUOI 189] JopUN 14/D DISD/23 YY JO soyeı ALLOY 7 IP L 264 Bonner zoologische Beitráge 54 (2005) 3. RESULTS 3.1. Agelastica alni The results of the feeding experiments are listed in Ta- ble 2. Decreasing total numbers of individuals are due to the fact that some larvae managed to escape from the test boxes. The nearly non-existing mortality in the con- trol sample proves that the experimental conditions were appropriate. In addition to the observed numerical results we found that those larvae who had taken up ex- tract from dried or fresh Athyrium filix-femina produced conspicuously less defensive secretion when disturbed than those larvae fed on untreated normal food plant or on normal food plant masked with extract of the other two fern species. Moreover, duration of larval stage 2 lasted for 12 days when the larvae were fed on normal food plant masked with extract from dried Athyrium filix-femina as compared to nine days in the control group, or 8.5 to 10 days in the other tests with normal but treated food plant. The logistic regression revealed a significant influence of food (which plant matter was offered) and the mask- ing of the test food (with the exception of Dryopteris felix-mas, the extract of which had no significant influ- ence), whereas the treatment of the test food and of the masking (dried and swelled or fresh) did not lead to sig- nificant differences. In Table 3 we present the results of comparisons be- tween different experimental settings, for young and old larvae. Due to the high mortality in some tests, only those data are presented completely where enough lar- vae survived up to day 10. Table 3. Comparison of the mortality rates under different test conditions against control (test no. 24). Probabilities of error cal- culated with Chi? or Fisher’s Exact 3a: Young larvae Test No. Probability of Error | (F.p. masked with dried D.f.) on day 10 > control p < 0.001 2 (F.p. masked with dried D.a.) on day 10 > control p < 0.001 3 (F.p. masked with dried A.f.) on day 10 > control p < 0.001 4 (Dried D.f. masked with f.p.) on day 3 > control p < 0.001 5 (Dried D.a. masked with f.p.) on day 3 > control p < 0.040 6 (Dried A.f. masked with f.p.) on day 3 > control p < 0.001 i (Dried D.f. swelled) on day 3 > control p < 0.001 8 (Dried D.a. swelled) on day 3 > control p < 0.001 9 (Dried A.f. swelled) on day 3 > control p < 0.001 10 (Dried f.p. swelled) on day 10 > control p < 0.001 11 (F.p. masked with fresh D.f.) on day 10 > control n.s. 12 (F.p. masked with fresh D.a.) on day 10 > control p < 0.002 13 (F.p. masked with fresh A.f.) on day 10 > control p < 0.040 14 (Fresh D.f. masked with f.p.) on day 3 > control p < 0.001 15 (Fresh D.a. masked with f.p.) on day 3 > control p < 0.001 16 (Fresh A.f. masked with f.p.) on day 3 > control p < 0.001 17 (Fresh D.f. swelled) on day 3 > control p< 0.001 18 (Fresh D.a. swelled) on day 3 > control p < 0.001 19 (Fresh 4.f swelled) on day 3 > control p < 0.001 20 (Fresh f.p. swelled) no difference to control 21 (Fresh D.f untreated) on day 3 > control p < 0.001 22 (Fresh D.a. untreated) on day 3 > contro p < 0.001 23 (Fresh A.f. untreated) on day 3 > control ns. Michael SCHMITT & Sigrun Bopp: Leaf Beetles (Chrysomelidae) Feeding Fern 265 3b: Old larvae Test No. Probability of Error #23 1 (£p. masked with dried D.f.), 2 (f.p. masked with dried D.a., no differences to control 3 (£.p. masked with dried A.f.) on day 10 4,5,6 4 (Dried D.f. masked with f.p.), 5 (Dried D.a. masked with no differences to control f.p.), 6 (Dried 4.f masked with f.p.) on day 3 , 7 (Dried D.f. swelled) on day 3 > control p < 0.040 8 (Dried D.a. swelled), 9 (Dried A.f. swelled) on day 3 no differences to control 10 (Dried f.p. swelled) on day 10 > control p < 0.040 11 (F.p. masked with fresh D./.), 12 (F.p. masked with fresh D.a.), no differences to control 13 (F.p. masked with fresh A.f.) on day 10: 14 (Fresh D.f masked with f.p.) on day 3 > control p < 0.001 1:5 (Fresh D.a. masked with f.p.) on day 3 > control p < 0.040 16 (Fresh A.f. masked with f.p.) on day 3 > control ns. ¡ty (Fresh D.f. swelled) on day 3 > control p < 0.001 18 (Fresh D.a. swelled), 19 (Fresh 4.f swelled) on day 3: no differences to control 20 (Fresh f.p. swelled) on day 10 > control ns. 21 (Fresh D.f untreated) on day 3 > control p < 0.030 22 (Fresh D.a. untreated) on day 3 > control p < 0.001 23 (Fresh A.f. untreated) on day 3 > control p < 0.001 Table 4. Comparison of the mortality rates of young and old larvae under different test conditions. Probabilities of error calcu- lated with Chi? or Fisher’s Exact Test No. m OM AND NA BWN m = 159) —- © — Ww DD u RR Re Re eR == O0v00 Du 42 Y Y t t 23 Probability of Error (F.p. masked with dried D.f.) young > old on day 10 (F.p. masked with dried D.a.) young > old on day 10 (F.p. masked with dried A.f.) young > old on day 10 (Dried D.f masked with f.p.) young > old on day 3 (Dried D.a. masked with f.p.) young > old on day 3 (Dried A.f. masked with f.p.) young > old on day 3 (Dried D.f swelled) young > old on day 3 (Dried D.a. swelled) young > old on day 3 (Dried 4.f swelled) young > old on day 3 (Dried f.p. swelled) young > old on day 10 (F.p. masked with fresh D.f.) (F.p. masked with fresh D.a.) (F.p. masked with fresh A.f.) (Fresh D.f. masked with f.p.) young > old on day 3 (Fresh D.a. masked with f.p.) young > old on day 3 (Fresh 4.f. masked with f.p.) young > old on day 3 (Fresh D.f swelled) young > old on day 3 (Fresh D.a. swelled) young > old on day 3 (Fresh 4.f swelled) young > old on day 3 (Fresh f.p. swelled) young > old on day 10 (Fresh D.f. untreated) young > old on day 3 (Fresh D.a. untreated) young > old on day 3 (Fresh 4.f untreated) young > old on day 3 p < 0.050 n.s. p < 0.010 p < 0.010 ns. n.s. p < 0.050 p < 0.001 p < 0.002 p<0:010 no difference between young and old on day 10 no difference between young and old on day 10 no difference between young and old on day 10 p<0.001 p<0.001 p < 0.001 p < 0.001 p < 0.001 p < 0.001 ns. p < 0.010 ns. nS: 24 (Fresh fp. untreated) young > old on day 10 p< 0.030 266 Bonner zoologische Beitráge 54 (2005) Table 5. Comparison of mortality rates on the three fern species. Probabilities of error calculated with Chi? or Fisher’s Exact 5a: Young larvae Test No. Probability of Error | (F.p. masked with dried D.f.) > 2 (F.p. masked with dried D.a.) on ns. day 10 3 (F.p. masked with dried A.f.) > 1 (F.p. masked with dried D.f.) on ns. day 10 3 (F.p. masked with dried 4.f.) > 2 (F.p. masked with dried D.a.) on n.s day 10 4 (Dried D.f. masked with f.p.) > 5 (Dried D.a. masked with fp.) on p<0.001 day 3 4 (Dried D.f masked with f.p.) > 6 (Dried A.f. masked with f.p.) on p<0.001 day 3 6 (Dried 4.f masked with f.p.) > 5 (Dried D.a. masked with f.p.) on n.s day 3 8 (Dried D.a. swelled) > 7 (Dried D.f. swelled) on day 3 p < 0.010 9 (Dried 4.f swelled) > 7 (Dried D.f swelled) on day 3 p > 0.050 8 (Dried D.a. swelled) > 9 (Dried 4.f. swelled) on day 3 p < 0.050 22 (Fresh D.a. untreated) > 21 (Fresh D.f untreated) on day 3 n.s. 21 (Fresh D.f. untreated) > 23 (Fresh 4.f untreated) on day 3 p < 0.001 22 (Fresh D.a. untreated) > 23 (Fresh Ay. untreated) on day 3 p < 0.001 5b: Old larvae Test No. Probability of Error | (F.p. masked with dried D.f.) > 2 (F.p. masked with dried D.a.) on nis. day 10 3 (F.p. masked with dried A.f.) > 1 (F.p. masked with dried D.f.) on day — nis. 10 3 (F.p. masked with dried A.f.) > 2 (F.p. masked with dried D.a.) on p< 0.050 day 10 5 (Dried D.a. masked with f.p.) > 4 (Dried D.f masked with f.p.) on =p < 0.050 day 3 4 (Dried D.f masked with f.p.) > 6 (Dried 4.f masked with fp.) on day 3 nis. 5 (Dried D.a. masked with f.p.) > 6 (Dried 4.f masked with f.p.) on — ns. day 3 14 (Fresh D.f. masked with f.p.) > 15 (Fresh D.a. masked with f.p.) on p< 0.020 day 3 14 (Fresh D.f. masked with f.p.) > 16 (Fresh 4.f masked with fp.) on p< 0.010 day 3 15 (Fresh D.a. masked with f.p.) > 16 (Fresh 4.f masked with f.p.) on nas. day 3 17 (Fresh D.f swelled) > 18 (Fresh D.a. swelled) on day 3 p < 0.001 17 (Fresh D.f. swelled) > 19 (Fresh A.f swelled) on day 3 p < 0.001 19 (Fresh 4.f swelled) > 18 (Fresh D.a. swelled) on day 3 n.s. 22 (Fresh D.a. untreated) > 21 (Fresh D.f. untreated) on day 3 p < 0.050 22 (Fresh D.a. untreated) > 23 (Fresh A.f. untreated) on day 3 ns. (Fresh 4.f. untreated) > 21 (Fresh D.f untreated) on day 3 n.s. Comparisons of mortality of young and old larvae are given in Table 4. In some tests, it did not differ signifi- cantly, as long as only numbers of surviving larvae are counted. It was, however, evident that young larvae died more frequently from starvation because they refused to feed on certain food at all. Old larvae took up leaf mat- ter in several tests but died afterwards, either due to in- toxication or from starvation after refusal of further feeding. There was no significant difference in mortality rates when tests with dried against fresh fern leaves were Michael SCHMITT & Sigrun Bopp: Leaf Beetles (Chrysomelidae) Feeding Fern 267 compared. Mortality was significantly higher on dried as compared to fresh food plant, and on normal food plant masked with extract from dried as compared to fresh Dryopteris filix-mas and Athyrium filix-femina. In order to reveal possible differences between the three fern species, the respective settings have been com- pared. The results are given in Table 5. All comparisons not listed in Table 5 did not yield significant differences between the fern species. 3.2. Phratora vitellinae As with Agelastica alni, those larvae who had taken up extract from dried or fresh Athyrium filix-femina pro- duced conspicuously less defensive secretion when dis- turbed than those larvae fed on untreated normal food plant or on normal food plant masked with extract of the other two fern species. This applied especially to the first larval instar. In addition, there occurred severe distortions of moult- ing, elytron formation and melanisation when the larvae were fed on willow leaves masked with extract from Dryopteris filix-mas (Fig. 1). Fig. 1. Adult Phratora vitellinae showing massive wing de- formation after having fed on willow leaves masked with ex- tract of Dryopteris-filix-mas. 4. DISCUSSION 4.1. Agelastica alni Young and old larvae fed on alder leaves even when they were covered with fern extract. This result is at odds with the generally held assumption that the forag- ing behaviour of herbivore insects is guided by olfactory and gustatory stimuli of the plants (e.g., DETHIER 1970; FRAENKEL 1959; HARBOURNE 1982; HSIAO 1969; SCHOONHOVEN 1972, 1973). JERMY (1966) stated that oligophagy is characterised rather by the avoidance of repellent substances than by search for attractants. This is rather unlikely since in our experiments the larvae avoided untreated fern leaves and accepted masked al- der leaves as well as masked fern leaves which they should have despised according to JERMY’S statement. Obviously, feeding was triggered by the presence of, feeding stimulants from alder leaves. Mortality after feeding on alder leaves masked with ex- tract from dry fern leaves demonstrates that at least toxic substances — and most probably also possible re- pellents and attractants — were in the aqueous extract at effective concentrations. Of course, the mortality of young larvae fed on fresh or soaked fern leaves (see Table 4) could be caused by starvation. This definitively does not apply to the old larvae feeding on fern leaves masked with extract from leaves of the normal food plant (test nos. 4, 5, 6), who fed on leaves of all three fern species. The dramatic mortality (all test larvae were dead after five days) must be due to incompatible or toxic compounds. The significantly higher mortality of old larvae in the control situation (test no. 24) is most probably due to the higher damage of the leaves they fed on as a conse- quence of their chewing. The partly damaged leaves dried faster than those the young larvae fed on. Masking alder leaves with extract from fresh fern leaves (tests no. 11, 12, 13) did not lead to significant differ- ences of mortality rates in young and old larvae. Thus, the extract cannot contain effective repellents and the toxic components of the extract do not affect young and old larvae to different extents. However, since in tests no. 12 & 13 with young larvae mortality was signifi- cantly higher than in the control, some substance in the aqueous extract of Dryopteris austriaca and Athyrium filix-femina must have had an effect on the young lar- vac. The most interesting results are those of tests 14, 15, and 16 (fresh fern leaves masked with extract from normal food plant): young larvae did not feed on these leaves at all, old larvae ingested leaf material, on day 3 mortality was significantly higher on Dryopteris filix-mas, while after seven days no larva had survived. This is clear evi- dence that (1) some stimulus from the normal food plant caused the larvae to feed on fern leaves, and (2) that the ingestion of fern material causes death either by starva- tion or through toxic components. This explanation is supported by the results of tests no. 21, 22, and 23, where untreated fern leaves — with one exception — have not been ingested at all. In test no. 21, young larvae took up material of D.f.-leaves, however at decreasing amounts per day until on day 4 no larva had survived. This strange behaviour was most probably caused by two factors, in contrast to the other two fern species: the glandular 268 trichomes are not exposed on the surface in D.f. and the larvae could, therefore, not that easily detect repellent or toxic substances from these glands, and young larvae possess probably less chemoreceptors than old ones. As SWAIN (1977) reports, the larvae of phytophagous insect are equipped with a lower number of chemoreceptors than adults. It seems plausible that the number of chemo- receptors increases during larval development. Since eggs are normally deposited on the larval food plant, and since first instar larvae are much less mobile than older ones, we can expect that young larvae are less choosy with re- spect to food plants than older ones. 4.2. Phratora vitellinae Only a few larvae were available, so the results of the feeding experiments do not allow for statistical analy- ses. The obvious result that adult beetles from those lar- vae who ingested components of Dryopterix filix-mas showed elytral deformations points in the same direc- tion as the results of 4. alni: D. filix-mas either contains more or more toxic substances than the other two ferns tested, or these substances reach higher quantities in the aqueous extract. The small sample size does not permit sound generalisa- tions from these results. However, there is hardly any other explanation for the distorted moulting than that the aqueous extract of D. filix-mas contained effective amounts of phytoecdysone. This is surprising since the study of SELVARAJ et al. (2005) revealed that steroids were only in the chloroform and ethanol extracts and in the hexane fraction, but not in the aqueous fraction of Pteridium aquilinum. 4.3. Conclusions The larvae in our experiments fed on ferns only if these leaves were covered with an aqueous extract of the original food plant, A/nus glutinosa or Salix spp., re- spectively (with the one mentioned exception of young Agelastica alni larvae ingesting — and dying — from un- treated leaves of Dryopteris filix-mas. Whenever larvae took up fern leave material, they suffered conspicu- ously, by dying, by less production of defensive secre- tion, or by wing deformation during pupal development. This means that (1) phytoecdysones contained in the fern leaves are not effective as repellents but as toxins. They can, therefore, indeed act as defence against phy- tophagous insects, however not unless these insects have ingested leave material. Thus, both points of view apply — that phytoecdysteroids do not deter insects from feeding on a fern plant (JONES & FIRN 1978) and that phytoecdysones form an effective protection against at- tacks of phytophagous insects (e.g., RUSSELL 1977). (2) Feeding activity on a certain plant is most probably triggered by the presence of feeding stimulants rather Bonner zoologische Beitráge 54 (2005) than prevented by the presence of repellents. (3) The differences found between the effectiveness of Dryop- teris filix-mas and the other two fern species provides a further caveat against uncritical generalisations of results from studies with bracken (Preridium aquilinum), which dominate the literature on fern-insect interaction. Acknowledgements. We thank Gabriele Uhl, Bonn, for her help with the statistics and for critically commenting on the manuscript, and Bradley J. Sinclair, Bonn, for linguistic and substantial improvement of the text. REFERENCES BALICK, M. J.; FURTH, D. G. & COOPER-DRIVER, G. 1978. Biochemical and evolutionary aspects of arthropod predation on ferns. Oecologia 35: 55-89. DETHIER, V. G. 1970. Chemical interactions between plants and insects. Pp. 83-102 in: SONDHEIMER, E. & SIMEONE, J. B. (eds.) Chemical Ecology. Academic Press, New York. D'ROZARIO, A. & BERA, S. 2003. Studies of glandular structures in Preridium aquilinum (L.) Kuhn with spe- cial reference to aphid infestation from Lachen Valley, north Sikkim. Phytomorphology 53: 37-42. FRAENKEL, G. S. 1959. The raison d’etre of secondary plant substances. Science 129: 1466-1470. GHATE, H.V.; Borowiec, L.; RANE, N. S.; RANADE, S. P. & PANDIT, S. 2003. Tortoise beetles and their host plants from Pune (Maharashtra State, India) and nearby places (Coleoptera: Chrysomelidae: Cassidinae). Ge- nus 14: 519-539 GILMAN, A. V. & COOPER-DRIVER, G. 1998. Japanese bee- tle attacks bracken in Vermont. American Fern Journal 88: 47-48. HARBOURNE, J. B. 1982. Hormonal interactions between plants and animals. Pp. 83-102 in: HARBOURNE, J. B. (ed.) Introduction to Ecological Biochemistry. Aca- demic Press, London. HENDRIX, S. D. 1980. An evolutionary and ecological per- spective of the insect fauna of ferns. American Natu- ralist 115: 171-196. HILL, M. P. 1998. Herbivorous insect fauna associated with Azolla species (Pteridophyta: Azollaceae) in southern Africa. African Entomology 6: 370-372. Hsıao, T. H. 1969. Chemical basis of host selection and plant resistance in oligophagous insects. Entomologia experimentalis et applicata 12: 777-788. JERMY, T. 1966. Feeding inhibitors and food preference in chewing phytophagous insects. Entomologia experi- mentalis et applicata 9: 1-12. JONES, C. G. & FIRN, R. D. 1978. The role of phytoecdys- teroids in bracken fern, Pteridium aquilinum (L.) Kuhn as a defense against phytophagous insect attack. Jour- nal of Chemical Ecology 4: 117-138. MARTINS, R. P., LEWINSOHN, T. M. & LAWTON, J. H. 1995. First survey of insects feeding on Pteridium aquilinum in Brazil. Revista Brasileira de Entomologia 39: 151-156. MESIBOV, R. 2001. Bracken and bugs. The Understorey Network Newsletter No. 22: 7. Michael SCHMITT & Sigrun Bopp: Leaf Beetles (Chrysomelidae) Feeding Fern 269 MUKHOPADHYAY, A. & THAPA, D. 1994. Species richness of ferns and associated insects from Darjeeling plains. Journal of the Bombay Natural History Society 91: 86- 90. RUSSELL, G. B. 1977. Plant chemicals affecting insect devel- opment. The New Zealand Entomologist 6: 229-234. SCHOONHOVEN, L. M. 1972. Secondary plant substances and insects. Recent Advances in Phytochemistry 5: 197-224. SCHOONHOVEN, L. M. 1973. Plant recognition by lepidop- terous larvae. Symposia of the Royal entomological Society London 6: 87-99. SELVARAJ, P., DE BRITTO, A. J. & SAHARAYAJ, K. 2005. Phytoecdysone of Pteridium aquilinum (L.) Kuhn (Dennstaedtiaceae) and its pesticidal property on two major pests. Archives of Phytopathology and Plant Protection 38: 99-105. SPECK, T. 1992. Pflanzen. Pp. 44-95 in: SCHMITT, M. (ed.) Lexikon der Biologie vol. 10 (Biologie im Uberblick). Herder Verlag, Freiburg im Breisgau etc. SWAIN, T. 1977. Secondary compounds as protective agents. Annual Review of Plant Physiology and Plant Molecular Biology 28: 479-501. SWEZEY, O.H. 1922. Insects attacking ferns in the Hawai- ian Islands. Proceedings of the Hawaiian Entomologi- cal Society 5: 57-65. WEINTRAUB, J. D., LAWTON, J. H. & SCOBLE, M. J. 1995, Lithinine moths on ferns: A phylogenetic study of in- sect-plant interactions. Biological Journal of the Lin- nean Society 55: 239-250. WINTERBOURN, M. J. 1987. The arthropod fauna of bracken (Preridium aquilinum) on the Port Hills, South Island, New Zealand. New Zealand Entomologist 10: 99-104. Authors’ addresses: Michael SCHMITT (corresponding author), Zoologisches Forschungsmuseum Alexander Koenig, Adenauerallee 160, D-53113 Bonn, Germany, E-mail: m.schmitt@uni-bonn.de; Sigrun Bopp, Univer- sitát, Abteilung Systematische Botanik und Okologie, Albert Einstein-Allee 11, 89081 Ulm, Germany, E-mail sigrun.bopp@uni-ulm.de. x Bonner zoologische Beitráge Band 54 (2005) | Heft 4 | Seiten 271-286 Bonn, Oktober 2006 The Genus Achaenops Suffrian, 1857 (Chrysomelidae: Cryptocephalinae), Designation of Neotypes and Description of New Species’ Matthias SCHOLLER Berlin, Germany Abstract. Described in 1857 by Eduard SUFFRIAN, the identity of the monotypic genus Achaenops remained doubtful. The type specimens were never re-examined after the original description and later lost when the collection of the Natu- ral History Museum in Hamburg was destroyed. In this study, a neotype is designated for the generic type species 4. dorsalis. Two other species described later in Achaenops were not found to be congeneric with 4. dorsalis, and were transferred: Lophistomus mandibularis (Jacoby, 1901: 240) comb. nov. and Acolastus nigrolineatus (Bryant, 1944: 336) comb. nov. Four species originally described in Cryptocephalus are transferred to Achaenops: Achaenops ruficornis (Suffrian, 1857: 75) comb. nov., Achaenops obscurellus (Suffrian, 1857: 76) comb. nov., Achaenops sericinus (Suffrian, 1857: 80) comb. nov., and Achaenops gilvipes (Suffrian, 1857) comb. nov. Protinocephalus weiseanus Reineck, 1913: 648 was found to be a junior synonym of A. gilvipes, and consequently Achaenops Suffrian, 1857 (= Protinocephalus Reineck, 1913 syn. nov.). A neotype is designated for C. obscurellus. Lectotypes are designated for Cryptocephalus ruficornis and C. gilvipes. Two species are described as new to science from South Africa, Achaenops punctatellus sp. nov. from the Cape Peninsula and Achaenops monstrosus sp. nov. from the Western Cape. Male and female genitalia are figured for the first time. A map showing the collection sites is given. Key words. Coleoptera, Achaenopina, South Africa, taxonomy 1. INTRODUCTION The genus Achaenops was described by monotypy by SUFFRIAN (1857), who gave a detailed diagnosis for it. CLAVAREAU (1913) erected the tribe Achaenopini for this genus and no more genera were added to this tribe that is currently ranked as a subtribe Achaenopina of Cryptocephalini. There is no published record of a re- examination of the generic type species A. dorsalis after its original description. Consequently, the study of A. dorsalis is of interest for the understanding of the sys- tematics of the Cryptocephalini. More specifically, it is the only subtribe of Cryptocephalini endemic to the Afrotropical Region. The designation of a neotype for A. dorsalis is necessary. Two species that were later described in Achaenops (JACOBY 1901; BRYANT 1944) are examined as well as types of Afrotropical species of Cryptocephalus and undetermined specimens in order to evaluate the diversity and distribution of Achaenops. 2. MATERIAL AND METHODS The dried adults were dissected by separating the abdo- men in water, the contents were soaked in cold diluted KOH and then washed in water. The eye length was measured in lateral view, the interocular space in frontal view. Paper presented to the 6th International Symposium on the Chry- somelidae, Bonn, Germany, May 7, 2004. Included in this study are specimens located in the fol- lowing nine collections. The letter codens used in the text are according to the list “Abbreviations for Insect and Spider Collections of the World” (EVENHUIS & SAMUELSON 2005). BMNH Natural History Museum, London, Great Britain (J. Beard and S. Shute) MESC Matthias Schóller personal Collection, Berlin, Germany MLUH Martin-Luther-Universität Halle/Saale, Germany, Wissenschaftsbereich Zoologie (K. Schneider) NHMB Naturhistorisches Museum Basel, Switzer- land (E. Sprecher and D. Burckhardt) NMW Naturhistorisches Museum Wien, Austria (H. Schönmann und H. Schillhammer) SAMC Iziko Museums of Capetown (South Afri- can Museum) Cape Town, South Africa (M. Cochrane) SANC South African National Collection of In- sects, Pretoria, South Africa (E. Grobbe- laar) TMSA South Africa, Gauteng, Pretoria, Trans- vaal Museum (R. Müller) ZMHB Museum für Naturkunde der Humboldt- Universität, Berlin, Germany (J. Frisch and M. Uhlig) 272 Bonner zoologische Beitráge 54 (2005) | 10 11 Figs. 1-11. Achaenops dorsalis. 1 habitus; 2 head, frontal; 3 thorax ventral; 4 claw, fore tibia; 5 aedeagus, dors eral: 7 aedeagus, ventral; 8 tergalapodem; 9 kotpresse, dorsal; 10 kotpresse, lateral; 11 kotpresse, ventral. al; 6 aedeagus, lat- Matthias SCHOLLER: The Genus Achaenops 273 3. RESULTS Achaenops dorsalis Suffrian, 1857 (Figs. 1-13) Achaenops dorsalis Suffrian, 1857 (SUFFRIAN 1857: 236) Type specimen. Neotype (male, NHMB): / Graaff- Reinet-Karroo Nat. Res., 830-1300 m, 24.X.1988 [white] / S. Afr., Cape Prov., W. Wittmer [white] / Neo- typus Achaenops dorsalis SUFFRIAN, des. Matthias Schöller [red, printed] /. A neotype is designated here in order to ensure the name’s proper and consistent application. Other specimens studied: 1f (MESC): South Africa, East Cape, Tsitsikamma nat. park, Storm river mouth, 34°01’S, 23°52’E, 5-11-1999, R. Constantin; Im (BMNH): Mossel Bay, Cape Province, October 1921, R. E. Turner, Brit. Mus., 1921-450; 2m (BMNH): Mossel Bay, 1.-14.x1.1921, R. E. Turner, Brit. Mus., 1921-476; 1m (BMNH): Mossel Bay, 18.-30.xi.1921, R.E. Turner, Brit. Mus., 1922-2; Im If (MESC) 2f (NHMB): Cape Province, Graaff-Reinet-Karroo Nature Reserve, 830-1300 m, 24.X.1988, leg. W. Wittmer; 1m (NHMB): Cape Province, Heidelberg, 12-24 km W, 05.X1.1988, leg. W. Wittmer & M.J.D. Brendell. Type locality. The type locality of the neotype is ap- proximately 32°41°S, 24°29’E, the collection locality of the lost types of Suffrian was Kaffernland, which means the territory of the Xhosa people in South Africa. Diagnosis. A small orange-brown species with puncta- tion of pronotum dense and coarse. SUFFRIAN (1857) listed the short, ovate body and the distant eyes with shallow canthus as characters to dif- ferentiate A. dorsalis from species of Cryptocephalus. Description of neotype (male) Habitus. Body small, shape oval (Fig. 1), surface gla- brous, size [mm]: length 1.8, width of elytra at humeri 1.0, length of pronotum 0.5, width 1.0. Head. Head orange, regularly vaulted, glabrous, shiny, punctation sparse and coarse, head sunk into prothorax but still visible in dorsal view; clypeus punctate; eyes evenly convex, both laterally and longitudinally, eyes small, canthus shallow, triangular, eyes distant (Fig. 2), ratio of distance between upper lobes to eye length is 2.7 : 1.8., eye length about 2.5 x gena, gena with a rim for the first antennomere; antenna (Fig. 1) orange, an- tennomeres without circular sensillate depressions, an- tennomere | swollen, antennomeres 5-11 apically di- lated, brown, longer than wide, clypeal area not distinctly separated, labial palpi acute. Thorax. Pronotum: orange, punctation dense and coarse, interstices glabrous, shiny, all margins even, 1.e., basal margin of pronotum without a row of teeth, lateral margins with a fine, narrow carina, only basal parts in posterior angles simultaneously visible in dorsal view: median lobe of basal margin truncate at apex but not raised; prosternal cavities closed; prosternal process transverse; ratio minimum width : length of prosternal process like 2:1.4, sides concave, apically and laterally with a fine ridge, apex concave (Fig. 3); scutellum tri- angular, elongate, glabrous, impunctate, without basal ridge, apically raised above elytra; elytra striate, scutel- lar row + 9 rows + marginal row, interstices glabrous; elytra apically rounded, reaching upper third of py- gidium; epipleura long and broad, 4/5 the length of the elytra, with few punctures only, basal margin very fine; legs orange except for black claws; external edge of front tibiae not distinctly grooved; tarsi stout, first tar- somere of mid- and hind tarsi shorter than the following together; claws slightly dentate (Fig. 4). Abdomen. Pygidium fuscous, side margins orange, sparsely and coarsely punctured, with short white setae, pygidium not visible in dorsal view; sternite 3 as long as sternites 4-7 along midline; lateral lobe at base of ab- domen truncate; sternites orange with margins fuscous, densely and coarsely punctured, dull, with short white setae. Length of aedeagus 0.65 mm, apex with a fine tip (Fig. 5), bent in lateral view (Fig. 6) ventral side exca- vated with a central longitudinal ridge and four nodes (Fig. 7). Tergalapodem small, Y-shaped with apical branches very short (Fig. 8). Description of female: Habitus. Size [mm]: length 2.3, width of elytra at hu- meri 1.3, length of pronotum 0.6, width 1.1. Head. Ratio of distance between upper lobes to eye length is 3.0 : 1.9. Thorax. Pronotum: centre of basal margin black; Elytra: additional punctures between scutellar row and suture. Abdomen. Pygidium orange, base fuscous, sparsely and coarsely punctured, pit of female abdomen very shal- low, almost as long as wide, diameter shorter than length of sternite, base of pit weakly limited and cov- ered with long setae; kotpresse (Figs. 9-11): dorsal sclerites large, length about 2 X width (Fig. 11), not at- tached to the broad sclerotization of the lateral fold (Fig. 10); posterior of the ventral band two weakly sclerotized areas are present which are not sharply delimited, one close to the middle of posterior margin of ventral band and the other, rectangular one not attached to the ventral band (Fig. 9). 274 tra $4 Fig. 12. Chromatic variability of Achaenops dorsalis. Variability. The chromatic variability is shown in Fig. 12 a-e. Distribution and biology. Known from four localities at the eastern part of the Western Cape and the western part of the Eastern Cape (Fig. 13). Figure 14 shows the habitat in Tsitsikama National Park (2xm). Note. The neotype of A. dorsalis is in accordance with the original description by SUFFRIAN (1857). The de- scription of A. dorsalis was based on a series of speci- mens located in the collection of the Natural History Museum Hamburg, Germany. The types in Hamburg were not found and are presumed to have been de- stroyed during World War II together with almost the complete collection of the museum in 1945. Moreover, the collections in Halle/Saale, Berlin (Germany) and Stockholm (Sweden), which harbour type specimens of SUFFRIAN were consulted, but unfortunately none of them had specimens of A. dorsalis. comb. nov. Achaenops ruficornis (Suffrian, 1857) (Figs. 13, 15-16) Cryptocephalus ruficornis Suffrian, 1857 (SUFFRIAN 1857: 75) Type specimen. Lectotype Cryptocephalus ruficornis (female, ZMHB): / 23639 [white] / Pt. Natal Boh. [blue] Bonner zoologische Beitráge 54 (2005) / ruficornis (Boh*) Suffr.* [blue] / Type [red] / Crypto- cephalus ruficornis Suffrian, 1857 Lectotypus designiert Matthias Schóller [red]/ Achaenops ruficornis (Suffrian, 1857), det. M. Schóller 2005 [white]/. A lectotype was designated here in order to ensure the name’s proper and consistent application. Type locality. South Africa, Port Natal (Durban, 29°49’S 30°56’E). Size. Length female lectotype 2.60, width of elytra at humeri 1.45, length of pronotum 0.90 and width 1.4. Head. Eyes relatively large, ratio of minimum distance between upper lobes to eye length is 3.0 : 2.4. Thorax. Claws relatively large, simple. Female. Kotpresse with two small dorsal sclerites sepa- rated from the broad sclerotizations of the lateral fold (Fig. 15) and a ventral band (Fig. 16) terminating in two small apodemes. Distribution and biology. Known only from the type locality. Note. SUFFRIAN (1857: 75-76) gave a detailed descrip- tion of this species. He mentioned male and female specimens communicated by Bohemann in ZMHB and Stockholm. I was only able to trace one female in Matthias SCHOLLER: The Genus Achaenops 275 26 28 30 32 34 1738 117 11738 [1720 Fr 7a Targa 143 Faz 1827 1830 Eee 1831 1892 1833 18 Ki [1929 17930 {7937 115 1933 | 19, + (eat PE | 2023 2029 | 20 il E E 5 = ER —— 2081 [20827 2083 120 20. 2123 214871 2129 12130 (2137 | TEE 2191 [292 2133 21 135 le NER 2223 , | zi a Nf ae a i 230 | 2: 2282 | 2283 | 22 y 5 2323 | 23 me | | 4 pS ¡ea 2391 = 23 312 2483 134i ee o] HA : EEE ee ae a ere poets | a 1 A Y 57 258 [PARO [2521 we ll bf | 2506 [Baer [88 E 2500 | Ber och | | r 2419 12420 12421 T de = 5 | 4 269 |2620) [2621 |2gap [2673 ER [87 |? 8:8 [2620 [2650] 2607 fa 2719 12730 arm [2722 12723 T | | al j9 12820 BEZ 12823 — + A a a | daa x 40 2880 [2881 [2882 | r | | | 9 [2920 [2981 | ere 2923 ; 29 ; 2980 Tzggı T Eve i 4 3021 [8022 [3023 [3024 3026 [3927 + 7 BEE — | 7 se Bre a2 jonas 722 [Sit [3126 [3127 [3108 [aib9 [130 =< dd TF Tula A lest 7 a = EN Du WP SPRL {3202 [9223 [8 3226 [3227 [3228/8209 le + t—+— } 4 ES ||| | 18322 [3323 13324 33) feo. | a x a Ai I 308 Fig. 13. Collection localities of Achaenops Suffrian; MW = A. dorsalis, O = A. ruficornis (Natal), © = A. obscurellus, O = A. sericinus, U = A. punctatellus, X= A. gilvipes, # = A. monstrosus. ZMHB. Achaenops ruficornis is the first species SUF- FRIAN (1857: 75) listed in his second species-group (2. Rotte) of Cryptocephalus, which comprised five species from South Africa. Other specimens studied: Im (MESC): South Africa, W. Cape pr. Cape Town, 34°13S 18°28E, Simon’s Town, Miller Pt, 31.10.1999, R. Constantin; Im (ZMHB), 1f (SANC), If (MESC): S. Africa, Western Cape, Cape Peninsula, Hout Bay, slope 500m S., Achaenops obscurellus (Suffrian, 1857) comb. nov. 34°03'S, 18°21'E, 20 m, 18.10.2003, R. Constantin. Er) Type locality. The type locality of the neotype is Cryptocephalus obscurellus Suffrian, 1857 (SUFFRIAN 1857: 76) Type specimen. Neotype (male, SAMC): / South Af- rica, W. Cape pr. Cape Town, 34°13S 18°28E, Simon’s Town, Miller Pt, 31.10.1999, R. Constantin [white la- bel]/ Neotypus Cryptocephalus obscurellus des. Mat- thias Schóller [red, printed] / Achaenops obscurellus (SUFFRIAN, 1857), det. Matthias Schóller 2005 [white, printed] /. A neotype is designated here in order to ensure the name’s proper and consistent application. Simon’s Town, Miller Point (34°13’S 18°28’E), the col- lection locality of the lost types of SUFFRIAN was Kaf- fernland, which means the territory of the Xhosa people in South Africa. Diagnosis. A small black species with shining, impunc- tate pronotum and pronotal and elytral side margins si- multaneously visible in dorsal view, elytra with regular rows of punctures, and brown legs, differs from 4. dor- salis and A. punctatellus in the structure of the prono- tum, the aedeagus and the colouration. 276 Bonner zoologische Beitráge 54 (2005) Fig. 14. Habitat of Achaenops dorsalis, Tsitsikamma National Park, South Africa. The “Mouth trail” along the seaside dominated by Erica fourcadei, Erica spp., many Asteraceae shrubs (Chrysanthemoides monilifera) on the border of a relict indigenous rain forest of Podocarpus falcatus and P. latifolius. Figs. 15-16. Achaenops ruficornis. 15 kotpresse, dorsal; 16 kotpresse, ventral. Description of neotype (male) Habitus. Body small, shape oval; size [mm]: length 1.90, width of elytra at humeri 1.00, length of pronotum 0.50, width 1.00. Head. Head visible in dorsal view, shiny, with coarse punctures; clypeus simple; labrum brown, mandibles brown, symmetric; eyes large and upper lobes ap- proached (Fig. 17), ratio of minimum distance between upper lobes to eye length is 1.9 : 2.0; eye length about 4 X gena; canthus shallow and sinuate; antenna light brown, segments 6-11 black and apically expanded, an- tennae inserted low on frons; labial palpi acute. Thorax. Pronotum shining and impunctate, with distinct margins, lateral margins with a fine, narrow carina and a row of punctures, simultaneously visible in dorsal view, median lobe of basal margin convex (Fig. 18); proster- nal cavities closed, prosternal process transverse; scutel- lum elongate triangular (Fig. 19), without basal ridge, not apically raised above elytra, base without emargina- tion; elytra with nine regular rows of punctures plus scutellar row which is not reaching the middle of the elytra plus marginal row, rows vanishing towards the base, interstices glabrous, lateral margins entirely simul- taneously visible in dorsal view, elytra apically rounded, epipleuron half the length of the elytra, impunctate; legs brown, no tibial spurs, claws simple, small, apical mar- gin of tarsomere 2 wider than tarsomere 1, fore tibia straight. Matthias SCHOLLER: The Genus Achaenops 277 25 Figs. 17-26. Achaenops obscurellus. 17 head, frontal; 18 basal margin of pronotum; 19 scutellum; aedeagus, lateral; 22 tergalapodem; 23 kotpresse, dorsal; 24 kotpresse, ventral; 25 spermatheca; 26 egg. 20 aedeagus, dorsal; 21 Bonner zoologische Beitráge 54 (2005) Fig. 27. Habitat of Achaenops obscurellus, Cape Peninsula, East coast South of Simon's Town at Miller's Point. A costal shrub dominated by Erica spp and Felicia spp. among some indigenous Protea and alien Pinus tree, South Africa. Abdomen. Venter dark brown, sternites and pygidium with coarse punctures and short white setae, lateral lobe at base of abdomen rounded; elytra covering 50% of pygidium; aedeagus simple (Fig. 20), with ventral side regularly vaulted (Fig. 21), length of aedeagus 0.6 mm, tergalapodem Y-shaped (Fig. 22). Description of female: Habitus. Size [mm]: length 2.50, width of elytra at hu- meri 1.35, length of pronotum 0.65, width 1.25. Head. As in male, but eyes more distant, ratio of minimum distance between upper lobes to eye length is 2.5 : 1.9. Abdomen. Kotpresse: dorsal sclerites triangular, not at- tached to the sclerotization of the lateral fold, dorsal fold slightly sclerotized (Fig. 23), posterior of the ven- tral band a trapeziform, sclerotized area, and another sclerotized area close to the middle of anterior margin of ventral band (Fig. 24); spermatheca (Fig. 25), sper- mathecal ductus coiled up and base extended; an egg found during dissection was longitudinal oval (Fig. 26). Variability. No significant variability was detected. Distribution and biology. Known from two localities on the Cape Peninsula (Fig. 13). Figure 27 shows the habitat on Miller Point. Note. The neotype of A. obscurellus is in accordance with the original description by SUFFRIAN (1857). The description of A. obscurellus was based on a single fe- male specimen located in the collection of the Natural History Museum Hamburg, Germany, but it is presumed destroyed (see above under 4. dorsalis). Moreover, the collections in Halle/Saale, Berlin (Germany) and Stock- holm (Sweden), which harbour type specimens of SUF- FRIAN were consulted, but none of them had a specimen of A. obscurellus. Achaenops obscurellus is the second species SUFFRIAN (1857: 75) listed in his second species-group (2. Rotte) of Cryptocephalus, which comprised five species from South Africa. Matthias SCHOLLER: The Genus Achaenops 279 I NA 3 yr Wh iy OSA Figs. 28-35. Achaenops gilvipes. 28 head, frontal; 29 aedeagus, dorsal; 30 aedeagus, lateral; 31 spermatheca; 32 kotpresse, dor- sal; 33 kotpresse, lateral; 34 Kotpresse ventral; 35 lateral lobe at base of abdomen. 280 Achaenops gilvipes (Suffrian, 1857) comb. nov. (Figs. 13, 28-35) Cryptocephalus gilvipes Suffrian, 1857 (SUFFRIAN 1857: 77) Protinocephalus weiseanus Reineck, 1913 (REINECK 1913: 648) syn. nov. Type specimens. Lectotype Cryptocephalus gilvipes (male, MLUH): /29252 (red ink) [white]/ MLU Halle, WB Zoologie, S.-Nr. 7/1/5 [white]/ Cryptocephalus gil- vipes Suffrian, 1857 Lectotypus designiert Matthias Schöller [red]/ Achaenops gilvipes (Suffrian, 1857), det. M. Schóller 2005 [white]/. A lectotype was designated here in order to ensure the name’s proper and consistent application. In his notebook preserved in Halle, Germany, SUFFRIAN wrote for the specimen no. 29252: “von Thorey erhalten (received from Thorey)”, Thorey was also mentioned in the original description. 3 Paralectotypes (females, MLUH): /22791 (red ink) [white/ MLU Halle, WB Zoologie, S.-Nr. 7/1/5 [white]/; 23711 (red ink) [white]/ MLU Halle, WB Zo- ologie, S.-Nr. 7/1/5 [white]/; 29251 (red ink) [white]/ MLU Halle, WB Zoologie, S.-Nr. 7/1/5 [white]/; all three with my labels: Cryptocephalus gilvipes Suffrian, 1857 Paralectotypus designiert Matthias Schóller [red]/ Achaenops gilvipes (Suffrian, 1857), det. M. Schóller 2005 [white]/. Holotype Protinocephalus weiseanus (male, MLUH): /Caffrar, Thorey (ink) [blue]/ gilvipes Suffr. Typ (ink) [blue]/ ex coll. J. Weise [white]/ Type [red]/ Protino- cephalus Weiseanus Rk. (ink) [white]/ (my label) Holo- typus Protinocephalus weiseanus REINECK vid. Schóller XI 1996 [red]/ Achaenops gilvipes (Suffrian, 1857), det. M. Schóller 2005 [white]/. Other specimens studied: | female, NHMW: Cape, 1867. Note: this record was published in Reineck, 1915 (as Cryptocephalus gilvipes), 2 females BMNH: E. Coll. Chevt., 6756, 273 and 6756, 275; 1 male BMNH, Cape Province, George, 15.-17.X1.1921. Characters not mentioned in previous descriptions. Head. Male with enlarged mandibles (Fig. 28) and gena with triangular projections. Abdomen. Apex of aedeagus (Fig. 29) slightly bend upwards, with a small apical tip, ventral side regularly vaulted (Fig. 30). Length of aedeagus 0.6 mm. Female. Spermatheca (Fig. 31) hook-shaped, slender, spermathecal ductus long and thin, densely coiled up; kotpresse: dorsal sclerites large, two slender sclerotized areas posterior to the dorsal sclerites, dorsal fold slightly Bonner zoologische Beitráge 54 (2005) sclerotized (Fig. 32), lateral fold strongly sclerotized (Fig. 33), posterior of the ventral band a rectangular sclerotized area, and another sclerotized area close to the middle of anterior margin of ventral band (Fig. 34); lateral lobe at base of abdomen rounded (Fig. 35). Variability. No significant variability was detected. Distribution. All specimens except for one male from the Western Cape (Fig. 13) were collected in the 19" century without exact data. Note. Achaenops gilvipes is the third species SUFFRIAN (1857: 75) listed in his second species-group (2. Rotte) of Cryptocephalus, which comprised five species from South Africa. Achaenops sericinus (Suffrian, 1857) comb. nov. (Figs. 36-40) Cryptocephalus sericinus Suffrian, 1857 (SUFFRIAN 1857: 80) Type specimen. Holotype (female, MLUH): 21756 [white label, red ink]/ HOLOTYPUS Cryptocephalus sericinus Suffrian, 1857; vid. Matthias Schóller [red, printed] / Achaenops sericinus (Suffrian, 1857) det. Matthias Schóller 2003. Other specimens studied: 1f (SAMC) / Kimberley Bro. Power 1912 [white] / South African Museum Cape Town (SAMC) [white] /; If (SAMC) / CT 10.86 [white] / South African Museum Cape Town (SAMC) [white] /. Size [mm]: length 3.1-3.4, width of elytra at humeri 1.6- 1.7, length of pronotum 0.95-1.05, width 1.6 Head. Head shagreened and punctation dense, head sunk into prothorax; eyes evenly convex, both laterally and longitudinally, eyes relatively small, distant, ratio eye width : length 1.4 : 2.4, ratio of distance between upper lobes to eye length is 1.9 : 1.2, canthus shallow; antennal segments 1-5 yellowish orange, 6-11 brown and apically expanded, antenna (Fig. 36) 0.36 x body length. Thorax. Pronotum: shagreened and punctation dense; lateral margins even, simultaneously visible in dorsal view; median lobe of basal margin truncate, slightly raised; prosternal process narrow, ratio minimum width: length like 1:2.5, sides concave, apically with a fine ridge, apex concave; scutellum triangular, glabrous, sha- greened and impunctate, without a basal ridge, not api- cally raised above elytra; elytra semistriate, center of rows 1-3 confused, and other rows might be disturbed by additional punctures, too; interstices shagreened, micro- structures round, elytra apically rounded, reaching upper third of pygidium; epipleura long, % the length of the elytra, with few small punctures only; external edge of tibiae distinctly grooved (Fig. 37); claws appendiculate. Matthias SCHOLLER: The Genus Achaenops 281 39 40 Figs. 36-40. Achaenops sericinus. 36 right antenna; 37 right fore leg; 38 spermatheca; 39 kotpresse, dorsal; 40 kotpresse, ventral. Abdomen. Pygidium densely punctured, shagreened, dull, female abdomen with deep longitudinal pit of equal diameter to length of sternite, base of pit straigth, sternite 3 as long as sternites 4-7 along midline; lateral lobe at base of abdomen rounded; sternites dark brown, chagreened, with sparse punctation and short white se- tae. Spermatheca and kotpresse: spermathecal ductus long, simple (1.e. not spiral), coiled up voluminous ball- like close to the spermatheca, insertion to vaginal sac slightly inflated (Fig. 38). kotpresse: dorsal sclerites large, dorsal fold slightly sclerotized (Fig. 39), lateral fold strongly sclerotized, posterior to the dorsal sclerites a tooth-like extension, anterior and posterior of the ven- tral band a triangular sclerotized area (Fig. 40). 282 Bonner zoologische Beitráge 54 (2005) Variability. No significant variability was detected. Distribution and biology. Indicated from two locali- ties, Kimberley (28°43’S 24°44’E) in the eastern North- ern Cape and Cape Town (33°55’S 18°26’E), but both records are not unambiguous collection localities and need confirmation. Note. Achaenops sericinus is the fifth species SUFFRIAN (1857: 75) listed in his second species-group (2. Rotte) of Cryptocephalus, which comprised five species from South Africa. Achaenops punctatellus sp. nov. (Figs. 41-45) Type specimens. Holotype (female, MLUH): 19196 [white label, red ink]/ Holotypus Achaenops punctatel- lus MIHI, des. Matthias Schöller [red, printed] /. Note: a new piece of cardboard was used to glue the specimen, and the original cardboard was kept on the original pin, too. 3 Paratypes: Im (BMNH) / Table Mt. Cape of G. Hope, W. Bevins, 1906-167 [white, blue line]/; Im (SAMC) / Cape Town J. Pureell 1886 (ink) / South African Mu- seum Cape Town (SAMC) [white] /; 1f (BMNH) / Camps Bay, Cape Peninsula, 1-20.x.1920 [white] / S. Africa E. Turner 1920-423. [white, blue line]/; all para- types with my label Paratypus Achaenops punctatellus MIHI, des. Matthias Schöller [red, printed] /. Diagnosis A brown species with tibia and femur yellowish orange and pronotum shagreened and punctured, differs from A. dorsalis in the structure of the pronotum and the colouration, and from A. sericinus in the colour of the legs and the envaginated clypeus. Description of Holotype (female) Size [mm]: length 2.40, width of elytra at humeri 1.30, length of pronotum 0.60 and width 1.20. Head. Head visible in dorsal view, shagreened, with coarse punctures, labrum and mandibles yellow; labial palpi acute; eyes small and upper lobes distant, there- fore ratio of distance between upper lobes to eye length is 2.25 : 1.00, eyes evenly convex, relatively small, can- thus deep; segments 1-3 of antennae yellowish orange, 4-11 brown, 5-11 apically expanded, antennae inserted low on frons. Thorax. Pronotum with distinct margins, basal margin of pronotum without a row of teeth, lateral margins even, not simultaneously visible in dorsal view; median lobe of basal margin truncate, slightly raised, expanded but not impressed prior to the expansion; punctures only on disk of pronotum, very shallow and moderately dense, interstices glabrous; prosternal cavities closed; scutellum triangular, not apically raised above elytra; elytra with punctation striate, 9 regular rows plus scutel- lar row which is not reaching the middle of the elytra, interstices glabrous; epipleuron half the length of the elytra; tibia and femur yellowish orange, external edge of tibiae simple, tarsi brown, claws simple, small; no tibial spurs. Abdomen. Venter brown, sternites and pygidium with coarse punctures and short white setae, pygidium sha- greened; kotpresse (Fig. 41): dorsal sclerite not attached to the sclerotization of the lateral fold, an extension di- rected towards the dorsal fold, posterior of the dorsal sclerite, a linear twig of the sclerotization of the lateral fold directed towards the dorsal fold; dorsal fold slightly sclerotized, posterior of the ventral band a diffuse, weakly sclerotized additional band is present, and an- other sclerotized area close to the middle of anterior margin of ventral band; spermatheca relatively small, spermathecal ductus coiled up densely close to the spermatheca (Fig. 42). Description of male. Head: clypeus V-shaped envagi- nated, mandible asymmetric (Fig. 43); size [mm]: length 2.05, width of elytra at humeri 1.10, length of pronotum 0.6, width 1.05; length of aedeagus 0.55 mm, tip moder- ately acute, orificium large (Fig. 44), with four internal sclerites visible, ventral side regularly vaulted (Fig. 45). Variability. In the female paratype, segments 1-4 of the antennae and the first two tarsal segments are yellow. Etymology. The name punctatellus was proposed by SUFFRIAN in his notebook preserved in Halle, Germany. It is a Latin adjective. Distribution and biology. Known from the surround- ings of Cape Town only. No information on the biology is available. Achaenops monstrosus sp. nov. (Figs. 46-53) Type specimens. Holotype (male, SANC): / SOUTH AFRICA, C.P. Middelberg Pass nr Citrusdal, 32.38S 19.09E. 19.1x.1986 R. Oberprieler [white] / collected on Protea nitida [white] / NATIONAL COLL. OF IN- SECTS Pretoria, S. Afr. [white] / Holotypus Achaenops monstrosus MIHI, des. Matthias Schöller [red] /. 5 Paratypes: If (SANC) / SOUTH AFRICA, C.P. Mid- delberg Pass nr Citrusdal, 32.388 19.09E. 19.1x.1986 R. Oberprieler [white] / collected on Protea nitida [white] /, / Im, 1f (SANC) / SOUTH AFRICA, C.P. Gydo Pass near Prince Alfred, 33.148 19.20E. 19.1x.1986 R. Ober- prieler [white] / collected on Protea nitida [white] /; 1f (NHMB) / Súd Afrika Cape Prov. W. Wittmer [white] / Algeria Cederberg 520/1200m 10.X.1990 [white] /; /1m (TMSA)/S.Afr. ¿SW Cape Prov., Wiedow farm Matthias SCHOLLER: The Genus Achaenops 283 ¿e 44 45 Figs. 41-45. Achaenops punctatellus. 41 kotpresse (lateral, ventral, dorsal); 42 spermatheca; 43 head, frontal; 44 aedeagus, dor- sal; 45 aedeagus, lateral. 31.46 S — 18.46 E [white] / 12.9.1987; E-Y: 2492 fyn- bos vegetation leg. Endródy-Younga [white] /; all para- types with my label Paratypus Achaenops monstrosus MIHI, des. Matthias Schóller [red, printed] /. Diagnosis A large black species with tip of mandibles and base of antennae brown, male with extremely enlarged clypeus and mandibles and pronotum not shagreened. Description of Holotype (male) Habitus. Body large, shape cylindrical, size [mm]: length 3.50, width of elytra at humeri 1.80, length of pronotum 1.20, width 1.90. Head. Head visible in dorsal view, shiny, punctation sparse and coarse, clypeus with two acute thorns and a median triangular extension (Fig. 46), labrum brown, mandibles large, black with reddish-brown apex, apex with three teeth, asymmetric with left mandible larger; eyes comparatively small and upper lobes distant, there- fore ratio of minimum distance between upper lobes to eye length is 2.0 :1.0, eye length about 2 x gena (Fig. 47); canthus normal and sinuate; antenna black, first two segments brown, segments 4-11 apically expanded, antennae inserted at lower third of frons; labial palpi acute. Thorax. Pronotum shiny and impunctate, but surface of pronotum irregularly undulated (seen at high magnifica- tion), with distinct margins, lateral margins with a fine, narrow carina and a row of punctures, simultaneously visible in dorsal view, median lobe of basal margin con- vex; prosternal process transverse; scutellum elongate triangular, without basal ridge, not apically raised above elytra, base without emargination; elytra with nine regu- lar rows of punctures plus scutellar row which is not reaching the middle of the elytra plus marginal row, in- terstices glabrous, with very fine wrinkles, lateral mar- gins not simultaneously visible in dorsal view, basal margin without carina, elytra raised around scutellum, elytra apically rounded, epipleuron 2/3 the length of the elytra, apically with a row of punctures; legs black, no 284 Bonner zoologische Beitráge 54 (2005) = 32 Figs. 46-53. Achaenops monstrosus; 46 head frontal; 47 head, lateral; 48 aedeagus, dorsal; 49 aedeagus, lateral; 50 tergalapo- deme; 51 kotpresse, dorsal; 52 kotpresse, ventral; 53 spermatheca. tibial spurs, fore tibiae bend, about 1/3 longer than mid and hind tibiae, apically extended, claws relatively small, dentate, apical margin of tarsomere 2 wider than tarsomere |. Abdomen. Venter black, sternites and pygidium with coarse punctures and short white setae; elytra covering 50% of pygidium; aedeagus with dorsal side weakly sclerotized along the median line (Fig. 48), with a rounded tip, almost straight in lateral view (Fig. 49), with ventral side regularly vaulted, length of aedeagus 0.9 mm; tergalapodeme simple, y-shaped (Fig. 50). Description of female Clypeus prominent, with shallow projections ventral to the antennae, but without thorn-like projections; eyes small and upper lobes distant, therefore ratio of mini- mum distance between upper lobes to eye length is 2.4 : 1.3; fore tibiae only slightly longer than hind tibiae, and Matthias SCHOLLER: The Genus Achaenops 285 only slightly bent; abdominal pit shallow, almost as long as wide, sides straight; kotpresse: dorsal sclerites slender, with an extension directed towards the dorsal fold (Fig. 51), posterior of the dorsal sclerite, a rectan- gular bent twig of the sclerotization of the lateral fold directed towards the dorsal fold, dorsal fold slightly sclerotized; posterior of the ventral band (Fig. 52), two sclerotized patches are present, and another two scle- rotized areas attached to the anterior and posterior mar- gins of the ventral band, respectively; spermatheca (Fig. 53), spermathecal ductus coiled up. Variability. Size [mm] (mean (max., min., n)): length of male 3.27 (3.50, 3.00, 3.00), female 3.53 (3.60, 3.50, 3.00), width of elytra at humeri in male 1.73 (1.80, 1.70), female 1.77 (1.80, 1.70), length of pronotum in male 1.07 (1.20, 0.90), width 1.80 (1.90, 1.80) length in female 1.00 (1.00, 1.00) and width 1.92 (1.95, 1.90). The puncturation on the clypeus is more dense and coarse in the male from Wiedow farm. Etymology. This species is named after the monstrous appearance of the male’s head. It is a Latin adjective. Distribution and biology. Known from the northern part of the Western Cape (Fig. 13); collected on wagon tree or waboon, Protea nitida Philip Miller, a shrub or tree up to 9 m which is growing typically on drier, lower slopes from 0-1200 m NN on gravely acid sandy soil. Species to be transferred to other genera. Two more species described in Achaenops (JACOBY, 1901; BRYANT, 1944) were found neither to be con- generic nor to belong to Achaenops. This illustrates the need for a neotype for A. dorsalis in the sense of article 75.3 of the International Code of Zoological Nomencla- ture. The two species have to be transferred to other genera: Lophistomus mandibularis (Jacoby, 1901) comb. nov. Achaenops mandibularis Jacoby, 1901 (JACOBY 1901: 240) Acolastus nigrolineatus (Bryant, 1944) comb. nov. Achaenops nigrolineatus Bryant, 1944 (BRYANT 1944: 336) 4. DISCUSSION SUFFRIAN (1857) placed Achaenops dorsalis between Cryptocephalus and Acolastus because of its habitus, shape of the head and the movability of the pronotum, the latter was a character which was of special impor- tance to SUFFRIAN for generic classification. Achaenops dorsalis has an oval habitus, whereas the species de- scribed in Cryptocephalus by SUFFRIAN and transferred to Achaenops in this study have a more or less cylindri- cal habitus. All species of SUFFRIAN’s (1857) second species group (=2. Rotte) of Cryptocephalus were trans- ferred to Achaenops, except for C. atratulus, the holo- type of which was examined during this study and found to belong to Cryptocephalus. The short diagnosis, of the second species group is in accordance with the diagnosis of Achaenops by SUFFRIAN, except for the cy- lindrical habitus. The female genitalia, studied here for the first time, are homogeneous. The spermatheca is hook-shaped, and the spermathecal ductus is coiled up densely. The arrangement of the sclerites of the kot- presse 1s homogeneous, too. The lateral fold is strongly sclerotized, and a pair of dorsal sclerites is present which are not attached to the lateral fold. Ventrally, a band-like sclerite is present. The species differ in the shape of the dorsal sclerites as well as in the presence or absence, or shape of sclerotized areas close to the ven- tral band. The male genitalia show few external charac- ters, however, the shape differs well enough to separate species, especially in lateral view. The ventral side of the aedeagus is regularly vaulted in all species except for A. dorsalis. The history of the genus Protinocephalus is curious; REINECK (1913) published the genus based on a single male specimen. In 1915, he published a large paper on Afrotropical Cryptocephalinae including females of A. gilvipes, but he did not recognise that the female speci- mens belonged to the male he described as P. weise- anus. Protinocephalus was published too late to be included in the catalogue of Cryptocephalinae (CLAVA- REAU 1913) and was never re-examined. | found the holotype in a drawer with in-litteris specimens in ZMHB. The only known plant Achaenops was collected from is Protea. Future studies will show if Achaenops is associ- ated with the fynbos-vegetation. In this publication, two new species were described to illustrate the variability of the genus. There are more undescribed species, a total of at least 25 species. A key to the species of Achaenops and a revised diagnosis for the genus will be given later together with the descrip- tion of the remaining species. Acknowledgements. I would like to thank Dr. Robert Constantin, Saint-Lo, France, for the communication of specimens and habitat pictures, and the colleagues listed in Materials and Methods for the allowing the loan of specimens from the respective museum collections. Dr. Horst Kippenberg, Herzogenaurach, Germany and an anonymous reviewer made valuable suggestions to im- prove the manuscript. 286 REFERENCES BRYANT, G. E. 1944. New species of African Chrysomeli- dae (Coleoptera). Annals & Magazine of Natural His- tory Series 11: 335-340. CLAVAREAU, C. H. 1913. Cryptocephalinae. Pp. 85 — 209 in: Junk, W. & Schenkling, S. (eds.) Coleopterorum Catalogus 53. EVENHUIS, N. L. & SAMUELSON, G. A. 2005. Abbreviations for Insect and Spider Collections of the World. The In- sect and Spider Collections of the World website. http://hbs. bishopmuseum.org/codens/codens-r-us. html. JACOBY, M. 1901. A further contribution to our knowledge of African Phytophagous Coleoptera. Transactions of the Royal Entomological Society London 1901: 209- 256. Bonner zoologische Beitráge 54 (2005) REINECK, G. 1913. Eine neue Cryptocephaliden-Gattung aus Siid-Afrika. Deutsche Entomologische Zeitschrift 1913: 647-648. | REINECK, G. 1915. Uber die áthiopischen Vertreter der Gattungen Cryptocephalus und Melixanthus (Anteris- cus) des Kónigl. Zoolog. Museums in Berlin und eini- ger anderen Museen und Sammlungen. (Coleopt. Chrysomelidae.). Mitteilungen aus dem Zoologischen Museum in Berlin 7: 391-469. SUFFRIAN, E. 1857. Zur Kenntniss der Afrikanischen Cryptocephalen. Linnaea Entomologica 11: 57-260. Author’s address: Dr. Matthias SCHOLLER, Grünberger Str. 33, 10245 Berlin, Germany. E-mail: schoeller@ tricho.b.shuttle.de Bonner zoologische Beitráge Band 54 (2005) | Heft 4 | Seiten 287-295 Bonn, Oktober 2006 Failure-time Analyses of the Effectiveness of Larval Shield Defenses in Tortoise Beetles (Chrysomelidae: Cassidinae)' Fredric V. VENCL'” $: Bengt J. ALLEN’ "Department of Ecology and Evolution, State University of New York at Stony Brook, , Stony Brook, NY, USA The Smithsonian Tropical Research Institute, Balboa, Ancón, Republic of Panama Abstract. Plant chemistry and predation are thought to be major factors responsible for the recurrent evolution of die- tary specialization in herbivorous insects. However, their relative importance and the degree to which they interact to drive diet evolution remain unknown. The present study aims to test predictions of the ‘nasty host plant hypothesis’, which posits that an herbivore’s diet becomes more restricted as its reliance upon novel host plant compounds that con- fer protection from predators increases. The tortoise beetle larval shield system affords a unique opportunity to examine how predation and host plant chemistry interact. Shields can be micro-manipulated, including removal, chemical modi- fication and reattachment, without harm to the larvae. We subjected larvae of different diet breadths produced from basal and derived hosts to a predation bioassay and compared the relative effectiveness of their shields under different treatment conditions. Failure-time analyses, the most appropriate statistical approach for right-censored temporal data, revealed that specialist larvae were consistently less susceptible to predation than were generalists feeding on the same plant. Although generalists were as competent as specialists at handling non-polar host chemistry, specialists were better at manipulating more polar host-derived compounds, which are more likely to include novel chemistry. Host shifts may be constrained to only those plants that possess novel, polar compounds. The interaction between plant chemistry and beetle diet evolution may be one of escalation driven by predation, wherein specialists are increasingly more effective than generalists in the assimilation of host plant polar compounds into shield defenses. Key words. Chrysomelidae, larval shield, Azteca, tortoise beetle, Hispinae, predation, chemical defense, failure-time analysis INTRODUCTION Approximately three-quarters of Earth's biodiversity is involved in a tri-trophic interaction among plants, her- bivorous insects, and insect enemies (STRONG et al. 1984; SOUTHWOOD 1996). However, herbivory repre- sents an ecological obstacle that few groups have been able to surmount and only nine of twenty-nine insect orders have succeeded in colonizing the plant resource spectrum. The vast majority of herbivorous insects are members of either the Lepidoptera or the Coleoptera. Chief among the many obstacles to plant-feeding are (1) bottom-up, plant related physiological factors, and, (2) top-down, ecological factors (FUTUYMA & KEESE 1992; DYER & FLOYD 1993). The major bottom-up obstacle is plant secondary chemistry, 1.e., those compounds not involved in photosynthesis or respiration. Over 20 thou- sand such compounds have been discovered so far and most of them have known resistance or defensive func- tions. The major top-down obstacle is predation. What is the relationship between the astonishing array of plant secondary compounds and insect enemies? The fact is that the vast majority of herbivorous insects are dietary specialists that can feed on only one or a few re- lated plants (EHRLICH & RAVEN 1964; FUTUYMA & ' Paper presented to the 6" International Symposium on the Chry- somelidae, Bonn, Germany, May 7, 2004. KEESE 1992; BERNAYS & CHAPMAN 1994). Why an in- sect would evolve a narrower, rather than a broader diet, presents an ecological conundrum: the likelihood of failing to locate a suitable host plant increases in the short life span of an individual insect with a narrow diet range. How plant chemistry and predation interact and their relative importances in the recurrent evolution of dietary specialization remains unknown. Selection by both natural enemies and plant chemistry could in concert, result in a net narrowing of the herbi- vore's host range, providing that range limitation affords better protection. We wished to test the ‘nasty host plant hypothesis’ (hereafter nhph), an hypothesis derived and modified from the parasitoid/insect host literature for application to the broader host plant/insect herbi- vory/predator reality of the Earth (from GUALD et al. 1992). At the micro-evolutionary time scale, nhph pre- dicts that an insect will evolve a narrower diet, if, through the assimilation of the noxious compounds ac- quired from its host, it is rendered less vulnerable to its natural enemies. In macroevolutionary time, nhph also predicts that host shifts will always be to more, rather than less noxious plants. Specifically nhph predicts that, (1) when grown on the same host, the specialist should “handle”, i.e. sequester, the host’s chemistry better than the corresponding generalist does, both quantitatively and qualitatively; (2) specialists should fare better than generalists when subjected to natural predators in bioas- 288 says; (3) derived specialists will fare better than basal specialists do in bioassays, and finally; (4) derived spe- cialists will have novel compounds in their defenses that account for the difference in efficacy. While evidence exists both for (SMILEY et al. 1985; STAMP & BOWERS 1992; DYER & FLOYD 1993; MONTLLOR & BERNAYS 1993; METCALF 1994; DYER 1995; VENCL & MORTON 1998) and against nhph (STAMP 1992; RANK et. al. 1996; DOBLER et al. 1997; KOPF et al. 1998; BECERRA & VENABLE 1999), a test of its ability to explain how narrow diets evolve, or to pre- dict the direction of host shifts, is still lacking. Such a test would require a combinatorial approach that in- cludes chemical, and mechanistic analyses, field ecol- ogy, within a framework of phylogenetically informed experiments (FUTUYMA 2000). The present study pre- sents preliminary results from an ongoing research pro- gram underway in Panama testing the specific predic- tions of nhph. Our study system, the larval tortoise beetle shield, is well suited for the investigation of how plant chemistry Bonner zoologische Beitráge 54 (2005) and predation might interact to favor the evolution of narrow diet breadths. Tortoise beetle larvae are soft- bodied, leaf surface grazers, and as such, are very ap- parent and predictable targets for predators and parasi- toids. Their shields are composite structures formed from the exuviae and accumulated fecula (Fig. 1A). Shields are attached to a mobile infrastructure, the furca, which emanates from the tip of the abdomen (Fig. 1B). Tortoise beetle larvae possess a bizarre telescoping anus that serves to precisely deposit fecal material on the furca-exuvia complex (Fig. 1C). Shields can be aimed and rapidly waved in the path of an attacking enemy (Fig. 1D). In addition to being physical barriers, shields have been shown to contain a plant-derived, chemical component that significantly enhances their effective- ness as an anti-predator defense (GOMEZ 1997; GOMEZ et al. 1999; MULLER & HILKER 1999; VENCL et al. 1999). Shields are suitable for examining the host plant chemistry/predation interaction because they can be eas- ily removed, chemically modified, and then reattached without otherwise harming the larvae (OLMSTEAD & DENNO 1993; VENCL et al. 1999), Fig. 1. Tortoise beetle larval shield system. (A) Dorsal aspect of shield (dark structure) and larval body with anterior oriented to the right. Scoli project laterally around the larval body and are probably sensory receptors. Three cast skins (exuviae) clearly visible within shield matrix; (B) De-shielded larvae with two-tined furca emanating from tip of abdomen and projecting above the larva; (C) Lateral view of larva with telescoping anus extended and applying fecula to shield; and (D) Shield tilted and wav- ing in direction of perturbation (cat whisker) touching anterior end of larva. Fredric V. VENCL & Bengt J. ALLEN: Larval Shield Defenses in Cassidinae 289 Our objective was to test the predictions of nhph by ex- amining the relative susceptibilities of generalist and of specialist tortoise beetle larvae to predation by a ubiqui- tous generalist ant predator, Azteca, in the Panamanian rain forest. We compared the effectiveness of shield de- fenses of three tortoise beetle species with contrasting diet ranges. To control for the potentially confounding effects of host plant chemistry on the behavior of preda- tors, larvae of both diet-range types were reared on the specialist's host plant, members of the morning glory family, Convolvulaceae. We made two specialist-gene- ralist dietary contrasts: (1) Acromis sparsa, a specialist on Merremia umbellata, against the generalist, Chely- morpha alternans, hereafter Cm, also reared on M. um- bellata ; and (2) Stolas plagiata, a specialist on /pomoea phillomega, once more against the generalist, C. alter- nans, hereafter Cp, also reared on J. phillomega. To ad- dress the importance of larval shield chemistry on ant behavior, shields of each species-host plant combination were subjected to micromanipulation and one of four leaching treatments prior to bioassay trials. 2. METHODS Bioassays were conducted in Gamboa, Republic of Panama, between 8:00 AM and 12:00 PM, during July, August, and September of 2003. Azteca ants (Hymenop- tera: Formicidae: Dolichoderinae) are common, fierce, generalist predators in Neotropical lowland rainforests (CARROLL 1983; HOLLDOBLER & WILSON 1990). We used A. lacrymosa as an assay agent. This species builds large carton nests attached to the boles of trees. They are extremely aggressive, strongly recruiting, and for- aged primarily in the area in and around their home tree. One month prior to the onset of the bioassay experi- ments, we set potted individuals of M. umbellata and 1. phillomega at the base of the home tree. Host vines were placed in contact with the tree trunks to enable the ants to use the host plants as foraging areas. We encour- aged routine patrolling of M. umbellata and 1. phillo- mega vines by the ants by regularly baiting host plant leaves with larva-sized tuna fragments. Shield micromanipulation. Larval sibships of each species-host plant combination were equally divided at random among the following treatment groups: (1) wa- ter (H,O); (2) methanol (MeOH); (3) both (HO fol- lowed by MeOH); and (4) unleached (intact) control. So that we could clearly observe the effects of diet range on shield chemistry in the absence of larval behavior, ant bioassay experiments were done using fourth-instar lar- vae freshly killed by freezing for 5 min. We immedi- ately removed each larval shield by placing fine forceps between the tines of the furca and gently lifting the shield away from the body. Shields were then soaked for 25-30 min in a solvent bath agitated every five min (or two consecutive baths of 12-15 min each in the case of larvae assigned to the treatment with both HO and MeOH). After soaking, shields were dried on paper toweling under an incandescent light bulb and slow fan for 45 min. Each shield was re-attached to the larval furca using rapid-setting, fumeless, water-insoluble craft glue (DAP) that had been warmed to 28° C for five min to minimize setting time. Larvae with re-attached shields were allowed to stand and dry for at least 20 min before bioassays were begun. Controls consisted of the shield removal and reattachment manipulations, but no leaching. To ensure high levels of ant activity on the host plants during bioassay trials, 45 min before the experiments began we baited each plant with pieces of tuna of about the same size as a fourth-instar larva. Each trial con- sisted of the presentation of an individual larva to forag- ing ants on either M. umbellata or I. phillomega. Inı- tially, individuals from each treatment group were randomly assigned to different host plants with dice. Following the first round of testing the delegation of larvae to host plants was constrained by the previous round, such that no treatment group was tested on the same plant more than once nor tested consecutively. Only one trial was done on a given host plant before moving to the next tree, and a minimum of five min elapsed between each trial. Using soft forceps, we placed each experimental larva near the center of a host plant leaf along the mid-vein. This formed the bioassay test arena. To avoid contami- nation, forceps were dedicated to a single treatment and were dipped in water and whipped to dryness between trials. A trial was started if there were at least two, but no more than five ants foraging on the leaf. Each trial lasted five minutes, or until the test larva was captured. A capture event was considered to be the movement of the test larva > 1 cm toward the leaf petiole by the ants. Trials were recorded with a Panasonic digital video camera (PV-DV951) mounted on a tripod positioned such that the entire test leaf was included in the field of view. We started video recording at first contact of an ant with the experimental larva and measured the num- ber of seconds elapsed between the start of the trial and a capture event (or the end of the trial period, whichever came first). Between 35 and 45 replicates of each sol- vent treatment were done for each of the four larval spe- cies-host plant combinations. Statistical analyses. We examined the effects of larval diet range and shield chemistry on the time to capture by Azteca ants using failure-time statistics (PROC LIFETEST; SAS v. 9.0) (reviewed by Fox 2001). In contrast to classical methods such as ANOVA that compare either the total number of captures at the end of the experimental time interval or the mean time to capture among treatment groups, failure-time methods 290 —O— A. sparsa —®— C. alternans (Cm) proportion not captured 250 300 0 50 100 150 Bonner zoologische Beiträge 54 (2005) 1.0 “E TS 0.8 0.6 0.4 —O— S. plagiata —@®— C. alternans (Cp) time (s) Fig. 2. Cumulative capture curves for specialist (open circles) and generalist (filled circles) tortoise beetle larvae with intact, unleached shields in Azteca ant bioassay. Larvae of both diet ranges were raised on either (A) Merremia umbellata (Contrast #1) or (B) Ipomoea phillomega (Contrast #2). Data points are the cumulative fraction of the cohort not yet captured (mean +SE) for each 10 s interval in the bioassay. Both contrasts were statistically significant at the 0.05 level based on Bonferroni-corrected pairwise comparisons following a Wilcoxon signed-rank test for heterogeneity among all four groups (P = 0.0040). compare the distributions of capture times over the entire experimental period. Times to the occurrence of a failure event (e.g., capture of a larva by ants) do not typically meet the distributional assumptions required by tradi- tional parametric approaches. Such data often lack equal variances and normal distributions. In addition, many of the trials ended before a capture event was recorded (so- called right-censored data). As a consequence, the ulti- mate fate of the experimental larva beyond the 5 min cen- sus interval was unknown. Analysis of variance and re- lated tests are unable to account for censored data, however, failure-time methods are not so limited. Cumu- lative capture functions were compared using a Wilcoxon signed rank test followed by pairwise multiple compari- sons to determine specific differences between treatment groups (KALBFLEISCH & PRENTICE 1980). Significance levels were corrected using the sequential Bonferroni technique (Dunn-Sidak method; SOKAL & ROHLF 1995). This method is less conservative than the standard Bon- ferroni technique but ensures that an appropriate experi- ment-wise error rate is maintained. Differences among groups in the proportion of larvae cap- tured were assessed with likelihood-ratio chi-squared tests of independence (SOKAL & ROHLF 1995). 3. RESULTS Failure-time analyses revealed pronounced differences between generalist and specialist larvae with both un- treated and solvent-leached shields in susceptibility to predation by Azteca ants. The relative performance through time of generalists and specialists with un- treated shields is shown in Figure 2. In both diet range contrasts, the specialists were consistently less suscepti- ble to predation than were generalist larvae. Specialist larvae were also significantly less likely than generalists to have been captured by the end of a trial (Contrast #1: G = 5.78, P = 0.016, Contrast #2: G’ = 5.92, P = 0.015). Tortoise beetle shield chemistry. The effects of sol- vent leaching of shields on larval susceptibility to preda- tion were striking. In both contrasts, regardless of diet range, the decay in capture curves for shields leached by some or all solvents (H20, MeOH or both) was signifi- cantly steeper than in curves for intact, unleached shields (Figs 3 and 4; Table 1). For both specialists, shield leaching by H,O had stronger effects on larval capture rates than leaching by MeOH. In contrast, there appears to be an interaction in the effects of different solvents on shields of generalist larvae (although our experimental design unfortunately does not allow for a statistical test of this possibility). Leaching by MeOH had a larger (negative) effect than leaching by H20 on capture rates of C. alternans grown on Merremia um- bellata (Contrast #1) compared to untreated controls. However, the opposite pattern exists for C. alternans grown on /pomoea phillomega (Contrast #2) (Figs. 3 and 4; Table 1). Fredric V. VENCL & Bengt J. ALLEN: Larval Shield Defenses in Cassidinae 291 Acromis sparsa Chelymorpha alternans (Cm) 1.0 @&R —O— intact Oo —@— MeOH (0) —4— H20 pu = ©: © © a [a (= ño E O o P<0.0001 O pum a 0 50 100 150 200 250 300 0 50 100 150 200 250 300 time (s) Fig. 3. Cumulative capture functions for specialist (4. sparsa) and generalist (C. alternans, Cm) tortoise beetle larvae raised on Merremia umbellata (Contrast #1) with intact and solvent-leached shields in the Azteca ant bioassay. P-values are from a Wil- coxon signed-rank test for heterogeneity among groups. Statistical results of Bonferroni-corrected pairwise comparisons are pre- sented in Table 1. Error bars were eliminated for clarity. Stolas plagiata Chelymorpha alternans (Cp) —O— intact —@— MeOH —A— H20 0.8 oO fe) — = —4— both o o O 0.6 + O € c 0.4 S 5 oO. P < 0.0001 0.2 O — Y 0.0 0 50 100 150 200 250 300 0 50 100 150 200 250 300 time (s) Fig. 4. Cumulative capture functions for specialist (S. plagiata) and generalist (C. alternans, Cp) tortoise beetle larvae raised on Ipomoea phillomega (Contrast #2) with intact and solvent-leached shields in the Azteca ant bioassay. P-values are from a Wil- coxon signed-rank test for heterogeneity among groups. Statistical results of Bonferroni-corrected pairwise comparisons are pre- sented in Table 1. Error bars were eliminated for clarity. 292 Bonner zoologische Beitráge 54 (2005) Table 1. Pairwise multiple comparisons of cumulative capture curves for specialist (4. sparsa and S. plagiata) and generalist (C. alternans) tortoise beetle larvae with intact and solvent-leached shields in the Azteca ant bioassay (Figs 3 and 4). Larvae of both diet ranges were raised on either Merremia umbellata (Contrast #1) or Ipomoea phillomega (Contrast #2). To keep the experi- ment-wise error rate at the 0.05 level, comparisons were done using a sequential Bonferroni approach (Dunn-Sidák method; So- kal and Rohlf 1995) following Wilcoxon signed-rank tests. Individual comparisons marked with an asterisk (*) were statistically significant. Contrast #1 Contrast #2 Shield type A. sparsa Cm S. plagiata Cp intact v. MeOH 4 ns ns ns intact v. H,O pe ns e = intact v. both y = de MeOH v. H,0 a ns a e MeOH v. both + ns E) $ HO v. both ns ns ns 4. DISCUSSION 4.1. Are specialists better defended than generalists? Our findings support the central tenet of nhph, the ‘nasty host plant hypothesis’, which posits that when both are feeding on the same host plant, specialist tor- toise beetle larvae will be better defended against preda- tion by their shields than are generalist larvae in an ant bioassay. We found that specialists with unleached, in- tact shields clearly outperformed their generalist coun- terparts in both diet contrasts we tested. Furthermore in both contrasts, we observed that all solvent leaching treatments (HO, MeOH, and both) significantly de- graded shield effectiveness, regardless of diet range. This latter finding supports the idea that shields have a critically important chemical component that consists of both polar and non-polar compounds. 4.2. Is the chemical defense derived from the host plant? Although a definitive answer to the question of the provenance of shield compounds must await compari- sons of plant and shield chemistries, evidence from this study supports the idea that shields are fortified with host-derived metabolites. If a shield system were based on autogenous synthesis, shield chemistry would be the same regardless of diet and thus one would not expect to observe differences in shield performance. When we compared the shield performances of the same beetle, C. alternans, reared on different host plants, we observed that effect of a particular leaching treatment depends on which plant the generalist fed upon, supporting the idea that at least part of larval shield chemistry is host- derived. In all the other fecal shield-bearing species, including several tortoise beetles studied so far, shield chemistry is based on precursors obtained from the host plant (GOMEZ 1997; MORTON & VENCL 1998; VENCL & MORTON 1998; MULLER & HILKER 1999). Fecula-laced shields may well represent a type of sequestered de- fense. If their chemical constituents were found to be host-derived, the shields of the tortoise beetles in this study would also strongly resemble sequestering types of defenses. 4.3. Do specialists sequester host chemistry better than generalists? Our findings lend support to the mhph prediction that specialists are more competent than are their generalist counterparts at handling host plant chemistry. Our data show that the H,O leaching treatment, which removed many of the more polar compounds from the larval shields, significantly increased the susceptibility of both specialist larvae to predation. In contrast, the MeOH leaching treatment had a greater impact on the generalist feeding on one host but not the other. Superior chemical sequestration might involve one or more of the follow- ing strategies: greater bio-concentration of a particular compound, differential sequestration of a variety of compounds, or the modification of the compounds. Many specialist herbivores are known to have enhanced mechanisms for the sequestration and/or the transforma- tion of plant metabolites into defensive compounds (BOWERS 1988; PASTEELS et al. 1983, 1988; DENNO et al. 1990). Some specialist herbivores have been found to more efficiently excrete or egest host plant secondary compounds (SELF et al. 1964; BERENBAUM 1983; FER- GUSON et al. 1985; METCALF 1994). We think that the tortoise beetle shield system is a type of sequestration Fredric V. VENCL & Bengt J. ALLEN: Larval Shield Defenses in Cassidinae 293 process that is relatively inexpensive (OLMSTEAD & DENNO 1993), possibly because tortoise beetle special- ists are superior at harvesting and modifying host me- tabolites compared to their generalist counterparts feed- ing on the same hosts. Since they cannot readily transit cell membranes, non- polar compounds are arguably more difficult to mani- pulate because they first must be modified in order to contain them within the larval gut (DUFFY 1980). The process of gut compartmentalization requires that a compound be made more polar through mechanisms like hydroxylation or conjugation (DUFFY 1980; BOWERS 1988). Our data suggest that given identical dietary in- puts, specialists are more competent at transforming less polar into more polar compounds, and thus dispropor- tionately fortifying their fecula with more polar com- pounds, compared to their generalist counterparts. The identity and defensive characteristics of these more po- lar shield constituents must await future structural elu- cidation. Suffice it to say that many classes of polar substances are well known to have deterrent and toxic characteristics. Some of these more polar compound classes include pyrrolizidine alkaloids, phenolics, cardenolides, sapogenines, and flavonoids. 4.4. Adaptation or accident? How herbivorous insects use their host plants for chemical defense, 1.e., by processes of sequestration, may entail quite different physiological mechanisms from those used by tortoise beetles for processing host compounds into shield fecula. Does shield formation re- quire special adaptations or are shields passive conse- quences of host consumption? Based on findings from shield-forming leaf beetles in other chrysomelid sub- families, VENCL & MORTON (1998) have suggested that shields are not 'default' waste tanks, but instead are ad- aptations for defense. They argue that since shields con- tain a highly culled subset of ingested host derived pre- cursors, some of which are nutrients and some of which are modified within the larval gut, fecal shield forma- tion must have entailed the evolution of specialized en- zymes that now serve the triple defensive functions of: (1) selective compound egestion through compartmen- talization, (2) compound bio-concentration, and (3) compound bio-activation (VENCL et al. 1999). The un- usual telescoping anus of tortoise beetles is de facto evidence of a specialized adaptation for the precise deposition of fecula on the shield framework (see Fig. 1B). There is good evidence from another tortoise bee- tle, Plagiometriona clavata, supporting the contention that shields with fecular retention represent specialized adaptations for predator defense (VENCL et al. 1999). For example, palmitic acid is one of the 'discarded' com- pounds that eventually ends up in P. clavata's shield. An erstwhile nutrient, it occurs in the fecula in relatively higher concentrations than it does in the host. Palmitic acid also elicits necrophoresis (undertaker behavior) in ants, whereby anything emitting it gets placed on the ant nest's dump (BLUM 1970). Palmitic acid therefore ap- pears to be more beneficial as part of the larval defense system than as a dietary nutrient. y Our findings for tortoise beetles are the first instances where specialists have been shown to derive an advan- tage (enemy-free space) over their respective generalists when reared on the specialist's host plant. This conclu- sion is in overall agreement with previous studies on Lepidoptera (STAMP & BOWERS 1992; DYER & FLOYD 1993; CORNELIUS & BERNAYS 1995, but see STAMP 1992). It is important to note that the Lepidoptera and herbivorous Coleoptera represent well over half the in- sects attacking plants and most of these herbivores are dietary specialists (STRONG et al. 1984). More work is necessary to determine if the relationship between the effectiveness of plant-derived, anti-predator chemical defenses and specialization is a general one that has in- fluenced the evolution of diet range in beetles and pos- sibly in other phytophagous insects. We are in the process of determining the origins and elucidating the chemical structures of the compounds responsible for the effects observed in this study. At a macroevolutionary level, nhph predicts that derived specialists will have novel, more polar compounds in their defenses that account for increased shield efficacy when compared to generalists or to basal specialists. If so, then the remarkably robust tortoise beetle radiation might have been fostered by the colonization of increas- ingly more chemically complex dicotyledonous plants. A process of defensive escalation may have enhanced the likelihood of beetle speciation. Whether host shifts have always been to plants containing more, rather than less potent chemistry must await further studies of more basal specialists and generalists in the tortoise beetle ra- diation. It is also essential to determine if selection on shield chemistry is diffuse, or whether each predator's selective impact is idiosyncratic, in which case, we would expect specificity in targeting of particular shield chemicals. We are currently undertaking studies to clar- ify these related issues. Acknowledgements. We wish to thank Kamal Kamalrage, Anayansi Valderrama, Jessica Gurevich, Donald Windsor and Gordon Fox for their help, encouragement, and thoughtful suggestions. We want to thank Fumio Aoki for his invaluable assistance. This 1s contribution # 1123 from the Graduate Program in Ecology and Evolution at Stony Brook University. This study was funded by NSF IBN # 108213 to the senior author. 294 REFERENCES BECERRA, J. X. & VENABLE, D. L. 1999. Macroevolution of insect-host associations: The relevance of host bio- geography to host affiliation. Proceedings of the Na- tional Academy of Sciences of the USA 96: 12626- 12631. BERNAYS, E. A. & CHAPMAN, R. F. 1994. Host-plant Se- lection by Phytophagous Insects. Chapman and Hall, New York. BERENBAUM, M. R. 1983. Coumarins and caterpillars: a case for coevolution. Evolution 37: 163-179. BLuM, M. S. 1970. The chemical basis of insect sociality. Pp. 61-93 in: BEROZA, M. (ed.) Chemicals Controlling Insect Behavior. Academic Press, New York. Bowers, M. D. 1988. Chemistry and coevolution: iridoid glycosides, plants, and herbivorous insects. Pp. 133- 165 in: SPENCER, K. C. (ed.) Chemical Mediation of Coevolution. Academic Press, New York. CARROLL, C. R. 1983. Azteca. Pp. 752-753 in JANZEN, D. H. (ed.) Costa Rican Natural History. University of Chicago Press. CORNELIUS, M. L. & BERNAYS, E. A. 1995. The effect of plant chemistry on the acceptability of caterpillar prey to the Argentine ant /ridomyrmex humilis (Hymenop- tera: Formicidae). Journal of Insect Behavior 8: 579- 393. DENNO, R. F., LARSSON, S. 8 OLMSTEAD, K. L. 1990. Role of enemy-free space and plant quality in host-plant se- lection by willow beetles. Ecology 71: 124-137. DOBLER, S, MARDULYN, P., PASTEELS, J. M. & ROWELL- RAHIER, M. 1996. Host-plant switches nd the evolu- tion of chemical defense and life history in the leaf- beetle genus Oreina. Evolution 50: 2373-2386. Dyer, L. A. 1995. Tasty generalists and nasty specialists? A comparative study of anti-predator mechanisms in tropical Lepidoptera. Ecology 76: 1483-1496. Dyer, L. A. & FLOYD, T. 1993. Determinants of predation on phytophagous insects: the importance of diet breadth. Oecologia 96: 575-582. DUFFY, S. S. 1980. Sequestration of plant natural products by insects. Annual Review of Entomology 25: 447- 477. EHRLICH, P. R. & RAVEN, P. H. 1964. Butterflies and plants: a study in coevolution. Evolution 18: 568-608. Fox, G. A. 2001. Failure-time analysis. Studying times to events and rates at which events occur. Pp. 235-266 in: SCHEINER, S. M. & GUREVICH, J. (eds.) Design and Analysis of Ecological Experiments. Chapman and Hall, New York. FERGUSON, J. E, METCALF, R. L., & FISCHER, D. C. 1985. Disposition and fate of cucurbitacin B in five species of diabroticites. Journal of chemical Ecology 11: 1307-1321. FUTUYMA, D. J. 2000. Some current approaches to the evo- lution of plant-herbivore interactions. Plant Species Biology 15: 1-9. FUTUYMA, D. J. & KEESE, M. C. 1992. Evolution and co- evolution of plants and phytophagous arthropods. Pp. 439-475 in: ROSENTHAL, G. A. & BERENBAUM, M. (eds.) Herbivores: Their interactions with plant secon- dary metabolites VII: Evolutionary and Ecological Processes. Academic Press, San Diego, CA. GAULD, I. D., GASTON, K. J. & JANZEN, D. H. 1992. Plant allelochemicals, tritrophic interactions and the ano- Bonner zoologische Beitráge 54 (2005) molous diversity of tropical parasitoids: the 'nasty' host hypothesis. Oikos 65: 353-357. GOMEZ, N. E. Q. 1997. The fecal shield of larvae of tor- toise beetles (Cassidinae:Chysomelidae. a role for chemical defense using plant derived secondary com- pounds. Ph. D. dissertation, Technischen Universitat Carolo-Wilhelmina, Braunschweig. GOMEZ, N. E., WITTE, L. & HARTMANN, T. 1999. Chemi- cal defense in larval tortoise beetles: essential oil com- position of fecal shields of Eurypedus nigrosignata and foliage of its host plant, Cordia curassavica. Jour- nal of Chemical Ecology 25: 1007-1027. HOLLDOBLER, B. & WILSON, E. O. 1990. The Ants. Belknap Press of Harvard University, Cambridge MA. KALBFLEISCH, J. D. & PRENTICE, R. L. 1980. The Statisti- cal Analysis of Failure-time Data. Wiley, New York. KOPF, A., RANK, N. E., ROININEN, H., JULKUNEN-TITTO, R., PASTEELS, J. M. & TAHVANAINEN, J. 1998. The evolution of host-plant use and sequestration in the leaf beetle genus Phratora (Coleoptera: Chrysomeli- dae). Evolution 52: 517-528. METCALF, R. L. 1994. Chemical ecology of diabroticites. Pp.153-169 in: JOLIVET, P. H., Cox, M. L. & PETITPI- ERRE, E. (eds.) Novel Aspects of the Biology of the Chrysomelidae. Kluwer Academic Publishers, Dord- recht etc. MONTLLOR, C. B. & BERNAYS, E. A. 1993. Invertebrate predators and caterpillar foraging. Pp. 170-202 in: STAMP, N. E. & CASEY, T. M. (eds.) Caterpillars: Eco- logical and Evolutionary Constraints on Foraging. Chapman and Hall, London. Morton, T. C. & VENCL, F. V. 1998. Larval beetles form a defense from recycled host plant chemicals dis- charged as fecal wastes. Journal of chemical Ecology 24: 765-785. MULLER, C. & HILKER, M. 1999. Unexpected reactions of a generalist predator towards defensive devices of cassidine larvae (Coleoptera, Chrysomelidae). Oecolo- gia 118: 166-172. OLMSTEAD, K. L. & DENNO, R. F. 1993. Effectiveness of tortoise beetle larval shields against different predator species. Ecology 74: 1394-1405. PASTEELS, J. M., ROWELL-RAHIER, M., BRACKMAN, J. C. & DUPONT, A. 1983. Salicin from host plant as precur- sor of salicylaldehyde in defensive secretion of chry- somelid larvae. Physiological Entomology 8: 307-314. RANK, N., SMILEY, J. & KÓPF, A. 1996. Natural enemies and host plant relationships for chrysomelid leaf bee- tles feeding on Salicaceae. Pp. 147-171 in: JOLIVET, P. H. A. & Cox, M. L. (eds.) Chrysomelidae Biology. SPB Academic Publishing, Amsterdam. SELF, L. S., GUTHERIE, F. E., & HODGSON, E. 1964. Adap- tations of tobacco hornworms to the ingestion of nico- tine. Journal of Insect Physiology 10: 907-914. SMILEY, J. T., HORN, J. M. & RANK, N. E. 1985. Ecological effects of salicin at three trophic levels: new problems from old adaptations. Science 229: 649-651. SOKAL, R. R. & ROHLF, F. J. 1995. Biometry. Freeman and Co., New York. STAMP, N. E. 1992. Relative susceptibility to predation of two species of caterpillar on plantain. Oecologia 92: 124-129. STAMP, N. E. & Bowers, M. D. 1992. Behavior of special- ist and generalist caterpillars on plantain (Plantago lanceolata). Ecological Entomology 17: 273-279. Fredric V. VENCL & Bengt J. ALLEN: Larval Shield Defenses in Cassidinae 295 STRONG, D. R., LAWTON, J. H. & SOUTHWOOD, T. R. E. 1984. Insects on Plants. Harvard University Press, Cambridge, MA. SOUTHWOOD, T. R. E. 1996. Insect-plant relations: over- view from the symposium. Entomologia Experimen- talis et Applicata 80: 320-324. VENCL, F. V. & MORTON, T. C. 1998. The Shield Defense of the Sumac Flea Beetle Blepharida rhois (Chrysomeli- dae: Alticinae). Chemoecology 8: 25-32. VENCL, F. V., MORTON, T. C., MUMMA, R. C. & SCHULTZ, J. ©. 1999. Shield defense of a larval tortoise beetle. Journal of Chemical Ecology 25: 549-566. Authors’ addresses: Fredric V. VENCL (corresponding author) Department of Ecology and Evolution, State University of New York at Stony Brook, Stony Brook, NY, 11794-5245, USA and The Smithsonian Tropical Research Institute, P.O.Box 2072, Balboa, Ancón, Re- public of Panama, e-mail: fvencl@life.bio.sunysb.edu; , Bengt J. ALLEN, Department of Ecology and Evolution, State University of New York at Stony Brook, Stony Brook, NY, 11794-5245, USA. Bonner zoologische Beiträge Band 54 (2005) | Heft 4 | Seiten 297-303 On Phyletic Closeness Between South American and New Caledonian Spilopyrines (Chrysomelidae, Eumolpinae, Tribe Spilopyrini)' Krishna K. VERMA! & Pierre JOLIVET” 5 "Borsi, Durg, India “Paris, France, and Florida State Museum of Entomology, Gainesville, FL, USA. Abstract. Among members of the eumolpine tribe Spilopyrini (Reid 1995), the South American genera Hornius and Stenomela and the New Caledonian Bohumiljania are especially similar, while the Australian and the New Guinean forms seem to stand a little apart. In this communication, by tracing taxonomic history of spilopyrines, and by taking into account their food plants, palaeogeological history of the Southern Hemisphere, anatomical features of spilopyrines, and their developmental stages, a possible phyletic closeness between the South American and the New Caledonian Bonn, Oktober 2006 genera has been inferred. Key words. Taxonomy, ecology, zoogeography 1. INTRODUCTION REID (2000) separated some members of the chry- somelid subfamily Eumolpinae, and put them in a new subfamily, Spilopyrinae. Earlier these genera had been regarded by REID (1995) as a tribe under Eumolpinae, the Tribe Spilopyrini. These primitive Eumolpinae are being referred to in this communication as spilopyrines, following VERMA & JOLIVET (2004). The spilopyrin genera include: Spilopyra Baly, 1860 (Australia and New Guinea), Macrolema Baly, 1861 (Australia and New Guinea), Richmondia Jacoby, 1898 (Australia), Cheiloxena Baly, 1860 (Australia), Stenomela Erichson, 1847 (Chile), Hornius Fairmaire, 1848 (Chile and Ar- gentina), and Bohumiljania Monros, 1958 (New Cale- donia). As pointed out by VERMA & JOLIVET (2004), the Chilean and the New Caledonian spilopyrines are espe- cially similar. This communication aims to emphasize a possible phyletic closeness between the South American and the New Caledonian spilopyrines. In this context the following are specifically notable. 2. TAXONOMIC HISTORY JOLIVET (1954), describing the wing venation in the New Caledonian spilopyrin, referred to it as Stenomela caledonica. MONROS (1958) created a new generic name for this spilopyrin, Bohumiljania. SEENO & WIL- Cox (1982), following MONRÓS (1958), placed Stenomela and the New Caledonian Bohumiljania to- gether under the Tribe Stenomelini, a tribe under Eu- molpinae. The Tribe Hornibiini, with a single genus Hornius, has been placed close to the Tribe Stenomelini. Paper presented to the 6" International Symposium on the Chry- somelidae, Bonn, Germany, May 7, 2004. These bits of taxonomic history clearly support the no- tion of phyletic closeness between the S. American and the New Caledonian forms. 3. FOOD PLANTS Hornius of Chile, as larva and adult, is a specialised feeder on leaves and bark of Nothofagus trees (Fagaceae) (JEREZ 1996). Stenomela of the same region, on the other hand feeds on different species of Myrtaceae (Blepharo- calyx crukshanksii Niedenzu and Luma spp.). Bohumil- Jania caledonica in the distant New Caledonia, also feeds on Myrtaceae (Sizygium spp.). This is a point of similarity between Stenomela and Bohumiljania. But in spilopyri- nes, Cheiloxena in Australia also feeds on Myrtaceae (Eucalyptus spp.) and Spilopyra on Sapindaceae (Cu- paniopsis and Guioa). Host plants of Macrolema and Richmondia are not on record (JOLIVET et al. 2003). 4. PALAEOGEOLOGICAL HISTORY In the context of distribution of spilopyrines, the Gond- wana Hypothesis has been invoked by JEREZ (1996) and REID (2000). HOLLOWAY (1979) and VERMA & JOLIVET (2004) have pointed out that in the late Cretaceous (75 mya) continental shelves of S. America and of the Aus- tralian Plate were still connected with those of the Ant- arctica, while Africa and India, that also resulted from the break, had moved away northward considerably re- moved from Gondwanaland. In a recent paper SANMARTIN & RONQUIST (2004) gave a detailed and interesting account of Gondwanian history, supported by explicit maps of the southern hemisphere in different geological periods. The fol- lowing parts of the history, as mentioned in that paper 298 Bonner zoologische Beitráge 54 (2005) Gondwanaland e - 60 my Africa Madagascar India New Zealand New Caledonia South America Australia „ Floral/faunal exchanges +“ continued into Pleistocene probably through intermediate islands. a New Guinea - 30 my Fig. 1. Cladograms showing sequence of separation of land masses in the Southern Hemisphere (based on data in SANMARTIN & RONQUIST 2004, and in a summary of her studies provided by I. Sanmartin, Uppsala, pers. comm. 2004). and in a summary of their work, provided by Isabel Sanmartin (Uppsala, pers. comm. 2004), are of special interest in the present context. (1) “Consistent with other studies, the animal data are congruent with the geological sequence of Gond- wana breakup...... Trans-Antarctic dispersal (Aus- tralia > southern South America) is also sig- nificantly more frequent than any other dispersal event in animals, which may be explained by the long period of geological contact between Austra- lia and South America via Antarctica.” (SANMAR- TIN & RONQUIST 2004). (1) In Palaeocene (60 mya) New Zealand + New Caledonia separated from Australia. (111) In mid to late Tertiary (mid Tertiary 40 to 30 mya) New Caledonia separated from New Zealand. (iv) During the period 60 to 30 mya Australia and South America remained in contact across Antarc- tica (Fig. 5), and this continued up to Eocene- early Miocene. At this time the Antarctica had a warm temperate climate, and had a rich flora, dominated by Nothofagus, as evidenced by fossils of this plant found in coastal parts of the Antarc- tica. Ready dispersal of Nothofagus to Australia and South America is probable. I. Sanmartin (Uppsala, pers. comm. 2004) points out, “This (= a warm temperate climate in the Antarctica) presumably established a significant dispersal route for the exchange of southern temperate biota between South America and Australia” (v) Opening of the Drake passage between Antarctica and South America at the boundary between Eo- cene and Oligocene (30 to 28 mya led to the onset of the Circum-Antarctic Current, and this resulted in cooling of the Antarctica. JEREZ (1996) has pointed out that Nothofagus arose in the Australian part of Gondwana, and from there it dis- persed to New Zealand and S. America. As mentioned above, Hornius is a specialized feeder on Nothofagus. It appears that primitive spilopyrines dis- persed to Australia, New Caledonia and S. America, along with temperate flora, like Nothofagus, and subse- quently, due to cooling and development of subantarctic conditions in the southern part of S. America, and due to separation of New Caledonia as an island from the Aus- tralian Plate, the primitive spilopyrines in these parts became shielded from competition with more modern forms, and have survived as Hornius and Stenomela in southern S. America, and as Bohumiljania in New Cale- donia. One may ask: New Guinea is also an island, and it shares Spilopyra and Macrolema with Australia, then how island isolation of New Caledonia could help the survival of Bohumiljania as a primitive spilopyrin? One answer to this question: New Zealand + New Caledonia separated from Australia in Palaeocene (60 mya), whereas floral and faunal exchanges between Australia and New Guinea could continue even into the Pleisto- cene (I. Sanmartin, Uppsala, pers. comm., 2004) (Fig. 5). Krishna K. VERMA & Pierre JOLIVET: South American and New Caledonian Spilopyrini (Chrysomelidae) 299 5. ANATOMICAL FEATURES New Caledonian and S. American spilopyrines have an elongated and cerambycid like body shape (Fig. 2). The Australo-Papuan genera in contrast have a more compact and robust body form like typical Eumolpinae. All the three S. American-New Caledonian genera are light green in colour on the host plant. On the other hand the Austra- lian genera show bright and diverse metallic colours. Fig. 2. New Caledonian and South American spilopyrines. (1) Bohumiljania (from JOLIVET et al. 2003); (11) Hornius (from JEREZ 1996); (111) Stenomela (from JEREZ 1996). 300 Bonner zoologische Beitráge 54 (2005) soe? Fig. 3. Aedeagi, lateral view. (1) of Bohumiljania; (ii) of Hor- nius; (111) of Stenomela (from VERMA & JOLIVET 2004). The aedeagal structure in spilopyrines in general is primitive, as discussed by VERMA & JOLIVET (2004). But in Hornius, Stenomela and Bohumuljania it is even more primitive; the primitive features of their aedeagi include: (1) Poor differentiation of the basal hood and the aedeagus proper (Fig. 3). (ii) Only moderate ventral curvature of the aedeagus, spread throughout the length of the organ, and not confined to the basal part of the aedeagus proper. In Stenomela, however the curvature is more marked in the basal part of the aedeagus proper. (1) The ventrally directed basal orifice of the aedeagus is relatively restricted anteroposteriorly. In contrast: (1) In Spilopyra, Macrolema, Cheiloxena and Richmondia the differentiation of the basal hood and the aedeagus proper is somewhat better marked than in the S. American and New Caledonian forms; (ii) In Richmondia, Cheiloxena and to some extent in Spi- lopyra the ventral curvature of the aedeagal tube is more marked than in S. American and New Caledonian spi- lopyrines, and in Richmondia and Cheiloxena this cur- vature is almost as strong as in higher Eumolpinae; and (111) the anteroposterior extent of the basal orifice is large, specially in Macrolema, Cheiloxena and Rich- mondia (For differences between typical eumolpine and spilopyrin aedeagi see VERMA & JOLIVET 2004). Thus the aedeagal structure in Hornius, Stenomela and Bohumiljania is more primitive than in the remaining spilopyrines, and almost of the chrysomeline type (For features of the chrysomeline type of aedeagus see VERMA 1996). Fig. 4. Spermathecal complex of Bohumiljania (from VERMA & JOLIVET 2004). Spermatheca in all spilopyrines shows a characteristic eumolpine feature, namely the proximal part of the spermathecal capsule is differentiated into a swollen or bulbous section, which receives the duct of the spermathecal gland and from which the spermathecal duct starts. In Bohumiljania this proximal part is some- what elongated and presents some irregular coiling (Fig. 4). Such a coiling is seen also in the spermatheca of Krishna K. VERMA & Pierre JOLiVET: South American and New Caledonian Spilopyrini (Chrysomelidae) South America Antarctic Peninsula ew Guinea el 7 = EN Australia . N Antarctica East Antarctica US ) Antarctica 11 301 Fig. 5. Southern hemisphere. (1) Gondwana (-140 my). End of Jurassic; (11) Polar view of the Southern hemisphere (-80 my). Mid-Cretaceous; (111) Polar view of the Southern hemisphere (- 60 my). Early Eocene. The Drake Passage is just forming. New Zealand, New Caledonia, Norfolk ridge and Lord Howe ridge are also figured (from Isabel Sanmartin summary (2002), pers. comm. 2004, with permission). 302 Bonner zoologische Beitráge 54 (2005) 2 Fig. 6. Bohumiljania caledonica (Jolivet). (1) starting to fly; (2) on a Syzygium cumini L. ( Myrtaceae) leaf, in New Cale- donia Megascelinae (SUZUKI 1988). Megascelinae are essen- tially Central and South American in distribution, and are closely related to Eumolpinae (JOLIVET 1957-1959; REID 1995; COX 1998). Should this situation be re- garded as covered by what has been described by MAYR & ASHLOCK (1991) in the following part of their text? “A serious problem is posed by propensity of genotypes to produce a certain phenotype, such as stalked eyes in certain acalyptrate dipterans, which is manifested in only some of the possessors of a genotype. Parallelism in this case may be defined as homologous similarity, since the common ancestor evidently had the genetic propensity even if it was not expressed phenotypically.” In this context it would be relevant to point out that Megascelis does not have a mediocubital patch in the hind wing like spilopyrines. Richmondia, however has a dark spot in this position, and not a well defined me- diocubital patch (VERMA & JOLIVET 2004). 6. DEVELOPMENTAL STAGES Eggs of Hornius are laid under an oothecal cover, made of excrement material, and a single cover includes 7-8 eggs (JEREZ & CERDA 1988). In Stenomela, eggs are also provided with excretory matter cover, each cover protecting two eggs. In Bohumiljania the ootheca is made up of excretory material mixed with vegetal fibres and a glandular secretion, and there are four eggs per ootheca (JOLIVET et al. 2003). Perhaps it will not be out place to mention that in Megascelis there is also a mem- branous protective cover for a batch of about 10 eggs (COX 1988). In remaining spilopyrines laid eggs have not yet been observed and reported. The neonate larva of Bohumiljania bores onto the stem of the host plant, but later instars are exposed feeders on leaf lamina (JOLIVET et al. 2003). There is a similar habit with the first instar larvae of Hornius and Stenomela; they pierce into leaf buds and leaf lamina, but later instars are well exposed, feeding on leaves. Larval habits have not been recorded so far for the Aus- tralian spilopyrines. Pupae of Hornius are formed in soil in a specially con- structed cell (JEREZ 1996). Pupae of Stenomela and Bo- humiljania have not been observed yet, but they too are believed to be formed in soil (JEREZ 1995; JOLIVET et al. 2003). 7. CONCLUDING REMARKS Phyletic closeness between spilopyrines of New Cale- donia and S. America has been hypothesized on the ba- sis of the above discussion. This hypothesis will hope- fully gain additional support through further studies on the biology of the Australian spilopyrines, and through DNA analysis studies. Acknowledgements. We must thank here Dr. Michael Schmitt, from Zoologisches Forschungsmuseum Alexander Koenig, Bonn, who welcomed the junior author at the Chrysomelid symposium held during the African Biodiver- sity Conference, in May 2004, and Dr. Michael Thomas, from the Florida Museum of Entomology, Gainesville, FL who has provided laboratory access to the junior author while in the USA. Our best thanks also to Dr. Isabel San- martin (Uppsala University, Sweden) who provided decu- mentation and illustrations. Krishna K. VERMA & Pierre JOLIVET: South American and New Caledonian Spilopyrini (Chrysomelidae) 303 REFERENCES Cox, M. L. 1998. The first instar larva of Megascelis puella Lacordaire (Coleoptera, Chrysomelidae, Megascelinae) and its value in the placement of the Megascelinae. Journal of Natural History 32: 387-403. HOLLOWAY, J. D. 1979. A survey of the Lepidoptera, bio- geography and ecology of New Caledonia. Dr. W. Junk Publishers, The Hague. 588 pages. JEREZ, V. 1995. Stenomela pallida Erichson, 1847. Rede- scription, ontogeny and affinities with genus Hornius (Chrysomelidae, Eumolpinae). Gayana Zoologia 59: 1- 122 JEREZ, V. 1996. Biology and phylogenetic remarks of the subantarctic genera Hornius, Stenomela and Dictyneis (Chrysomelidae, Eumolpinae). Pp. 239-258 in: JOLI- VET, P. H. A. & Cox, M. L. (eds.) Chrysomelidae Bi- ology, vol. 3, S. P. B. Academic Publishing, Amster- dam. JEREZ, V. & CERDA, L. 1988. Antecedentes morfologicos y biologicos de Hornius grandis (Phil.& Phil. 1864) (Chrysomelidae — Eumolpinae). Bosque 9(2): 83-86. JOLIVET, P. 1954 (1957-1959). Recherches sur l’aile des Chrysomeloidea (Coleoptera). D.Sc.Thesis (Paris Uni- versity) and Memoires de l'Institut Royal des Sciences Naturelles Belgique 51: 1-180 and 58: 1-152. JOLIVET, P., VERMA, K. K & MILLE, C. 2003. Biology and taxonomy of Bohumiljania caledonica (Jolivet) (Col- eoptera, Chrysomelidae). Nouvelle Revue d’Entomo- logie (N. S.) 20(1): 3-22. MAYR, E. & ASHLOCK, P. D. 1991. Principles of System- atic Zoology (Second Edn.). 475 pp., McGraw-Hill Inc., New York. Monros, F. 1958. Consideraciones sobre la fauna del sur de Chile y revision de la Tribu Stenomelini. Acta Zoo- logica Lilloana 15: 143-153. REID, C. A. M. 1995. A cladistic analysis of subfamilial re- lationships in the Chrysomelidae sensu lato (Chry- someloidea). Pp.559-631 in: PAKALUK, J & SLIPINSKI, S. A. (eds.) Biology, Phylogeny, and Classification of th Coleoptera- Papers celebrating the 80” Birthday of Roy A. Crowson. Warszawa Muzeum Instytut Zo- ologii PAN. REID, C. A. M. 2000. Spilopyrinae Chapuis: a new subfam- ily in the Chrysomelidae and its systematic placement (Coleoptera). Invertebrate Taxonomy 14: 837-862. SANMARTIN, I. & RONQUIST, F. 2004. Southern hemisphere biogeography inferred by event-based models: plant versus animal patterns. Systematic Biology 53(2): 216- 243. SEENO, T. N. & WILCOX, J. A. 1982. The Leaf Beetle Gen- era (Coleoptera, Chrysomelidae). Entomography Pub- lications, Sacramento, CA, USA, pp. 1-221. SUZUKI, K. 1988. Comparative morphology of the internal reproductive system of the Chrysomelidae (Col.). Pp. 317-355 in: JOLIVET, P. PETITPIERRE, E. & HSIAO, T. H. (eds.) Biology of Chrysomelidae. Kluwer Aca- demic Publishers, Dordrecht. VERMA, K. K. 1996. Inter-subfamily relations among Chrysomelidae as suggested by organization of the male genital system. Pp. 317-351 in: JOLIVET, P. H. A. & Cox, M. L. (eds.) Chrysomelidae Biology, vol. 3, S. P. B. Academic Publishing, Amsterdam. VERMA, K. K. & JOLIVET, P. 2002. Comments on Spilopy- rinae ( Col. Chrysomelidae). Nouvelle Revue d’Ento- mologie ( N. S.) 19(2):99-110. VERMA, K. K. & JOLIVET, P. 2004. The primitive Eumol- pinae and the Gondwana Hypothesis. Pp. 395-406 in: JOLIVET, P., SANTIAGO-BLAY, J. A. & SCHMITT, M. (eds.) New Developments in the Biology of Chry- somelidae. S. P. B. Academic Publishing by, the Hague. Authors’ addresses: Krishna K. VERMA, HIG1/327, Housing Board Colony, Borsi, Durg — 491001, India. E- mail: kkvermain@sancharnet.in; Pierre JOLIVET (corre- sponding author), 67 Boulevard Soult, 75012 Paris, France, and Florida State Museum of Entomology, DPI, Gainesville, FL, 32614-7100, USA. E-mail: timarcha@club-internet. fr. on sl pl pu p men f E) = D 4 Ñ $ E — nt | gual a) A re Bonner zoologische Beitrage Band 54 (2005) | Heft 4 | Seiten 305-312 Bonn, Oktober 2006 in Hungary (Coleoptera: Chrysomelidae)' Karoly VIG? & Viktor MARKO” ; 4) Department of Natural History, Savaria Museum, Hungary Department of Entomology, Corvinus University of Budapest, Hungary Abstract. The species richness and species composition of Coleoptera assemblages were investigated in deciduous tree canopies in Hungary. Apple and pear orchards were investigated in Nagykovacsi, Kecskemét and Sarospatak in 1990— 94, and limes and maples in Keszthely in 1999-2002. This study presents in detail the findings on leaf beetles. Earlier investigations in Hungary revealed surprisingly high diversity of Coleoptera assemblages in the canopy of apple and pear orchards. Altogether 324 species, almost 3 per cent of Hungary’s beetle fauna, were represented: 253 in apple and 188 in pear orchards. The majority of the species belonged to the Curculionidae, Chrysomelidae and Coccinelidae families. The proportion of leaf beetles ranged between 15 and 20 %. The commonest leaf-beetle species in the canopy of the commercial orchards investigated were Phyllotreta vittula, Phyllotreta atra, Phyllotreta nigripes, Oulema melanopus, and Aphthona euphorbiae. In the abandoned orchards, the commonest were Luperus xanthopoda, Smarag- dina salicina and Orsodacne lineola. Examination of the fauna of parks, avenues and other planted urban plant stocks has only begun to occupy researchers in the last decade. Analysis of the full Arthropoda assemblages of these plant stocks has still not been undertaken. The proportion of leaf-beetle species in the material gathered on maples and limes in Keszthely ranged between 17.0 and 21.3 per cent. Apart form leaf beetles, the bulk of the specimen material collected consisted of species of the Coccinelli- dae, Staphylinidae and Curculionidae families. The commonest leaf-beetle specimens collected in the lime canopy were Aphthona euphorbiae, Chaetocnema tibialis, Longitarsus lycopi, Longitarsus pellucidus, Longitarsus pratensis and Longitarsus succineus. The commonest on maple were Aphthona euphorbiae, Chaetocnema concinna, Chaetocnema tibialis, Longitarsus lycopi, Longitarsus pellucidus, Longitarsus succineus, Phyllotreta cruciferae and Phyllotreta vit- tula. It was concluded that leaf beetles contribute a high proportion of the biodiversity of the deciduous tree canopy, some- times occurring with high species richness and abundance. However, the reasons for this occurrence and their potential Species Composition of Leaf Beetle Assemblages in Deciduous Tree Canopies / role are poorly understood. Key words. Coleoptera, Chrysomelidae, Hungary, faunistics, deciduous tree canopy fauna 1. INTRODUCTION Attention focused on apple and pear orchards during the development of integrated plant-protection methods. Knowledge of the biology of some key pests and preda- tors (BALAZS 1992; BLOMMERS 1994; HERARD 1986) is extensive enough to describe the role of these organisms in the ecological processes of such orchards. Yet the species composition and diversity of the total arthropod assemblages are still poorly known, although the infor- mation could be essential to a general understanding of the biodiversity relations of the plantations. Two long-term investigations of the arthropod fauna of apple orchards in Hungary have been carried out in the last century (ZILAHI-SEBESS 1955; MESZAROS 1984). An additional investigation focused on the Coleoptera assemblages of apple and pear orchards in Hungary (MARKO et al. 1995). ' Paper presented to the 6" International Symposium on the Chry- somelidae, Bonn, Germany, May 7, 2004. Examination of the fauna of parks, avenues and other planted urban plant stocks has only begun to occupy re- searchers in the last decade. This research had several objectives, but plant-protection criteria were dominant. Nonetheless, analysis of the full Arthropoda assem- blages of these plant stocks has still not been under- taken. This study aims to compare the leaf-beetle samples col- lected in the canopy of deciduous trees in Hungary and explain the similarities and differences between the samples. 2. MATERIALS AND METHODS Investigations. of the canopy fauna of apple and pear or- chards were carried out in three geographical regions of Hungary. The samples were collected at the following localities. Nagykovacsi (3 plots) is situated in the centre of Hungary in woodland amidst hills of medium height and surrounded by mixed oak forest. Kecskemét (5 plots), situated in the Great Hungarian Plain, and Saro- 306 spatak (4 plots) in Northern Hungary both lie in a low- land agricultural environment. Some of the selected plots had been untreated, and others treated with broad- spectrum insecticides (mainly organophosphorus insec- ticides and pyrethroids), and also treated with selective insecticides (mainly IGR pesticides) in IPM orchards. The samples restricted to the canopies were collected Bonner zoologische Beitráge 54 (2005) between 1990 and 1994, from April to November, by beating methods, using the Winkler-type umbrella (© 0.7 m) or 0.25 m” plastic sheet. Sampling per year var- ied between 12 and 22 occasions. The number of trees per sampling varied between 10 to 30. Table 1 shows the parameters of each area investigated. Table 1. Description of the orchards investigated in Hungary (after MARKÓ et al. 1995) Woodland in hill of medium height Agricultural lowland Nagykovacsi Kecskemét Sárospatak Fruit species apple pear pear apple apple apple pear Age of plantation M M M Y Y M M Size of plantation 5.8 ha 1.1 ha 51 ha 2 ha 2 ha 5-6 ha 50.6 ha Untreated + Conventionally + + rT: st: + treated IPM applied + + + + No. of treatments - - 3-6 7-8 7-8 7-8 7-8 Collecting method U U U U S U U Years 1990-92 1992-4 1992-4 1992-3 1992-4 1992-4 1993-4 Sampling per year 12 12 12 12 23-24 21-22 4/1993 7/1994 Trees per sampling 10 10 12 10 30 branches 10 10 M = mature tree, more than 13 years old; Y = younger than 13 years old; U = umbrella; S = plastic sheet Investigation of lime and maple canopies was carried out in Keszthely and vicinity in 1999-2002. Keszthely lies in the middle of Western Hungary at the western end of Lake Balaton, in a basically agricultural lowland environment. Collections were made 8-10 times during the vegetation period, using pyrethroid spraying of the whole canopy and canopy netting (Y = 0.5 m). The number of trees per sampling varied between 4 to 10. The composition of the chrysomelid communities was compared by metric ordination (principal coordinate analysis—PCoA), based on the Horn index, after log2 transformation of the data, and the Jaccard similarity in- dex (KREBS 1989), using the Syntax 5.1 program (Po- DANI 1997). The starting data for these analyses were obtained by aggregating all the individuals collected in the canopy of the orchard investigated (apple and pear) or tree species (lime and maple). 3. RESULTS Tables 2, 3, 4 and 5 show the results obtained in the canopies of the apple and pear orchards. There were a total of 253 Coleoptera species in the canopies of apple trees and 188 Coleoptera species in those of pear trees (MARKO et al. 1995). At Nagykovacsi, altogether 14,917 specimens of Coleoptera were collected, of which 756 specimens were leaf beetles; there were 176 Coleoptera species collected, of which 36 were leaf- beetle species (Table 2). The number of leaf-beetle specimens collected in Nagykovacsi in 1990-94 appears in Table 3. The commonest leaf-beetle species in the canopy of conventionally treated pear orchard were Lu- perus xanthopoda (Schrank, 1781), Phyllotreta atra (Fabricius, 1775), Phyllotreta nigripes (Fabricius, 1775) and Phyllotreta vittula (Redtenbacher, 1849). In the abandoned apple orchards, the commonest was L. xan- thopoda, while in the untreated pear orchard, the com- monest were L. xanthopoda, Smaragdina (Monrosia) salicina (Scopoli, 1763) and Orsodacne lineola (Panzer, 1794). The high proportion of L. xanthopoda in all or- chards was probably due to the large number of wild cherry trees in the surrounding mixed oak forest. It is in- teresting that the number of P. vittula was high in the conventionally treated pear orchards in the years inves- tigated. This leaf beetle is not a pear-feeding phyto- phagous species, its diet being restricted to cruciferous plants and grasses (VIG 1998). The reason of its occur- rence in the pear canopy is unknown. Karoly VIG & Viktor MARKO: Leaf Beetles from Tree Canopies in Hungary Table 2. Number of Coleoptera specimens and species collected in apple and pear-orchard canopies in Nagykovacsi, Hungary (1990-94) Total Coleoptera specimens Total Coleoptera species No. of Chrysomelidae specimens No. of Chrysomelidae species 307 A = untreated pear (1990-92); B = untreated apple (1990-92); C = conventionally treated pear (1992-4) Table 3. Number of leaf-beetle specimens collected in apple and pear-orchards in Nagykovácsi, Hungary (1990-94) Species Altica sp. Altica oleracea (Linnaeus, 1758) Aphthona euphorbiae (Schrank, 1781) Cassida (s. str.) flaveola Thun- berg, 1794 Cassida (s. str.) nebulosa Linna- eus, 1758 Chaetocnema (Tlanoma) concinna (Marsham, 1802) Chaetocnema (Tlanoma) tibialis (Illiger, 1807) Clytra laeviuscula (Ratzeburg, 1837) Crepidodera aurata (Marsham, 1802) Cryptocephalus (s. str.) hypochae- ridis (Linnaeus, 1758) Cryptocephalus (Burlinius) chrysopus Gmelin, 1790 Cryptocephalus (s. str.) imperialis Laicharting, 1781 Cryptocephalus (s. str.) nitidus (Linnaeus, 1758) Cryptocephalus (s. str.) nitidus (Linnaeus, 1758) Gastrophysa polygoni (Linnaeus, 1758) Labidostomis longimana (Linna- eus, 1761) Lachnaia sexpunctata (Scopoli, 1763) th 6 A B C Total 7,430 4,506 2,981 - 14,917 125 95 70 176 168 181 407 756 24 17 19 36 Species A B C Luperus xanthopoda (Schrank, ay 133 57 1781) Orsodacne lineola (Panzer, 1794) 31 = | Oulema melanopus (Linnaeus, 2 3 5 1758) Pachybrachis tessellatus (Olivier, | | — 1791) Phyllotreta atra (Fabricius, 1775) = 1 32 Phyllotreta cruciferae (Goeze, | | A 1777) Phyllotreta diademata Foudras, _ _ | 1859 Phyllotreta nemorum (Linnaeus, 2 3 3 1758) Phyllotreta nigripes (Fabricius, 3 3 34 1775) Phyllotreta nodicornis (Marsham, I _ _ 1802) Phyllotreta undulata Kutschera, - - | 1860 Phyllotreta vittula (Redtenbacher, 4 = 245 1849) Psylliodes (s. str.) tricolor Weise, = = l 1888 Smaragdina (Monrosia) affinis (MU- — ] - liger, 1794) Smaragdina (Monrosia) aurita 1 = = (Linnaeus, 1767) Smaragdina (Monrosia) salicina 3] 3 | (Scopoli, 1763) Sphaeroderma testaceum (Fabri- - - | clus, 1775) A = untreated pear (1990-92); B = untreated apple (1990-92); C = conventionally treated pear (1992-4) 308 Bonner zoologische Beitráge 54 (2005) Table 4. Number of Coleoptera specimens and species collected in apple-orchard canopies in Kecskemét, Hungary (1992-94) Total Coleoptera specimens Total Coleoptera species No. of Chrysomelidae specimens No. of Chrysomelidae species A B C D E Total 2075 507 474 449 505 4010 85 54 45 39 46 131 36 24 13 18 16 107 14 8 5 7 5 19 A = mature, IPM applied; B = mature, conventionally treated with board-spectrum insecticides; C = young, conventionally treated; D = young, IPM applied; E = young, IPM applied Table 5. Number of Coleoptera specimens and species collected in pear-orchard canopies in Sárospatak, Hungary (1993-4) Total Coleoptera specimens Total Coleoptera species No. of Chrysomelidae specimens No. of Chrysomelidae species A = conventionally treated; B = untreated and IPM applied A B Total 490 427 919 31 31 45 8 19 27 2 5 6 Table 6. Number of Coleoptera specimens and species and number and proportion of leaf-beetle specimens and species collected in lime and maple canopies in Keszthely, Hungary (1999-2002) Total Coleoptera specimens Total Coleoptera species No. (proportion) of Chrysomelidae specimens No. (proportion) of Chrysomelidae species A B Total 4054 2404 6458 182 211 283 266 (6.6%) 441 (18.3%) 707 (10.94%) 31 (17.0%) 45 (21.3%) 53.18.73) A = lime; B = maple At Kecskemét, altogether 4010 Coleoptera specimens were collected of which 107 specimens were leaf bee- tles and there were 131 Coleoptera species collected of which 19 leaf beetle species were captured (Table 4.). Interestingly, P. vittula dominated in both the conven- tionally and selectively insecticide-treated apple or- chards at the end of June and in mid-July in the years investigated. At Sarospatak, altogether 917 Coleoptera specimens were collected, of which 27 specimens were leaf bee- tles. There were 45 Coleoptera species represented, in- cluding six leaf-beetle species (Table 5). Table 6 summarizes the results obtained during the in- vestigation of the canopies of lime and maple trees in Hungary. There were a total of 182 Coleoptera species in the lime canopies and 211 Coleoptera species in the maple. At Keszthely, altogether 6458 Coleoptera speci- mens were collected, of which 707 specimens were leaf beetles. Of the 283 Coleoptera species collected, 53 were leaf-beetle species. The commonest leaf beetle species in the lime canopy were Aphthona euphorbiae (Schrank, 1781), Chaetocnema (Tlanoma) tibialis (1- liger, 1807), Longitarsus (s. str.) Ivcopi (Foudras, 1860), Longitarsus (s. str.) pellucidus (Foudras, 1860), Longi- tarsus (s. str.) pratensis (Panzer, 1794) and Longitarsus (s. str.) succineus (Foudras, 1860), and in the maple canopy A. euphorbiae, Chaetocnema (Tlanoma) con- cinna (Marsham, 1802), C. tibialis, L. lycopi, L. pellu- cidus, L. succineus, Phyllotreta cruciferae (Goeze, 1777) and P. vittula (Table 7). The results of metric ordinations show clear differences between the treatments and tree families. The composi- tion (species composition and relative abundance) of leaf-beetle assemblages in the canopies of untreated ap- ple and pear trees was very similar and distinct from the Karoly VIG & Viktor MARKO: Leaf Beetles from Tree Canopies in Hungary Table 7. Number of leaf-beetle specimens collected in lime and maple canopies in Keszthely, Hungary (1999-2002) Species Altica oleracea (Linnaeus, 1758) Aphthona abdominalis (Duftschmid, 1825) Aphthona euphorbiae (Schrank, 1781) Aphtona nonstriata (Goeze, 1777) Aphthona semicyanea Allard, 1859 Aphthona venustula (Kutschera, 1861) Cassida (Mionychella) hemisphaerica Herbst, 1799 Cassida (Hypocassida) subferruginea Schrank, 1776 Chaetocnema (s. str.) aridula (Gyllenhal, 1827) Chaetocnema (Tlanoma) concinna (Mar- sham, 1802) Chaetocnema (s. str.) hortensis (Geoffroy, 1785) Chaetocnema (Tlanoma) picipes Ste- phens, 1831 Chaetocnema (Tlanoma) tibialis (Mliger, 1807) Chrysolina (Menthastriella) herbacea (Duftschmid, 1825) Clytra laeviuscula (Ratzeburg, 1837) Crepidodera aurata (Marsham, 1802) Cryptocephalus (Burlinius) planifrons Weise, 1882 Cryptocephalus (s. str.) sericeus (Linna- eus, 1758) Epitrix pubescens (Koch, 1803) Galeruca (s. str.) pomonae (Scopoli, 1763) Longitarsus sp. | Longitarsus sp. 2 Longitarsus (s. str.) kutscherae (Rye, 1872) Longitarsus (s. str.) lateripunctatus personatus Weise, 1893 A = lime (1999-2001; B = maple (1999-2002) tO ir) A tn 77 Ww NO) 309 Species A B Longitarsus (s. str.) lewisii (Baly, 1874) 1 2 Longitarsus (s. str.) luridus (Scopoli, 7 10 1763) / Longitarsus (s. str.) lycopi (Foudras, 19 24 1860) Longitarsus (s. str.) melanocephalus (De - | Geer, 1775) Longitarsus (s. str.) nasturtii (Fabricius, 8 7 1792) Longitarsus (s. str.) pellucidus (Foudras, 29 38 1860) Longitarsus (s. str.) pratensis (Panzer, 10 1] 1794) Longitarsus (s. str.) rubiginosus (Foudras, - 5 1860) Longitarsus (s. str.) substriatus Kutschera, ] 3 1863 Longitarsus (s. str.) succineus (Foudras, 35 34 1860) Neocrepidodera ferruginea (Scopoli, 2 6 1763) Phyllotreta astrachanica Lopatin, 1977 | 6 Phyllotreta cruciferae (Goeze, 1777) 9 17 Phyllotreta diademata Foudras, 1859 — 6 Phyllotreta nemorum (Linnaeus, 1758) — l Phyllotreta nigripes (Fabricius, 1775) 3 10 Phyllotreta procera (Redtenbacher, 1849) 3 7 Phyllotreta striolata (Fabricius, 1803) = l Phyllotreta undulata Kutschera, 1860 | 5 Phyllotreta vittula (Redtenbacher, 1849) 5 22 Psylliodes (s. str.) chrysocephalus (Lin- 2 7 naeus, 1758) Psylliodes (s. str.) illyricus Leonardi & = l Gruev, 1993 Psylliodes (s. str.) napi (Fabricius, 1792) | | Psylliodes (s. str.) picinus (Marsham, - 1 1802) 310 Bonner zoologische Beitráge 54 (2005) Chrysomelidae assemblages, PCoA, Horn similarity Ms. 03 e Tilia 2 0.25] +. °® Pear abandoned e Acer Apple abandoned 6 3 -0.3 e Pear treated 2 * Apple treated e Pear treated 1 0 Axis 1 Fig. 1. Examination of the similarity of leaf-beetle assemblages gathered on various deciduous trees, by metric ordination (Horn index). Chrysomelidae assemblages, PCoA, Horn similarity (log2 n) e Pear treated 1 3 ° Apple treated e Pear treated 2 Axis 2 Apple abandoned 4 e e Pear abandoned e Tilia Acer e -0.4 -0.3 -0.2 -0.1 0 0.1 0.2 0.3 0.4 Axis 1 Fig. 2. Examination of the similarity of leaf-beetle assemblages gathered on various deciduous trees, by metric ordination (Horn index, after log2 (x+1) transformation of the raw data). Karoly Vic & Viktor MARKO: Leaf Beetles from Tree Canopies in Hungary 31] assemblages of maple and lime trees (Fig. 1). However, the abandoned orchards were situated in the same dis- trict (Nagykovácsi) and the maple and lime trees in a different one (Keszthely). It can also be seen that char- acteristic chrysomelid assemblages were formed in commercial orchards, regardless of differences in their surroundings. On the other hand, the commercial pear orchard2 (No. 6 in the figures) is clearly distinct from the abandoned orchards, despite all being situated in the same district (Fig. 1). The results are similar if the analysis based on log2 transformed data are compared, or only the species compositions (Jaccard index). In these analysis, the role of the commonest, dominant species was lower (log2 transformation) or relative abundance was not taken into consideration (Jaccard similarity) in the comparisons (Figs. 2 and 3). So the re- sults show the basic structure of the chrysomelid assem- blages. The neighbouring habitats seem to be more im- portant in forming the species composition of the chrysomelid assemblages. The species composition in the commercial pear orchard2 was closer to the aban- doned ones from the same location than to the other commercial orchards. Similarly, the leaf beetle assem- blages in the canopies of lime and maple trees from the same location, Keszthely show remarkably similar composition (Fig. 3). Chrysomelidae assemblages, PCoA, Jaccard similarity 5 Pear treated 1 e 6 0 e Pear treated 2 4 e Apple abandoned 7 e Pear abandoned ® Apple treated 1 Tilia e Acer e 0.1 0.2 0.3 0.4 0.5 Axis 1 Fig. 3. Examination of the similarity of leaf-beetle assemblages gathered on various deciduous trees, by metric ordination (Jac- card index). 4. DISCUSSION The leaf-beetle assemblages examined consist of spe- cies that sometimes feed on the tree species con- cerned and much larger specimen and species num- bers of visiting species. The untreated apple and pear orchards typically had large numbers of specimens of species feeding on the trees (L. xanthopoda and O. lineola). Treatment obviously increases the propor- tion in the assemblages of visiting species less tied to a food plant (principally and sometimes exclusively Phyllotreta vittula), and this causes the distinction in the commercial plantations. On lime and maple spe- cies, however, visiting species were found almost ex- clusively, so that the different characteristics of the two species of tree do not give them distinct beetle faunas. The tourists have a decisive role in species composition. The relatively large number of tourist species (A. euphorbiae, Cassida (s. str.) nebulosa Linnaeus, 1758, C. concinna, Cryptocephalus (s. str.) nitidus (Linnaeus, 1758), Lachnaia sexpunctata (Sco- poli, 1763), Oulema melanopus (Linnaeus, 1758), P. atra, P. cruciferae, Phyllotreta nemorum (Linnaeus, 1758), P. nigripes, P. vittula and S. (M.) salicina in Nagykovacsi and 283 species in the case of Kesz- thely) have arrived from the environs, so that the spe- cies composition is more similar on trees found in the same environment. The explanation for this, as said 312 Bonner zoologische Beitráge 54 (2005) earlier, is that the environs of the trees have the deci- sive effect on the development of the species compo- sition. Leaf beetles contribute a high proportion of the biodi- versity of the deciduous tree canopy, sometimes occur- ring with high species richness and abundance. How- ever, the reasons for this occurrence and their potential role are poorly understood. Most of the species are cer- tainly tourists, which arrived in the canopy by chance, since these tree species are not their food plant. How- ever, it cannot be ruled out that some species may occa- sionally draw moisture from the leaves of the deciduous tree species examined, especially if the main food plant has become inedible. With some species, consumption of honeydew or sooty moulds is conceivable, although unsupported by observations specific to these species. Acknowledgements. Karoly Vig was supported by the Janos Bolyai Foundation of the Hungarian Academy of Sciences. REFERENCES BALAZS, K. 1992. Proceedings of the international sympo- sium on integrated plant protection in orchards, Gödöllö 1990. Acta Phytopathologica et Entomologica Hungarica 27(1-4): 1-690. BLOOMERS, L. H. M. 1994. Integrated pest management in European apple orchards. Annual Review of Entomol- ogy 39: 213-241. HERALD, F. 1986. Annotated list of the entomophagous complex associated with pear psylla, Psylla pyri (L.) (Homoptera: Psyllidae) in France. Agronomie 6(1): 1- 34. KREBS, C. J. 1989. Ecological Methodology. Harper and Row Publishers, New York, 654 pp. MARKO, V., MERKL, O., PODLUSSANY, A., VIG, K., KU- TASI, Cs. & BOGYA, S. 1995. Species composition of Coleoptera assemblages in the canopies of Hungarian apple and pear orchards. Acta Phytopathologica et En- tomologica Hungarica 30(3—4): 221-245. MESZAROS, Z. (ed.) 1984. Results of faunistical and flo- ristical studies in Hungarian apple orchards. Acta Phytopathologica et Entomologica Hungarica 19: 91— 176. PODANI, J. 1997. SYN-TAX 5.1: A new version for PC and Macintosh computers. Coenoses 12: 149-152. VIG, K. 1998. Host plant selection by Phyllotreta vittula (Redtenbacher, 1849). Pp. 233-251 in: BIONDI, M., DACCORDI, M. & FURTH, D. G. (eds.) Proceedings of the Fourth International Symposium on the Chry- somelidae. Proceedings of a Symposium (30 August, 1996, Florence Italy), XX International Congress of Entomology, Torino, Museo Regionale di Scienze Natural. ZILAHI-SEBESS, G. 1955. Mes recherches faunistiques dans les vergers de Tiszacsege. Acta Universitatis Debrece- niensis de Ludovico Kossuth Nominate 2: 1-15. Authors’ addresses: Karoly VIG (corresponding au- thor), Savaria Museum, Department of Natural History, H-9700 Szombathely, Kisfaludy S, u. 9., Hungary, E- mail: nathist.savmuz@axelerto.hu; Viktor MARKO, Cor- vinus University of Budapest, Department of Entomol- ogy, H-1118 Budapest, Ménesi ut 44., Hungary, E-mail: vmarko@omega.kee.hu Bonner zoologische Beitrage Instructions to Authors Bonner zoologische Beitráge publishes original papers, reviews, and scientific notes covering the whole field of zoology, with em- phasis on biodiversity, biogeography, history of zoology, mor- phology, phylogenetics, and taxonomy. The material must not be submitted for publication elsewhere. All manuscripts will be peer- refereed. Final acceptance of items for publication is the responsi- bility of the corresponding editor dealing with the respective taxon (see list of editors). Manuscripts not compatible with the following guidelines will be returned. 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Are the tables and figures sufficient? = Y u . _ u 7 4 0 | - | J A E $ A o Y = ] j 7 h 3 ‘ y bi 5 4 , y u A A x R i . u ; i Ve { 1 = \ -. ! mM u , [2 ' 7% 1 f { i ae Y y y * \ ñ ñ Y . 7 . . ‘ ya 7 t . ; E y u r : ' + - y , 1 ' 4 4 y e . r - 3 I i : f é o N hr \ ı - b I } vn 7 we i 4 o # 5 ' a Ni ‘Inhalt Special issue: Proceedings of the 6th International Symposium on the Chrysomelidae, held at Zoologisches Forschungsmuseum Alexander Koenig, Bonn, Germany, May 7, 2004. ASLAN, Irfan; BEENEN, Ron & ÖZBET, Hikmet: SMITHSONIAN INSTITUTIO 173 Biological Aspects of Galeruca circassica Reitter, 1889 = Cephalaria procera Fish. and Lall. (Dipsacaceae) in Anatolia 3 9088 0132 5 4 727 BEENEN Ron: 179 Translocation in Leaf Beetles (Coleoptera: Chrysomelidae) DUNGELHOFF, Susanne & SCHMITT, Michael: , 201 Functional Morphology of Copulation in Chrysomelidae-Criocerinae and Bruchidae (Insecta: Coleoptera) FURTH, David G.: 209 The Current Status of Knowledge of the Alticinae of Mexico (Coleoptera: Chrysomelidae) GEISER, Elisabeth: 239 Frozen Chrysomelids from Alpine Glaciers - Witnesses of the Postglacial Resettlement GROSS, Jürgen & FATOUROS, Nina E.: 247 Striking Differences in Behaviour and Ecology Between Populations of Chrysomela lapponica HUBWEBER, Lasse & SCHMITT, Michael: 253 Parameres - Similarities and Differences in Chrysomelidae and Cerambycidae (Coleoptera) SCHMITT, Michael & Bopp, Sigrun: 261 Leaf Beetles (Insecta: Coleoptera: Chrysomelidae) Suffer From Feeding on Fern Leaves SCHOLLER, Matthias: 271 The Genus Achaenops Suffrian, 1857 (Chrysomelidae: Chrysomelidae), Designation of Neotypes and Description of New Species VENCL, Fredric V. & ALLEN, Bengt J.: 287 Failure-time Analyses of the Effectiveness of Larval Shield Defenses in Tortoise Beetles (Chrysomelidae: Cassidinae) VERMA, Krishna K. & JOLIVET, Pierre: 297 | On Phyletic Closeness Between South American and New Caledonian Spilopyrines (Chrysomelidae, Eumolpinae, Tribe Spilopyrini) VIG Karoly & MARKO, Viktor: 305 Species Composition of Leaf Beetle Assemblages in Deciduous Tree Canopies in Hungary (Coleoptera: Chrysomelidae) Titelbild/ Cover illustration: Calligrapha fulvipes Stal, 1859 (Chrysomelidae), Sabanilla, Costa Rica 2005 (M. 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