AIDOTOOZ 10 ASSINS INAJA = a ADOTOOZ 10 IVNANO[ SSIMS REVUE SUISSE DE ZOOLOGIE TOME 114—FASCICULE 1 Publication subventionnée par: ACADEMIE SUISSE DES SCIENCES NATURELLES (SCNAT) VILLE DE GENEVE SOCIETE SUISSE DE ZOOLOGIE DANIELLE DECROUEZ Directrice du Muséum d’histoire naturelle de Genéve ALICE CIBOIS, PETER SCHUCHERT Chargés de recherche au Muséum d’histoire naturelle de Geneve Comité de lecture Il est constitué en outre du président de la Société suisse de Zoologie, du directeur du Muséum de Genève et de représentants des instituts de zoologie des universités suisses. Les manuscrits sont soumis à des experts d’institutions suisses ou étrangères selon le sujet étudié. La préférence sera donnée aux travaux concernant les domaines suivants: taxonomie, systématique, faunistique, phylogénie, évolution, morphologie et anatomie comparée. Administration MUSEUM D'HISTOIRE NATURELLE 1211 GENÈVE 6 Internet: http://www.ville-ge.ch/musinfo/mhng/page/rsz.htm PRIX DE L’ABONNEMENT: SUISSE Fr. 225.— UNION POSTALE Fr. 250.— (en francs suisses) Les demandes d’abonnement doivent étre adressées à la rédaction de la Revue suisse de Zoologie, Museum d’histoire naturelle, C.P. 6434, CH-1211 Genève 6, Suisse ANNALES de la : SOCIETE SUISSE DE ZOOLOGIE et du MUSEUM D'HISTOIRE NATURELLE de la Ville de Genève tome 114 fascicule 1 2007 E x kl GENEVE MARS 2007 ISSN 0035 - 418 X REVUE SUISSE DE ZOOLOGIE SWISS JOURNAL OF ZOOLOGY REVUE SUISSE DE ZOOLOGIE TOME 114—FASCICULE 1 Publication subventionnée par: ACADEMIE SUISSE DES SCIENCES NATURELLES (SCNAT) VILLE DE GENEVE SOCIETE SUISSE DE ZOOLOGIE DANIELLE DECROUEZ Directrice du Muséum d’ histoire naturelle de Genève ALICE CIBOIS, PETER SCHUCHERT Chargés de recherche au Muséum d’histoire naturelle de Genève Comité de lecture Il est constitué en outre du président de la Société suisse de Zoologie, du directeur du Muséum de Genève et de représentants des instituts de zoologie des universités suisses. Les manuscrits sont soumis à des experts d’ institutions suisses ou étrangères selon le sujet étudié. La préférence sera donnée aux travaux concernant les domaines suivants: taxonomie, systématique, faunistique, phylogénie, évolution, morphologie et anatomie comparée. Administration MUSEUM D’HISTOIRE NATURELLE 1211 GENEVE 6 Internet: http://www. ville-ge.ch/musinfo/mhng/page/rsz.htm PRIX DE L’ABONNEMENT: SUISSE Fr. 225.— UNION POSTALE Fr. 250.— (en francs suisses) Les demandes d’abonnement doivent étre adressées a la rédaction de la Revue suisse de Zoologie, Muséum d’histoire naturelle, C.P. 6434, CH-1211 Genéve 6, Suisse REVUE SUISSE DE ZOOLOGIE 114 (1): 3-12; mars 2007 Additions à la faune des scorpions de l’Etat du Amapa, Brésil (Chelicerata, Scorpiones) Wilson R. LOURENCO! & Jian-Xin QI? 1 Departement de Systématique et Evolution, USM 0602, Section Arthropodes (Arachnologie), Muséum national d’ Histoire naturelle, CP 053, 61 rue Buffon, F-75005 Paris, France. E-mail: arachne@mnhn fr 2 Department of Biological Sciences, National University of Singapore, 14 Science Drive 4, Singapore 117543, Singapore. E-mail: free8wind@yahoo.com.cn | Addition to the scorpion fauna of the Amapa State in Brazil (Cheli- cerata, Scorpiones). - This paper presents the results of our study of a small collection of neotropical scorpions now deposited in the Geneva Museum. The collection is composed of specimens of 6 species in 5 genera and 2 families. One new species, Auyantepuia amapaensis sp. n. (Chactidae), is described from the region of the Monte Tipac in Brazil. Keywords: Auyantepuia - new species - Brazil - Amapa State. INTRODUCTION La faune des scorpions de la région néotropicale, et en particulier celle du Brésil, peut étre considérée comme une des plus étudiées au monde. Dés le début du XIX® siècle, les scorpions du Brésil ont été l’objet de nombreuses contributions, abou- tissant a une premiere synthese réalisée par Mello-Leitäo (1945). Dans une perspective plus moderne, plusieurs travaux d’ensemble concernant la systématique et la biogéographie des scorpions brésiliens ont été réalisés au cours des trois dernières décennies, parmi lesquels nous pouvons citer Lourenço (1982a, b, 1986, 1994 et 1996). Une synthèse plus récente, réalisée en 2002 par le même auteur, mont- re qu’au sein de la région néotropicale, le Brésil présente une des faunes scorpioniques les plus diversifiées, ce que confirme de manière fréquente la découverte de nouveaux taxa, et de nouvelles stations pour des espèces déjà connues. Dans l’ensemble du Brésil, certaines régions demeurent peu explorées. C’est le cas de la région septentrionale, souvent désignée sous le nom de ‘Guyane brésilienne’, qui comprend l’Etat de l’ Amapä et ses environs. Le présent travail est le résultat de l’étude d’une petite collection de scorpions collectés dans l’Etat de l’Amapä, et déposée au Muséum d’histoire naturelle de Genève. La plupart des espèces identifiées sont déjà connues de la Guyane française voisine (Lourenço, 1983), et certaines sont citées pour la première fois du Brésil. En outre, une nouvelle espèce appartenant au genre Auyantepuia Gonzales-Sponga est décrite. La présentation du matériel étudié est faite ci-après dans l’ordre alphabétique des familles, genres et espèces. Manuscript accepted 21.11.2006 4 W. R. LOURENCO & J.-X. QI METHODES Les illustrations et les mensurations ont été réalisées à l’aide d’une loupe Wild MS, équipée d’une Camara Lucida et d’un micromètre oculaire. La nomenclature tri- chobothrial adoptée est celle de Vachon (1974). Les mensurations sont réalisées selon Stahnke (1970) et données en mm. La terminologie morphologique suit celle proposée par Vachon (1952) et Hjelle (1990). TAXA CONSIDERES DANS LE TRAVAIL FAMILLE DES BUTHIDAE C.L. Kocu, 1837 Genre Tityus C.L. Koch, 1836 Tityus cambridgei Pocock, 1897 MATERIEL: Brésil, Etat de 1’ Amapa, Serra do Navio (J. Lacroix leg.), X/1973, 6 mâles, 4 femelles. FAMILLE DES CHACTIDAE Pocock, 1893 Genre Broteochactas Pocock, 1893 Broteochactas delicatus (Karsch, 1879) MATÉRIEL: Brésil, Etat de l’Amapä, Serra do Navio (J. Lacroix leg.), X/1973, 1 mâle, 2 femelles. Genre Brotheas C.L. Koch, 1837 Brotheas gervaisii Pocock, 1893 MATERIEL: Brésil, Etat de 1’ Amapä, Serra do Navio (J. Lacroix leg.), IX/1973, 7 mâles, 4 femelles. Brotheas granulatus Simon, 1877 MATERIEL: Brésil, Etat de 1’ Amapa, Serra do Navio (J. Lacroix leg.), X/1973, 18 mâles, 6 femelles. Genre Hadrurochactas Pocock, 1893 Hadrurochactas schaumii (Karsch, 1880) MATÉRIEL: Brésil, Etat de l’Amapä, N. Serra do Navio (J. Lacroix leg.), XI/1973, 20 mâles, 3 femelles. REMARQUE: Espèce nouvelle pour le Brésil. Genre Auyantepuia Gonzalez-Sponga, 1978 HISTORIQUE SUR LA SYSTÉMATIQUE DU GENRE AUYANTEPUIA: Comme cela a été expliqué dans un travail récent (Lourenço & Souza Aratijo, 2004), le genre Auyantepuia a été créé par Gonzalez-Sponga (1978) pour accueillir une seule espèce, Broteochactas scorzai Dagert, 1957, endémique de la région de l’Auyantepui, for- mation montagneuse de la région Guayana, au Venezuela. Le genre est resté mono- typique jusqu’à la publication par Lourenço (1983), qui y ajoute trois espèces nou- velles de la Guyane française. Peu de temps après, Francke & Boos (1986), dans une publication sur les scorpions Chactidae de Trinidad et Tobago, réévaluent les caractères diagnostiques du genre Auyantepuia et le considèrent comme synonyme du genre Broteochactas Pocock. Mais dans leur analyse, ces deux auteurs ne tiennent compte ADDITIONS A LA FAUNE DES SCORPIONS DE L’ETAT DU AMAPA, BRESIL 5 Fics 1-6 Anneau V du metasoma et telson, vue ventrale (2) et latérale (1, 3-6). (1-2) Auyantepuia mottai (holotype femelle). On observe nettement l’arc postérieur formé par des granules spiniformes. (3) A. fravalae (holotype mâle). (4) A. gaillardi (holotype mâle). (5) A. sissomi (holotype femelle). (6) A. amapaensis sp. n. (holotype male). 6 W. R. LOURENCO & J.-X. QI que de la seule espèce type du genre, A. scorzai. Presque en même temps, Lourenco (1986) dans une étude biogéographique sur les scorpions de la région Guyano- Amazonienne, penche plutöt pour une subdivision par groupes d’especes au sein du vaste genre Broteochactas. Il définit ainsi, en leur attribuant des noms de genre, les groupes ‘Auyantepuia’, auquel est rattachée l’espèce Broteochactas parvulus Pocock, 1897 “Taurepania’, “Vachoniochactas’ et “Hadrurochactas’. Quelques années plus tard, deux nouvelles espèces sont décrites et rattachées au groupe ‘Auyantepuia’: Broteochactas kelleri Lourengo, 1997 et Broteochactas skuki Lourengo & Pinto da Rocha, 2000. Sissom (1990), dans sa classification des Chactidae, ignore les groupes d’espèces proposés par Lourenço (1986). Il fait uniquement mention d’un groupe “Vachoniochactas’ a l’intérieur du genre Broteochactas, groupe qui sera réhabilité en tant que genre (Lourengo, 1994). Monod & Lourenço (2001), analysant de manière précise l’ensemble des espèces de Broteochactas associées au groupe d’espèces ‘Hadrurochactas’, mettent en évidence les particularités tant morphologiques que bio- géographiques de ce groupe, mais hésitent à le rétablir en tant que genre valable, ce qui sera fait par Soleglad & Fet (2003). Bien entendu nous sommes d’accord avec cette dé- cision. Dans ce travail de Soleglad & Fet (2003) d’autres décisions moins cohérentes à nos yeux sont également prises, comme la description d’un nouveau genre Neochactas, dans lequel sont englobées, entre autres, plusieurs espèces du groupe ‘Auyantepuia’. Suite à la revalidation du genre Auyantepuia par Lourenço & Souza Aratjo (2004), Soleglad & Fet (2005) récidivent en considérant à nouveau Auyantepuia comme synonyme du genre Broteochactas. Cependant, comme il semble que les travaux de Soleglad & Fet (2003, 2005) présentent des nombreuses inadé- quations, tout récemment signalées par Prendini & Wheeler (2005), nous adoptons ici la position de ces derniers auteurs. Avec la description d’une nouvelle espèce, un modèle de distribution géographique disjoint et relictuel est confirmé pour le genre Auyantepuia, avec néanmoins une forte concentration d’espèces dans la région à l’est de la ‘Guayana’. Le modèle de distribution géographique observé pour les espèces du genre Auyantepuia est très certainement une conséquence des vicissitudes paléoclima- tiques survenues dès la fin du Cénozoïque et surtout au cours du Pléistocène (voir Lourenço, 1986). DIAGNOSE RÉVISÉE ET SIMPLIFIÉE POUR LE GENRE AUYANTEPUIA: Scorpions de petite taille, avec une longueur totale allant de 24 à 28 mm. Coloration générale chatain rougeätre, avec les pattes et les chélicères parfois d’une coloration plus claire, jaunätre. Tégument presque toujours lisse, chagriné. Pinces des pédipalpes avec les doigts très courts par rapport à la longueur de la main (cette morphologie des pinces aboutit à une position des trichobothries db et esb souvent au même niveau, voire plus basale que celle de Ets.) Trichobothriotaxie, néobothriotaxique majorante. Face ventrale de l’anneau V du metasoma avec des gros granules spiniformes qui forment un arc dans la région postérieure, ce caractère étant diagnostique pour le genre (Figs 1 et 2). Peignes réduits en taille, avec 5 à 8 dents. ADDITIONS A LA FAUNE DES SCORPIONS DE L’ETAT DU AMAPA, BRESIL 7 COMPOSITION DU GENRE AUYANTEPUIA: Auyantepuia amapaensis Sp. n. (Brésil) Auyantepuia scorzai (Dagert, 1957) (Venezuela) Auyantepuia fravalae Lourenço, 1983 (Guyane française) Auyantepuia gaillardi Lourenço, 1983 (Guyane française) Auyantepuia sissomi Lourenço, 1983 (Guyane française) Auyantepuia parvulus (Pocock, 1893) (Brésil) Auyantepuia kelleri (Lourenço, 1997) (Guyane française) Auyantepuia skuki (Lourenço & Pinto da Rocha, 2000) (Brésil) Auyantepuia mottai Lourenço & Araujo, 2004 (Brésil) DESCRIPTION DE LA NOUVELLE ESPÈCE Auyantepuia amapaensis sp. n. Figs 6-19 MATÉRIEL: Brésil, Etat de l’Amapä, région du ‘Monte Tipac’, 240 m alt., IV/1973 (F. Cardoso leg.). Holotype mâle. ETYMOLOGIE: Le nom spécifique fait référence à l’Etat de l’ Amapä où la nou- velle espèce a été collectée. DIAGNOSE: Scorpions de petite taille, avec une longueur totale d’environ 18 mm. Coloration générale de jaunätre à jaune-rougeatre, avec les pattes et les chélicères plus claires; carapace et tergites avec des taches brunâtres bien marquées; tégument presque lisse, chagriné; carènes latéro-ventrales esquissées sur les anneaux I et II du metasoma; face ventrale de l’anneau V avec des granules spiniformes qui forment un arc dans la région postérieure. Peignes de grande taille, avec 7-6 dents. Auyantepuia amapaensis Sp. n. est une espèce voisine de À. sissomi, dont elle se distingue par: (i) une coloration générale jaune clair avec des taches brunätres bien marquées sur la carapace et les tergites, (ii) une granulation plus faible sur la pince des pédipalpes, (iii) une taille globale nettement plus petite (voir Tableau I). DESCRIPTION (basée sur le mâle holotype): Coloration générale de jaunâtre à jaune-rougeätre. Prosoma: Plaque prosomienne jaune foncé avec des zones très assombries situées en avant et sur les régions latérales de la plaque; les zones des sillons et la région postérieure un peu plus claires; tubercule oculaire noirätre. Mesosoma: Tergites jaunâtres avec des plages confluentes brun foncé. Metasoma: Tous les anneaux d’une coloration de jaunätre à jaune-rougeätre, avec des taches brunätres présentes sur les faces latérales et dorsale; face ventrale jaunâtre, sans taches. Vésicule jaunätre avec l’aiguillon rougeätre. Sternites jaunätres; peignes et opercule génital, jaune clair; sternum, hanches et processus maxillaires de la même couleur que les sternites. Pattes jaune clair avec des taches brunätres, un peu diffuses. Pédipalpes jaune-rougeatre avec des taches brunätres sur le fémur et très partiellement sur le tibia. Chélicères jaunâtre avec des taches brunätres très diffuses à la base des doigts; doigts jaunätres avec des dents légèrement rougeätres. MORPHOLOGIE: Prosoma. Plaque prosomienne légèrement concave fronta- lement (Fig. 7). Tubercule oculaire en position antérieure par rapport au centre de la plaque prosomienne. Ecartement des yeux médians un peu supérieur à un diamètre W. R. LOURENCO & J.-X. QI Fics 7-8 Auyantepuia amapaensis Sp. n., holotype mâle. (7) Habitus. (8) Yeux latéraux en détail. ADDITIONS A LA FAUNE DES SCORPIONS DE L’ETAT DU AMAPA, BRESIL à & Et E % ? Fics 9-19 Auyantepuia amapaensis sp. n., holotype mâle. Trichobothriotaxie. (9-11) Pince du pédipalpe, vues externe-dorsale, ventrale et interne. (12-14) Tibia du pédipalpe, vues dorsale, externe et ventrale. (15) Fémur du pédipalpe, vue dorsale. (16) Sternum, opercule génital et peignes, vue ventrale. (17-18) Chélicére, vues dorsale et ventrale. (19) Tranchant du doigt mobile. 10 W. R. LOURENCO & J.-X. QI Fic. 20 Carte d’ Amérique du Sud Tropicale, avec la distribution des espéces du genre Auyantepuia dans les regions Guayana et Amazonienne. Auyantepuia scorzai (cercle avec étoile noire). A. fravalae (fleur noire). A. gaillardi (carré noir). A. sissomi (étoile noire). A. parvulus (cercle avec triangle noir). A. kelleri (triangle). A. skuki (cercle avec carré noir). A. mottai (cercle noir). A. amapaensis sp. n. (cercle avec fleur noire). N = Serra do Navio. oculaire. Deux paires d’yeux latéraux; présence d’une troisieme paire d’yeux vesti- giaux, situés derriere la deuxiéme paire (Fig. 8). Plaque prosomienne avec quelques micro granules sur les régions latérales (Fig. 8); autres zones lisses, chagrinées. Mesosoma. Tergites avec quelques rares granules trés épars dans leur région posté- rieure. Metasoma. Carènes dorsales et latéro-dorsales esquissées sur les anneaux I à IV; carenes latéro-ventrales esquissées sur les anneaux I et II; les autres carènes absentes. Face ventrale de l’anneau V avec une granulation spiniforme et des granules qui for- ment un arc dans la région postérieure. Vésicule aplatie avec des gros granules spini- formes sur la région antérieure de la face ventrale; aiguillon bien plus court que la vési- cule (Fig. 6). Sternites a stigmates arrondis; tégument lisse. Peignes de grosse taille avec 7-6 dents, sans fulcres (Fig. 16). Pédipalpes. Fémur à carènes dorsale externe, dorsale interne et ventrale interne moyennement marquées, presque completes; tibia a carene dorsale interne esquissée; pince pratiquement lisse, sans carénes; fémur avec une faible granulation sur la face interne; face dorsale de la pince avec quelques granules; les autres faces lisses. Tranchant du doigt mobile avec une série linéaire de granules divisée en 6 séries par des granules plus gros (Fig. 19). Pattes. Télotarses avec des nombreuse soies irrégulièrement distribuées. Chélicères avec la dentition caractéristique des Chactidae (Vachon, 1963); présence d’une faible serrula sur la face ventrale du doigt mobile (Figs 17-18). Trichobothriotaxie du type C; néobothriotaxie majorante (Vachon, 1974) (Figs 9 à 15). ADDITIONS A LA FAUNE DES SCORPIONS DE L’ETAT DU AMAPA, BRESIL 11 TABLEAU I: Mensurations (en mm) de l’holotype male d’Auyantepuia amapaensis sp. n. et de 1°- holotype femelle d’A. sissomi (d’apres Lourengo, 1983). Auyantepuia amapaensis Sp. n. Auyantepuia sissomi* Longueur totale 17,9 26,2 Prosoma - Longueur 2,8 3,6 - Largeur antérieure 179 24 - Largeur postérieure 2,8 3,6 Anneau caudal I - Longueur 173 145 - Largeur 1,8 2,4 Anneau caudal V - Longueur 2,2 310; - Largeur 1,4 1697 - Hauteur 1,1 1,4 Vesicule - Largeur 1,0 2,1 - Hauteur 0,7 1,4 Pédipalpe - Fémur longueur 17 2,1 - Fémur largeur 0,9 1,1 - Tibia longueur Dell DET] - Tibia largeur 1,0 1,3 - Pince longueur 3,5 54 - Pince largeur 1,3 2,0 - Pince hauteur 1,7 2,3 Doigt mobile - Longueur 2,0 2,6 * Valeurs données à titre comparatif. REMERCIEMENTS Nous sommes très reconnaissants à Jean-Paul Mauries, Dépt. de Systématique et Evolution, Muséum national d'Histoire naturelle, Paris, pour la révision des versions préliminaires du texte. RÉFÉRENCES BIBLIOGRAPHIQUES FRANCKE, O. F. & Boos, J. 1986. Chactidae (Scorpiones) from Trinidad and Tobago. The Journal of Arachnology 14(1): 15-28. GONZALEZ-SPONGA, M. A. 1978. Escorpiofauna de la region oriental del Estado Bolivar, en Venezuela. Roto-Impresos C.A., Caracas, 217 pp. HJELLE, J. T. 1990. Anatomy and morphology (pp. 9-63). In: POLIS, G. A. (ed.). The Biology of Scorpions. Stanford Univ. Press, Stanford, 587 pp. LOURENÇO, W. R. 1982a. Révision du genre Ananteris Thorell, 1891 (Scorpiones, Buthidae) et description de six espèces nouvelles. Bulletin du Muséum national d Histoire naturelle, Paris, 4e sér., 4 (A1/2): 119-151. LOURENÇO, W. R. 1982b. Révision du genre Rhopalurus Thorell, 1876 (Scorpiones, Buthidae). Revue Arachnologique 4: 107-141. LOURENÇO, W. R. 1983. La faune des Scorpions de Guyane française. Bulletin du Muséum national d'Histoire naturelle, Paris, 4e sér., 5 (A3): 771-808. 12 W. R. LOURENCO & J.-X. QI LOURENCO, W. R. 1986. Diversité de la faune scorpionique de la région amazonienne; centres d’endémisme; nouvel appui a la théorie des refuges forestiers du Pléistocéne. Amazoniana 9(4): 559-580. LOURENCO, W. R. 1994. Biogeographic patterns of tropical South American scorpions. Studies on Neotropical Fauna and Environment 29(4): 219-231. LOURENCO, W. R. 1996. The biogeography of scorpions. Revue suisse de Zoologie vol. hors sé- rie II: 437-448. LOURENCO, W. R. 2002. Scorpions of Brazil. Les Editions de L’IF, Paris, 320 pp. LOURENGO, W. R. & SOUZA ARAUJO, J. 2004. Nouvelles considérations sur le genre Auyantepuia Gonzalez-Sponga (Scorpiones, Chactidae) et description d’une nouvelle espéce pour la région de Manaus, en Amazonie brésilienne. Acta biolögica paranaense, Curitiba 33(1- 4): 1-11. MELLO LEITAO, C. 1945. Escorpiöes Sul Americanos. Arquivos do Museu Nacional, Rio de Janeiro 40: 1-468. Monop, L. & LOURENCO, W. R. 2001. A new species of Broteochactas Pocock, 1890 from Brazilian Amazonia (Scorpiones: Chactidae) (pp. 195-202). In: FET, V. & SELDEN, P. A. (eds.). Scorpions 2001, In Memoriam Gary A. Polis. British Arachnological Society, London, 404 pp. PRENDINI, L. & WHEELER, W. C. 2005. Scorpion higher phylogeny and classification, taxonomic anarchy, and standards for peer review in online publishing. Cladistics 21: 446-494. SıssoM, W. D. 1990. Systematics, Biogeography, and Paleontology (pp. 64-160). In: PoLis, G. A. (ed.). The Biology of Scorpions. Stanford Univ. Press, Stanford, 587 pp. SOLEGLAD, M. E. & FET, V. 2003. High-level systematics and phylogeny of the extant scorpions (Scorpiones: Orthosterni). Euscorpius 11: 1-175. SOLEGLAD, M. E. & FET, V. 2005. The genus Auyantepuia Gonzälez-Sponga, 1978 is confirmed as a synonym of Broteochactas Pocock, 1893 (Scorpiones: Chactidae). Euscorpius 29: 1-14. STAHNKE, H. L. 1970. Scorpion nomenclature and mensuration. Entomological News 81: 297-316. VACHON, M. 1952. Etudes sur les Scorpions. Institut Pasteur d'Algérie, Alger, 482 pp. VACHON, M. 1963. De l’utilité, en systématique, d’une nomenclature des dents des chélicères chez les Scorpions. Bulletin du Muséum national d Histoire naturelle, Paris, 2e sér., 35(2): 161-166. VACHON, M. 1974. Etude des caracteres utilisés pour classer les familles et les genres de Scorpions (Arachnides). 1. La trichobothriotaxie en arachnologie. Sigles trichobothriaux et types de trichobothriotaxie chez les Scorpions. Bulletin du Muséum national d'Histoire naturelle, Paris 3° sér., n° 140, Zool. 104: 857-958. REVUE SUISSE DE ZOOLOGIE 114 (1): 13-31; mars 2007 Aphanius almiriensis, a new species of toothcarp from Greece (Teleostei: Cyprinodontidae) Maurice KOTTELAT!, Roberta BARBIERI? & Maria Th. STOUMBOUDI? ! Route de la Baroche 12, Case postale 57, CH-2952 Cornol, Switzerland (address for correspondence) and Raffles Museum of Biodiversity Research, National University of Singapore, Kent Ridge, Singapore 119260. E-mail: mkottelat@dplanet.ch 2 Institute of Inland Waters, Hellenic Centre for Marine Research, P. O. Box 712, 19013 Anavyssos, Greece. E-mail: robertab@ath.hemr.gr, mstoum@ath.hemr.gr Aphanius almiriensis, a new species of toothcarp from Greece (Tele- ostei: Cyprinodontidae). - Aphanius almiriensis, new species, is described from a brackish water spring and from a lagoon (and its inflowing fresh- water spring) in the Peloponnese (Greece). It is distinguished by the yel- lowish caudal fin of the male that has a wide faint grey margin and by the colour pattern of the female (7-11 dark, roundish blotches on the side, more or less connected by an irregular dark midlateral stripe). Aphanius almirien- sis is critically endangered; it is possibly extinct at the type locality and the second locality is much impacted. The identity, type material and type locality of A. fasciatus are discussed, and a neotype is designated. Several species are possibly confused under the name A. fasciatus. Keywords: fish - Cyprinodontidae - Aphanius - Peloponnese - extinction. INTRODUCTION Tooth carps of the genus Aphanius are small fishes that inhabit fresh to hyper- saline waters around the Mediterranean and Red Seas and the coasts of the Indian Ocean eastwards to Pakistan and southwards to Somalia. Several endemic species are also known from inland waters in arid areas, many of them in Turkey on the Anatolian plateau (Vilwock, 1984; Wildekamp et al., 1999; Hrbek & Wildekamp, 2003). The small size of these fishes, their low dispersal ability, the isolation of their populations, the small size of the habitats and the even smaller size of the areas of occupancy, lead to the evolution of very localised endemics. A few species, usually coastal ones, have a distribution range that appears very wide, e.g., A. fasciatus (Valenciennes) that is recorded around most of the Mediterranean and its islands. In fact, the distribution of the latter is better described as a long succession of more or less isolated populations (Bianco & Taraborelli, 1988; Wildekamp, 1993; Bianco et al., 1996). While a number of studies were performed on the life history, ecology and genetics of local Aphanius populations (often within a national framework) (e.g., Kiener & Schachter, 1974; Boumaiza, 1980; Tigano, 1982; Ferrito et al., 2003), a large scale comparison of Manuscript accepted 1.11.2006 14 M. KOTTELAT ET AL. populations throughout the range of the species is still missing. Such a comparison is likely to reveal that several species have been confused under the name A. fasciatus. We describe here a new species of Aphanius from Greek waters, firstly discovered in 1997 in a brackish spring. We also clear the nomenclature and the identity of the ‘real’ A. fasciatus. There have been arguments in recent years on the name to be used for the current genus Aphanius, with Lazara’s (1995) attempt to pose Lebias as the valid name. To change the name of a genus, whose species are listed as endangered in numerous national and international legal instruments, was unfortunate, especially since a critical re-analysis of the case demonstrated this decision to be based on misinterpreted data (Kottelat & Wheeler, 2001). To close the case, Kottelat & Wheeler (2001) requested a ruling from the International Commission of Zoological Nomenclature [ICZN]. The ruling (ICZN, 2003) definitively conserves Aphanius and places Lebias on the list of names not available for the zoological nomenclature. METHODS Ideally, we tried to adopt the Phylogenetic Species Concept (Cracraft, 1989; Kottelat, 1997, 1999) as an operational tool for the evolutionary species concept (Wiley & Mayden, 2000; Mayden, 2002), which is closest to our perception of what a species is. The reality is that outside the academic world it is seldomly possible to investigate the phylogeny of a given group with enough detail to demonstrate the monophyly of all the included taxa. In the context of biodiversity inventories and conservation, alpha-level taxonomy is facing priorities strikingly different from those of academic research. The goals are different, and the approach has to be adjusted. Therefore, we often have to resort to the ‘Pragmatic Species Concept’ (Kottelat, 1995; see also Kottelat & Ng, 1994). Methods for counts and measurements follow Kottelat (2001). Standard length (SL) is measured from the tip of the snout to the end of the hypural complex. Length of caudal peduncle is measured from behind the base of the last anal-fin ray to the end of the hypural complex, at the mid-height of the caudal-fin base. Scales on lateral se- ries are counted from the anteriormost scale (the first one to touch the shoulder girdle) to the posteriormost one (at the end of the hypural complex). Scales on the caudal fin itself are indicated by “+”. Transverse scales are counted as the number of longitudi- nal scale rows. The scale row on the dorsal and ventral mid-lines is noted as “1/2”. All measurements are made point to point. Abbreviations used: CMK, collection of first author; FSJF, collection of Jorg Freyhof, Berlin; HCMR, Hellenic Centre for Marine Research (formerly National Centre for Marine Research), Athens; MHNG, Muséum d’histoire naturelle, Genève; MNHN, Muséum National d'Histoire Naturelle, Paris. COMPARISON MATERIAL Aphanius fasciatus: Italy: Sardinia: MNHN 2005-1975, neotype, 39.5 mm SL; CMK 18600, 85, 32.9-47.8 mm SL; Italy: Sardinia: Cagliari, stagno di Cagliari, Primo Bacino (salt- works); A. Cau, 22 March 2005.— MNHN A.3968, male, 29.1 mm SL; MNHN 187, 3 syntypes of Poecilia calaritana, 33.9-35.7 mm SL; MNHN 92, 1, 37.5 mm SL; CMK 18599, 17, 29.2- ALPHANIUS ALMIRIENSIS FROM GREECE 15 35.4 mm SL; CMK 18786, 3, 36.3-41.6 mm SL; FSJF 209, 6, 10.9-26.7 mm SL; Cagliari. — FSJF 183, 3, 18.1, 24.3 & 27.9 mm SL; Arbarax. Italy: MNHN 197, 1 female, 43.7 mm SL, possible syntype of Lebias flava; Napoli. — MNHN 1888-112-115, 4, 31.8-33.0 mm SL; MNHN 1899-249, 5, 40.7-46.0 mm SL; Venezia. France: Corsica: MNHN 1985-204, 1 (female), 44.2 mm SL. — MHNG uncat., 8, 26.1- 42.0 mm SL; MHNG uncat., 2, 35.1-43.3 mm SL; MHNG 1321.34-38, 5, 26.8-29.2 mm SL; Bonifacio. Greece: CMK 17015, 17; CMK 16976, 6; Messolonghi. - HCMR 1471, 14; CMK 18452, 2; Kato Basiliki. - HCMR 1121, 14; CMK 18541, 2; Prokopos. -HCMR 1188, 7; Kaiafa lake. - HCMR 1318, 170; Thermissia lagoon. - HCMR 1319, 48; Sachtouri lagoon. - HCMR 1321, 74; Metochi lagoon. - HCMR 1390, 16; Aitoliko. - HCMR 1418, 29; Gialova lagoon. Turkey: MNHN 1927-79-80, females, 31.9-54.1 mm SL; MNHN 1927-78, 20 males, 23.5-48.9 mm SL; MNHN A.3796, 10, 25.9-42.0 mm SL; Izmir. - MNHN 1928-210-211, 23; Konya. Israel: MNHN A.2809, 13, 15.4-24.9 mm SL; Ramle. Egypt: MNHN A.5254, 6, 25.6-41.1 mm SL; no locality. - MNHN 4991, 1 male, 35.0 mm SL; Suez. Tunisia: MNHN 1904-48, 8, 28.9-40.0 mm SL; no locality. - MNHN 1883-1031, 8, 26.7-36.4 mm SL.; Blidet Ahmar. Algeria: MNHN 4392, 8 syntypes of Cyprinodon doliatus, males, 22.8-34.7 mm SL; MNHN 3218, 5 syntypes of C. cyanogaster, females, 24.9-32.6 mm SL; Biskra. - MNHN A.2367,1 female, 36.7 mm SL; Touggourt. - MNHN 1999-542,2 females, 19.0-31.6 mm SL; Annaba. Aphanius sp.: MNHN 1923-29, 1 male, 28.4 mm SL; MNHN 1922-72, 1 female, 37.6 mm SL; Tunisia: Gabes. RESULT Aphanius almiriensis, new species Fig. 1 HOLOTYPE: MHNG 2654.087, 35.1 mm SL; Greece: Peloponnese: Korinthia Distr.: brackish water spring Kokosi at southern end of Almiri beach, at Kato Almiri, about 4 km south of Loutra Elenis; 37°50’32”N 23°00’58”E; R. Barbieri & A. Economou, 13 May 1997. PARATYPES: HCMR 1064, 14, 24.5-38.8 mm SL; CMK 17296, 3, 25.9-29.8 mm SL; same data as holotype. - HCMR 1473, 5, 13.4-22.1 mm SL; same locality as holotype; R. Barbieri & M. Tsatsas, 26 Oct 2003. ADDITIONAL MATERIAL (NON TYPES): HCMR 1478, 12, 14.9-29.4 mm SL; Greece: Peloponnese: channel close to sea at east side of Meligou Lagoon; 37°23’07”N 22°45’02”E; R. Barbieri & M. Stoumboudi, 29 July 2004. - HCMR 1314, 190, 20.9-37.7 mm SL; MHNG 2654.088, 3, 23.4-30.2 mm SL; CMK 18370, 11, 19.1-33.1 mm SL; same locality; R. Barbieri & A. Economou, 13 Oct 1998. - HCMR 1477, 36, 9.4-26.0 mm SL; Greece: Peloponnese: freshwater spring at edge of Meligou Lagoon; R. Barbieri & M. Stoumboudi, 29 July 2004. DIAGNOSIS: Aphanius almiriensis is distinguished from all other species of Aphanius in Europe by the yellow caudal of the male, with a wide faint grey margin (vs. hyaline, greyish, greyish blue, or bright yellow with or without a black subdistal bar) and by the colour pattern of the female (7-11 dark, roundish blotches on the side, more or less connected by an irregular dark midlateral stripe). Additional characters useful to distinguish the species (but not unique to it) are: male with 6-10 dark, broad, regularly set bars on the body; neuromasts in interorbital area in deep open grooves; 25-28+2 scales in lateral series on body; and 15-16 pectoral rays. DESCRIPTION: Main morphometric data of the holotype and 13 paratypes are given in Table 1. General appearance is shown in Fig. 1. Head with relatively massive appearance. Dorsal profile of head and body convex, with a slight nuchal concavity in largest males. Mouth subterminal, oriented upwards, lower jaw almost vertical. 16 M. KOTTELAT ET AL. TABLE 1. Morphometric data of holotype and 13 paratypes of Aphanius almiriensis from type locality and of neotype of A. fasciatus. Holotype data included in range of female values. A. almiriensis A. fasciatus female males (n= 7) females (n=7) male holotype range range neotype Standard length [mm] 3571 24.5 - 28.5 25.2 - 38.8 395 Total length [mm] 43.2 30.5 - 35.8 31.1 - 46.6 48.2 In percents of standard length Head length 31.9 30.1 - 32.1 29.9 - 31.9 29.4 Predorsal length 64.1 60.4 - 62.0 61.0 - 64.7 61.0 Prepelvic length 52.0 50.2 - 53.1 52.0 - 54.9 529 Preanal length 70.8 64.3 - 65.9 67.6 - 70.8 62.8 Body width 235 20.4 - 22.1 22.1 223:5 17.5 Head depth 19.1 18.6 - 19.8 19.1 - 19.2 21.0 Body depth 28.3 24.9 - 27.8 25.6 - 28.6 28.9 Depth caudal peduncle 16.5 16.0 - 17.6 15.2 - 16.5 17.0 Length caudal peduncle 22.8 23.4 - 25.3 22.2 - 24.1 23.8 Depth of 1st simple dorsal ray 14.7 15.0 - 16.5 1320155 13.9 Length of middle caudal rays 21.9 24.2 - 26.2 20.2 - 23.9 235 In percents of head length Head depth 59.8 58.6 - 63.8 59.8 - 63.8 71.6 Eye diameter Died 30.9 - 33.6 27.6 - 31.1 2579 Snout length 32.6 28.4 - 30.5 29.7 - 32.6 34.5 Interorbital width 49.6 47.3 - 52.6 47.7 - 51.0 39.7 Posterior extremity of maxilla under tip of snout. Eye diameter about equal to snout length. Depth of caudal peduncle 1.35-1.55 times in its length. Dorsal-fin origin in front of anal-fin origin and behind midlength of body. Dorsal fin with 9-11 (modally 10) rays; anal with 10-11 (modally 10) rays, pectoral with 15-16 rays. 25-28+2 scales in lateral series, !/2 8-9 1/2 in transverse row between dorsal origin and ventral midline in front of pelvic-fin base; !/2 6-7 1/2 on caudal peduncle. Small ctenii along posterior edge of scales on caudal peduncle in largest males. Cephalic neuromasts in interorbital area in deep open grooves (Fig. 2). Males slightly more robust than females, up to 28.5 mm SL in available material. Dorsal rounded, anal and caudal truncate. Females larger than males, up to 38.8 mm SL, body less laterally compressed. All unpaired fins rounded. Anus and genital openings separated. COLOUR PATTERN: Based on the type series: Males with 6-8 dark bars on body, wider above than below, more or less reaching ventral midline and more or less fused along dorsum in front of dorsal-fin origin. Caudal and anal fins yellow, caudal fin with a greyish distal band. Dorsal fin yellowish, with an obvious black margin along anterior and upper margins, and a few dark dots near its base. Females with 7-11 dark, roundish blotches on side, more or less connected by an irregular dark midlateral stripe; no remains of other colour marks. No observation on life colour of material from Almiri, but unpaired fins of males probably brighter yellow in life than in Fig. 1. In material from Meligou, all adult males with yellow caudal and anal fins. ALPHANIUS ALMIRIENSIS FROM GREECE 7, Fic. 1 Aphanius almiriensis; Greece: Kato Almiri; CMK 17296, paratype, male, 29.8 mm SL (above); holotype, MHNG 2654.087, 35.1 mm SL (below). VARIATION: Material from Meligou lagoon generally agrees with the above des- cription, except for a single very large female (37.7 mm SL) that has the flank colour pattern consisting in slightly elongated bars. Most males have a few broad bars, but there are a few individuals with a few more and narrower bars (7-10). We do not adopt the hypothesis that these individuals may be hybrids between A. almiriensis and A. fas- ciatus, because the latter (although it is widely distributed along the eastern coast of the Peloponnese) was never observed at Meligou lagoon despite intense search for it. DISTRIBUTION: Greece, only two populations are known to date, in the spring near Almiri (Saronikos Gulf) and in the Meligou lagoon (Argolikos Gulf). In 1997- 2004, we searched about fifteen springs and brackish water systems along the coast of the Peloponnese and Southwestern Greece and found Aphanius in eleven (Fig. 3). Only A. fasciatus was observed and/or collected in all but the two aforementioned localities, including the Thermisia and Metochi lagoons that are located between the Almiri spring (Fig. 4) and the Meligou lagoon. HABITAT: Almiri spring is a permanent brackish water spring (23%o) adjacent to the sea, which it joins after a few meters. The spring itself is now contained by a concrete wall and the enclosure is filled with typical halophytic aquatic vegetation. Meligou lagoon is a permanent brackish marsh (11-25%c). Until the second half of the 20‘ century it occupied an area of about 2.6 km2. Later on, reclamation works 18 M. KOTTELAT ET AL. Immun Fic. 2 Dorsal view of head (schematised) of: Aphanius almiriensis, CMK 17269, 29.6 mm SL (left); A. fasciatus, CMK 17015, 35.8 mm SL (right). Dark grey: canal and pores in interorbital area. Scale bar 1 mm. reduced its surface area to 1.5 km2. There is a spring along the west side of the lagoon, with permanent freshwater discharge. BIOLOGY: Aphanius almiriensis is benthopelagic, observed in shallow areas with slow current, among vegetation, as well as in the water column. It is a very fast swimmer and forms schools. It is a short-lived species and a fractional spawner that breeds in late spring - early summer, spawning 1-3 eggs a time. However, larvae were collected in Meligou in September 1998 (HCMR 1314), showing that the reproductive period may be prolonged until late summer. CONSERVATION STATUS: Aphanius almiriensis was discovered in 1997 in the Almiri spring, which at the time released brackish water of constant salinity. During various visits at this location, between 1997 and 2000, the observed salinity was 23%o. However, in 2002 the spring was releasing pure seawater and the only fish observed there were mugilids. At the same time, the spring was occupied by unidentified sea grasses and sea anemones. The species was then believed to be extinct, but a visit in October 2003 yielded 5 juveniles 13.4-22.1 mm SL (HCMR 1473). In July 2004, no A. almiriensis were observed in Almiri despite intensive search. After realizing the distinctness of the Almiri population, older collections were re-examined for possible additional localities with A. almiriensis and a sample obtained in Meligou lagoon in October 1998 was discovered. In that sampling ALPHANIUS ALMIRIENSIS FROM GREECE 19 FIG. 3 Distribution of Aphanius almiriensis (circles) and À. fasciatus (squares) in the water systems in- vestigated along the coast of Peloponnese and southwestern Greece. T, type locality. occasion, A. almiriensis was estimated as abundant in the main freshwater spring supplying the lagoon at its NW side. However, recently, this spring was dammed with rocks, to create a “natural swimming pool”. During a visit in July 2004, no fish were present in the modified spring area. However, A. almiriensis was collected from different areas of the lagoon. According to the above, the species is considered to be critically endangered (CR A2ace+3ce; Blab(i,ii,iii,iv,v)+B2ab(i,ii,iii,iv,v)c(iv) according to IUCN criteria. 20 M. KOTTELAT ET AL. Fic. 4 Brackish water spring at Kato Almiri (Greece), type locality of Aphanius almiriensis. DISCUSSION COMPARISON WITH A. FASCIATUS: Aphanius fasciatus has a wide distribution, occurring along the northern coast of the Mediterranean from the Rhône delta (France) and Corsica eastwards, and along the southern coast from Algeria eastwards. It is ALPHANIUS ALMIRIENSIS FROM GREECE 21 missing in France west of Camargue, along the Spanish coast and in western Algeria (Wildekamp, 1993: 48; Römer, 1991). The species is reported to be variable in shape, colour pattern and coloration (e.g., Wildekamp, 1993), but our observations (un- published) show that part of this variability in fact should be re-interpreted as inter- specific differences. The aim of the present paper is the description of A. almiriensis, which urgently needs a formal name. Conservation measures for a critically endangered species are impossible to be implemented if this does not have a formal name. A species without a name simply does not exist for policy makers, even though scientists recognise it. Our aim is not to revise A. fasciatus sensu lato, which we reserve for a later opportu- nity. In this context, we compare A. almiriensis with the populations of A. fasciatus present in coastal waters adjacent to the range of A. almiriensis, and with the popula- tion at the type locality of A. fasciatus (with the available data, these populations of A. fasciatus are conspecific). Our logic in doing so is that, as discussed below, we doubt the conspecificy of several of the populations usually referred to A. fasciatus. At the present stage what matters is to show that A. almiriensis is distinct from the species in adjacent waters, distinct from topotypical A. fasciatus, and distinct from the nominal species placed in the synonymy of A. fasciatus. To resolve these problems, a com- parison of A. almiriensis with remote populations (species) (e.g., from France, Algeria, Egypt) is irrelevant, whatever the academic interest might be. For the present discussion, data on A. fasciatus are based on material obtained in Greece (Messolonghi, Kato Basiliki, Prokopos) and Sardinia (Cagliari; type locality), unless otherwise stated. Three valid species of Aphanius have so far been recorded in European waters and A. almiriensis is easily distinguished from them. In A. iberus (Valenciennes) from Spain, the caudal fin of the male is hyaline to bluish-grey, with 2-5 dark grey bars; the female has numerous dark brown spots on the side and the back. In A. baeticus Doadrio, Carmona & Fernandez-Delgado, also from Spain, the caudal fin of the male is dark grey, with one distinct submarginal bar and 4-5 vertical rows of hyaline dots sometimes organised in regular bars; the female has a few large black spots on the side, usually organised in two rows, one on the middle of the flank, one at the level of the pectoral-fin base. In A. fasciatus, the male has 8-15 dark bars on the body; the female has 11-17 narrow, elongated dark bars occupying about half of the body depth on the side of the body, over a faint greyish midlateral stripe (Fig. 5b). The males of A. almiriensis and A. fasciatus share the yellow caudal fin, but in A. almiriensis the caudal fin has a greyish distal edge, while in A. fasciatus the caudal fin is plain yellow (bright yellow in life) with or without a black subdistal bar (Figs 5a, 6). The colour pattern of the female A. almiriensis is not observed in other peri- Mediterranean Aphanius. The female colour pattern of many populations referred to A. fasciatus remains undescribed or only succinctly described, and reported as very vari- able. Our examination of material from all around the Mediterranean shows that while there is some intra-population variability (as for any other character), this intra-popu- lation variability is limited, and that the inter-population variability is not random but corresponds to a zoogeographic pattern and defines a number of (diagnosable) clusters of populations. The above-mentioned A. fasciatus female colour pattern is that of the 2 M. KOTTELAT ET AL. Fic. 5 Aphanius fasciatus, CMK 18451; Greece: Prokopos lagoon; male, 35.7 mm SL (above); female, 40.6 mm SL (below). populations of the Tyrrhenian, Adriatic, Ionian and western Aegean basins. The eastern Mediterranean populations also differ from A. almiriensis; the female colour pattern consists, among others, in a row of very short, narrow, black, vertical bars. Aphanius almiriensis also differs from A. fasciatus by its general appearence (compare Figs | and 5), with A. almiriensis having the dorsal head profile less convex than A. fasciatus, the ventral head profile more angular, the dorsal and anal fins of the male not reaching the caudal-fin base (vs. reaching), a quadrangular caudal fin (vs. somewhat trapezoidal), all characters difficult to quantify. Further, A. almiriensis has a stouter caudal peduncle (depth 1.4-1.6 times in its length, vs. 1.4-1.9), a wider inter- orbital (14.6-16.9% SL, vs. 11.6-14.4; 47-53% HL, vs. 40-47) and a larger eye (diameter 27-34% HL, vs. 24-32). The largest examined specimen of A. almiriensis is 38.8 mm SL, while the largest recorded size of A. fasciatus in Greek waters is 75.5 mm TL (estimated 60 mm SL) (Leonardos & Sinis, 1999). This is an unusually large size, and most adult specimens are in the size range 35-50 mm SL. The above comparison is based on specimens of similar sizes (A. almiriensis 24.5-38.8 mm SL, A. fasciatus 25.2-42.3 mm SL) and excludes the hypothesis that the characters distin- guishing A. almiriensis from A. fasciatus are juvenile characters or size-dependant. We also observe differences between the two species in the number and orga- nisation of the neuromasts in the interorbital areas. In A. almiriensis, there are 2 neu- romasts on each side, located in deep grooves (Fig. 2a). In A. fasciatus from Greece and the Adriatic and Tyrrhenian basins (including Sardinia and the neotype), there are three neuromasts in closed canals pierced with 3 pores (Fig. 2b). A single specimen of ALPHANIUS ALMIRIENSIS FROM GREECE 23 A. almiriensis (the largest female, 38.3 mm SL) has the grooves partly closed, with two pores. In A. fasciatus, only few of the smallest males (less than about 30 mm SL) have the three neuromasts in wide, shallow grooves (vs. conspicuous, deep grooves in A. almiriensis). In juveniles of the eastern Mediterranean populations, there are three neuromasts in shallow grooves, which start closing anteriorly at about 30 mm SL; in most large specimens the posterior half of the groove remains open. All conditions are present in the material from Tunisia and Algeria. Examination of type material, topotypes or original descriptions of all nominal species usually listed in the synonymy of A. fasciatus (see, e.g., Kottelat, 1997) shows that A. almiriensis is distinct from all of them. Their identities are discussed below. ON THE IDENTITY AND TYPE LOCALITY OF A. FASCIATUS: Although it is not within the scope of the present paper to revise A. fasciatus sensu lato, it is necessary to briefly discuss the diversity within this ‘species’, as most Aphanius populations from the east- ern Mediterranean drainages have been at some point referred to A. fasciatus. Also it needs to be ascertained that A. almiriensis is distinct from all the nominal species listed as synonyms of A. fasciatus (see Kottelat, 1997). There has been a number of publications describing or discussing variability within A. fasciatus. We will not discuss them in detail, for most suffer from being only of limited, local interest, neglecting other populations or with problematic methodo- logies. Frequent problems are the use of morphometrics and meristics without defining the methods used, and misunderstanding of taxonomic procedures. Aphanius fasciatus was originally described (as Lebias fasciata) by Valenciennes (in Humboldt & Valenciennes, 1821: 160, pl. 51 fig. 4). Valenciennes indicated neither the origin nor the number of his specimens. Kottelat (1997: 161) com- mented that Valenciennes (in Cuvier & Valenciennes, 1846: 156) later stated that these specimens were from Cagliari (Sardinia). In fact, the case is more complex. Valenciennes’ 1846 account is somewhat ambiguous and it appears that the species was most likely based on a single specimen (thus holotype) of unknown origin and that in 1846 Valenciennes re-described A. fasciatus on the basis of additional (non-type) material from Cagliari obtained by Bonelli and Marmora. After describing the Cagliari specimens, Valenciennes stated (in Cuvier & Valenciennes, 1846: 158) that he compared them with the specimen (singular) “preserved for a long time in ... the museum”. In 2005 we searched for the holotype in MNHN but it is no longer extant. Valenciennes further commented that this specimen has broken dorsal and anal fins and that the artist should have made them higher, implying that this very specimen is the model of the plate. Valenciennes’ 1821 description indicates 10-12 whitish bars while the figure shows only 8, but in 1846 he commented that the colour pattern too is not correctly depicted and that the artist had “exaggerated the bars, especially the an- terior ones, believing that they were less contrasted because of the effect of alcohol”. Valenciennes (in Cuvier & Valenciennes, 1846: 159) commented “[in 1821] I gave only the description and figure of these two species [L. fasciata and L. rhom- boidalis], whose origin I ignored. By doing this, I only wanted to complete what Mr. Cuvier [1816] had left to be done in the first edition of his Regne animal, where the only two species of the genus then known are designated as new”. In a footnote to the 24 M. KOTTELAT ET AL. very brief diagnosis of his Lebias [a vernacular name], Cuvier (1816: 199) had mere- ly mentioned that he knew two species of this genus, both new, but he neither named them nor provided any data. The meaning of Valenciennes’ last sentence may seem am- biguous, but the context and his comments pp. 146-150 make it clear that he described the two unnamed species mentioned by Cuvier [implying he named them on the basis of the material on which Cuvier based his statement]. Identification of Lebias fasciata as the species presently called Aphanius fas- ciatus goes back to Valenciennes (in Cuvier & Valenciennes, 1846: 158) who simply stated that he compared the holotype with specimens from Cagliari and that he was “convinced of their being the same”. (It is puzzling that on p. 146 Valenciennes men- tioned “one of them [...] is my Lebias fasciata, which the late Mr. Delalande very recently brought back from the vicinity of Rio de Janeiro”. The meaning of this sen- tence should not be misunderstood as ‘the specimen being brought by Delalande from Brazil’, but that Delalande had since brought specimen(s), which Valenciennes considered conspecific with his L. fasciata. Delalande [1787-1823] collected in Brazil in 1817 [Cuvier, 1828: 254; Bauchot et al., 1990: 72], which was too late for him to have collected the holotype that was already examined by Cuvier before 1816. It is difficult to understand how Valenciennes could identify his L. fasciata being identical on the one hand with a Brazilian species and on the other hand with a Sardinian fish; this may relate with the small size of these fishes, the optical equipment, the state of preservation, etc.) As the holotype was already in the MNHN in 1816, its origin might be specu- lated to be in areas from which the museum had received fish collections before that date. Noteworthy within the distribution range of A. fasciatus s.l. are the collections obtained by Geoffroy Saint-Hilaire in Egypt in 1798-1799 and by Delalande in Toulon (France) in 1813. In conclusion, the type locality of A. fasciatus remains unknown and, as the holotype is now missing, it will remain impossible to establish from the original description either its identity or its type locality. This problem can be solved only by a neotype designation. We designate here MNHN 2005-1975, 39.5 mm SL (Fig. 6), as neotype. This specimen was collected in the salt-works of Cagliari in 2005, at the same locality as the material of Bonelli and Marmora, on which Valenciennes based his 1846 description. As this locality has long been erroneously considered as type locality of A. fasciatus (see above), the present neotype designation best creates stability in defini- tively linking the name to this locality. Morphometric data of neotype are listed in Table 1. Meristic data: 10 dorsal-fin rays; 9 anal-fin rays; 12 pectoral-fin rays; 26+2 scales in midlateral series; 1/2 10 1/2 scales in transverse row between dorsal origin and ventral midline in front of pelvic-fin base; !/2 7 1/2 scales in transverse row on caudal peduncle. WESTERN MEDITERRANEAN A. FASCIATUS S.L.: The material presently available to us and the published information are still insufficient to satisfactorily answer questions like the number of species hiding under the name A. fasciatus, their distribution and their diagnostic characters. Nevertheless, it is possible to comment on some popu- lations and on the nominal species traditionally placed in the synonymy of A. fascia- ALPHANIUS ALMIRIENSIS FROM GREECE 25 FIG. 6 Aphanius fasciatus, Italy: Sardinia: Cagliari; MNHN 2005-1975, neotype, male, 39.5 mm SL (above); CMK 18600, female, 42.6 mm SL (below). tus. Up to now, attention has focussed on the western Mediterranean populations (Corsica, Sardinia, Sicily, Tyrrhenian slope of Italy, Malta, Tunisia). The different studies have been based only on a few populations and a few characters and therefore there are no characters with data homogeneously available for all populations. The colour pattern, however, is described or figured for a number of populations large enough to allow a comparison. Valenciennes’ figure (in Humboldt & Valenciennes, 1821: pl. 60) of A. fascia- tus depicts a male, as shown by the conspicuous dark bars. An interesting character shown on the figure is the absence of the black margin of the dorsal fin. Even if the distal part of the fin was missing (as stated by Valenciennes, in Cuvier & Valenciennes, 1846), the membranes between the anterior rays should have been coloured dark or black. A perusal through the pictures of A. fasciatus in the literature shows all males to display this marking, except for material from Corsica in Kiener & Schachter (1974: 323), from Sardinia in Wagner (1828: 1055, pl. 12) and a population from Tunisia in van der Zee & Vonk (1991). Kiener & Schachter (1974) figured specimens from 3 populations in Corsica (Diana and Biguglia in northeastern Corsica; Porto-Vecchio in southeastern Corsica) and compared them with material from Comacchio (northern Italy, Adriatic coast). They mention having seen males from Corsica up to 65 mm TL (that is, large adults) and state that the most salient features of the northeastern individuals when compared to the Comacchio ones are the shorter dorsal fin, a less vivid coloration and the absence of the black stripe in the anterior part of the dorsal fin. The drawings in Kiener & 26 M. KOTTELAT ET AL. Schachter show deep-bodied fishes with a general appearance and markings similar to those of Valenciennes’ figure (their photograph -Fig. 6- shows a more slender indi- vidual). Kiener & Schachter (1974: 324) comment that a few of the Porto-Vecchio individuals show the characters of the Comacchio population, albeit attenuated. They also mention having seen material from Sardinia, but do not mention coloration; they only mention a shorter dorsal fin than the Comacchio specimens. Kiener & Schachter’s material is not known to have been preserved; it is not in MNHN collections. We could not find other published figures of Corsican populations, but S. Valdesalici provided us with photographs of fishes from Furiani (northeasten Corsica) without the black margin in the dorsal fin and from Porto-Vecchio (southeastern Corsica) with obvious black margin. Males from Bonifacio (southern Corsica; MHNG uncat.) have the obvious black margin. The account of A. fasciatus in Keith & Allardi (2001: 278) shows a fish with a faint dark distal margin (but no dark anterior margin). However, we cannot give any value to this drawing as it is obviously based on the one in Maitland (1976), itself based on unknown sources and not very realistic. Valenciennes (in Cuvier & Valenciennes, 1846: 156) re-described A. fasciatus based on fresh material from Cagliari, Sardinia. The colour pattern he described for the males agrees with what was reported by Kiener & Schachter (1974) from Comacchio and by van der Zee & Vonk (1990) as ‘short’ form from Tunisia. Valenciennes (in Cuvier & Valenciennes, 1846: 151) also described a Cyprinodon calaritanus from the same swamp in Cagliari as the neotype of A. fascia- tus, which is the female of his 1846 A. fasciatus (confirmed by examination of 3 syn- types MNHN 187). He recorded the presence of narrow bars on the body and one or two, sometimes three, black spots at caudal-fin base, which is also agreeing with the Comacchio females figured by Kiener & Schachter and others. (Lebias calaritana is a name available from a very brief diagnosis by Cuvier [1829: 280], from which one deducts that the name is based on material from Sardinia sent by Bonnelli and that most likely is the material used by Valenciennes in 1846.) Wagner (1828) figured individuals from Cagliari, Sardinia (his figures show his Lebias sarda and L. lineopunctata, unambiguously male and female, respectively, of the same species). The type material of both nominal species is lost (Kottelat, 1997). The male has a relatively deep body and a short dorsal fin. There is no indication of a black anterior margin in the dorsal fin. The female has very narrow bars on the body, the posteriormost ones replaced by a number of small spots over the whole depth of the caudal peduncle. This pattern is also distinct in the Corsican females figured by Kiener & Schachter (1974) and in some populations (not all) from Tunisia figured by Boumaiza (1980: 87) (the figured Tunisian males have a large dorsal fin with a black margin). The variability among the different accounts of material from Cagliari is intriguing, but can only be interpreted after more populations can be examined. Comparing Valenciennes’ and Wagner’s descriptions and figures and our material from Cagliari remains inconclusive. It might be that Wagner’s figure is faulty or based on a not well preserved specimen; or it might mean that two ‘forms’ or species are (or were) present in Sardinia; or, that Wagner might have based his descriptions on material with erroneous locality data. ALPHANIUS ALMIRIENSIS FROM GREECE 27 Lebias nigropunctata Schinz (1840: 334) is also described from Sardinia, but the text is uninformative and does not allow to objectively identify the species; there is no known surviving type material (Kottelat, 1997). The identity of these three nominal species based on non-extant Sardinian material can only be cleared by the designation of neotypes. We designate MNHN 2005-1975 (the neotype of A. fasciatus) as the common neotype of Lebias sarda Wagner, 1828, L. lineopunctata Wagner, 1828, and L. nigropunctata Schinz, 1840. This makes them objective synonyms of A. fasciatus. We have seen photographs of A. fasciatus from Sardinian localities provided by S. Valdesalici. Adult males from Stagno di Pilo, near Porto Torres (northwestern Sardinia) have no black margin on the dorsal fin, while the black margin is present in specimens from northeastern Sardinia (no precise locality data). Some data on the genetic diversity within the western Mediterranean popu- lations is available, however in a format presently not compatible with a taxonomic analysis. Nevertheless, they somehow provide information paralleling the observations on the colour pattern. An electrophoretic analysis based on 11 populations from Corsica, Sardinia, Sicily, the Tyrrhenian and Adriatic coasts of Italy, and Tunisia shows that the 3 Sicilian and Tunisian populations constitute one unit, the 2 Adriatic ones a second unit, and the Tyrrhenian, Sardinian and Corsican ones a third unit (Maltagliati, 1999). Within that last unit, the mainland and Corsican populations are closer to each other than to the Sardinian ones. Another allozyme study of 20 Corsican and Sardinian populations shows that the Corsican populations have closest similarity with some populations from the northeastern tip of Sardinia (Maltagliati, unpublished, pers. comm.). Tigano et al. (2004) noted differences in the number and location of NOR loci on chromosomes of Adriatic and Sicilian populations. Tigano et al. (1999) report differences in the morphology of pharyngeal bones between Sicilian and Adriatic populations. Ferrito et al. (2003) compared some mor- phological traits and allozymes of two Sicilian, one Sardinian and one Adriatic popu- lations. Although the number of populations is limited, the three units reported by Maltagliati (1999) also show up in these studies. Wildekamp (1993: 50) figures a specimen from Malta with more and narrower bars than the Adriatic ones, probably similar to the Tunisian ones. (Gulia [1861: 11] made a laconic description of two new species from Malta, Micromugil timidus and M. macrogaster, apparently the male and the female; there is no known type material.) Material from Sicily is figured by Wildekamp er al. (1999: 35) and its fins by Tigano (1982: 175). Material from Tunisia is figured by Boumaiza (1980) and van der Zee & Vonk (1991; see below). In all, the pre-dorsal bars tend to be vertically split in at least their lower extremity, and there is a tendency to have very thin bars intercalated between larger ones. This is not the case in material, figures and photographs we have seen from the Adriatic, Greece and Turkey (pers. obs.; Wildekamp, 1993 and unpub- lished photographs); in these populations, the bars tend to be fewer, wider and more regular. Cyprinodon cyanogaster and C. doliatus were both described by Guichenot (1859: 379) from Biskara in northeastern Algeria. They are respectively female and male, and apparently conspecific (8 syntypes of C. doliatus MNHN 4392 and 5 syn- types of C. cyanogaster MNHN 3218 examined). Presently, we are unable to distin- 28 M. KOTTELAT ET AL. guish them morphologically from the Sardinian populations of A. fasciatus and we consider them as conspecific. The Algerian and most of the Tunisian material that we examined, as well as that figured by Boumaiza (1980) and van der Zee & Vonk (1990; ‘short form’), also are conspecific. As observed in A. fasciatus, the syntypes of C. do- liatus (males) have the black margin in the dorsal fin and the black subdistal bar on the caudal fin and the syntypes of C. cyanogaster (females) have the small spots on the caudal peduncle. Van der Zee & Vonk (1990) have photographs of live specimens of an Aphanius from Tunisia with a deep body, short dorsal fin, pallid coloration and without dark margin in the dorsal fin. It differs from A. fasciatus, as now fixed by the neotype, in having more dark bars (about 20 vs. 8-15). Most interesting is that van der Zee & Vonk report the presence of two ‘forms’ of A. fasciatus in sympatry in Lake Kelbia drainage, the one mentioned above, which they call normal form (their figure 2), and a more stocky one, which they call ‘short’ form with a vivid coloration, fewer bars and a very conspicuous dark margin in dorsal fin. The two ‘forms’ reportedly also differ in their way of swimming (Wildekamp, 1993: 50). This is strongly suggesting that two species are involved: the ‘short’ form with an appearance identical to that of the Sicilian populations, and the ‘normal’ form possibly endemic. We had the possibility to exa- mine only two specimens that possibly belong to this ‘normal’ form (Gabes, Tunisia; MNHN 1922-72, 1923-29). Concluding, the data and material presently available to us justify investigating the possible existence of at least two species within A. fasciatus s.l. in the Western Mediterranean: 1) A. fasciatus from the Tyrrhenian basin, Sicilia, Malta, Tunisia and Algeria; and 2) an unnamed species from Tunisia. The status of some Corsican and Sardinian populations (without the black margin on the dorsal fin) also requires inves- tigation. There are no available names for the Tunisian species or the last mentioned Sardinian-Corsican populations. EASTERN MEDITERRANEAN A. FASCIATUS S.L.: Data on the Eastern Mediterranean populations are less numerous and less detailed. The earliest record of Aphanius from the Adriatic is apparently by Nardo (1827: 488) who recorded A. fasciatus and described a new species (A. nanus). Nardo did not give precise locality data, but his introductory paragraph makes it clear that his work was based mainly on his obser- vations along the shores of Istria and additional observations in Dalmatia and Romandiola (Ravenna Province, Italy). Although brief, his account of A. fasciatus refers to 8-9 bluish bars on body and dorsal fin black anteriorly. His A. nanus is diag- nosed by having 12-13 irregular black bars and black spots and refers to the female. Lebias flava Costa (1838: fasc. 19: 35, pl. 17 fig. 1) was described from lake Varano, on the Adriatic shore of southern Italy. Valenciennes (in Cuvier & Valenciennes, 1846: 159) saw no difference between Costa’s plate and his material from Sardinia. Wildekamp (1993: 48) figures a male from Dalmatia with a colour pattern sim- ilar to that of the Comacchio (near Ravenna, Italy) material figured by Kiener & Schachter (1974). The material of A. fasciatus (of unknown origin) figured by Gandolfi et al. (1991: 617) shows the same colour pattern. ALPHANIUS ALMIRIENSIS FROM GREECE 29 Cyprinodon desioi described by Gianferrari (1932: 214, fig. 1) from north- eastern Libya has been considered as a synonym of A. fasciatus (e.g. by Wildekamp, 1993: 49). Neither the description nor the.figure mention dark markings in the dorsal fin. The figure does not show any pattern on the caudal fin, but the description mentions that there are one, two or no brown bars. With the presently available material and data, we are unable to distinguish any of these population and nominal species from A. fasciatus as recognised above. Cyprinodon hammonis Valenciennes (in Cuvier & Valenciennes, 1846: 169) (type locality: Egypt [Siwa oasis, along the Libyan border] and Syria) is listed as a synonym of A. fasciatus by Wildekamp (1993: 48), who comments that the Syria specimens are A. dispar. Valenciennes’ description of colour pattern (dorsal, caudal and pectoral fins blackish, without bars or spots) does not seem to be based on A. fas- ciatus. Hrbek & Meyer (2003) examined the phylogeny of Aphanius using mtDNA. Their study included 7 populations that they identified as A. fasciatus; 6 of them pooled closely together (Prokopis [Prokopos], Greece; Ravenna and Palermo, Italy; Tarsus, Turkey; Corsica; Malta) and one slightly more distant (Lake Bafa, Turkey), but the data cannot be used to reach taxonomic conclusions. ACKNOWLEDGEMENTS We thank Alcibiades Economou (HCMR) for sharing field work, Romain Causse, Javier Gregorio, Patrice Pruvost (MNHN), Sonia Fisch-Muller, Claude Weber (MHNG) and Jörg Freyhof (FSJF) for access or loan of material under their care, Ruud Wildekamp for making photographs of various Aphanius species available, Stefano Valdesalici for his efforts in obtaining numerous photographs and material of Italian and Corsican populations of A. fasciatus, Angelo Cau for obtaining material from the type locality, and Ferruccio Maltagliati for permitting to cite unpublished observations. REFERENCES BAUCHOT, M.-L., DAGET, J. & BAUCHOT, R. 1990. L’ichtyologie en France au debut du XIX® siècle: l’histoire naturelle des poissons de Cuvier & Valenciennes. Bulletin du Museum National d'Histoire Naturelle, Paris Sect. A, Ser. 4, 12 (Suppl.): 1-142. BIANCO, P. G., AHNELT, H. & ECONOMIDIS, P. S. 1996. 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Atlas of the oviparous cyprinodontiform fishes of the world. Volume 1. The genera Adamas, Adinia, Aphanius, Aphyoplatys and Aphyosemion. American Killifish Association, Mishawaka, 311 pp. WILDEKAMP, R. H., KÜcük, F., UNLUSAYIN, M. & VAN NEER, W. 1999. Species and subspecies of the genus Aphanius Nardo 1897 (Pisces: Cyprinodontidae) in Turkey. Turkish Journal of Zoology 23: 23-44. WILEY, E. O. & MAYDEN, R. L. 2000. The evolutionary species concept (pp. 70-89). In: WHEELER, Q. D. & MEIER, R. (eds). Species concepts and phylogenetic theory - a debate. Columbia University Press, New York. RM ti ar + ru Pr er Ng a Apr REVUE SUISSE DE ZOOLOGIE 114 (1): 33-48; mars 2007 Pupal morphology in the subfamily Paussinae (Coleoptera: Carabidae) Andrea DI GIULIO!, Andreas KAUPP?, Simone FATTORINP, Augusto VIGNA TAGLIANTI* & Peter NAGEL? ! Dipartimento di Biologia, Universita degli Studi “Roma Tre”, Viale Guglielmo Marconi 446, I-00146 Roma, Italy. E-mail: digiulio@uniroma3.it 2 Institut für Natur- Landschafts- und Umweltschutz / Biogeographie, Universität Basel, Basel, Switzerland. 3 Via Casal dei Pazzi 49, I-00156 Roma, Italy. 4 Dipartimento di Biologia Animale e dell’Uomo, Universita di Roma “La Sapienza”, Roma, Italy. Pupal morphology in the subfamily Paussinae (Coleoptera: Carabidae). - Pupae of the Ozaenini lineage (Pachyteles vignai Deuve, 2000, P. digiulioi Deuve, 2000 and Physea setosa Chaudoir, 1868) and the Paussini lineage (Platyrhopalopsis melleii (Westwood, 1833) and Paussus kannegieteri Wasmann, 1896), are described and shown in figures, and compared with other adephagan taxa. In contrast to the customary assumption that pupal characters are uninformative with regard to systematics, this analysis shows the importance of pupal characters for phylogenetic reconstruction at different levels. Differences between basal and derived Paussinae occur in the arrangement of setae and in the folding of gonothecae, while some characters, like the microsculpture, the inversion of the last abdominal seg- ments and perhaps the reduction of the 7th abdominal spiracle, support Paussinae monophyly. The extent of pubescence is a morphocline among Adephaga, with reduction in several unrelated taxa. Finally, the possibility of recognising distinct abdominal segments and antennal articles at the pupal stage, in taxa where they appear fused at the adult stage, may provide an ontogenetic basis for elucidating phylogenetic relationships. Keywords: Coleoptera - Carabidae - Paussinae - pupae - morphology - phylogeny. INTRODUCTION Coleoptera have exarate pupae with free body appendages. The body is slightly bent, with the head sharply turned downwards. The legs (podothecae) lay in a back- ward position appressed to the ventral side of the body, the pterothecae (covers of the wings) are turned downwards and partially cover the hind legs. As stated by Jeannel (1941, 1949), and Sturani (1962), both pairs of metapterothecae are generally fully developed in Adephaga, even in the pupae of non-alate species. The antennothecae are turned backwards and lay between the mid legs and the mesopterothecae. The only residual larval characters are the sometimes remnant urogomphi and the pigment of the Manuscript accepted 15.11.2006 34 A. DI GIULIO ET AL. stemmata, which is only present in young pupae, disappearing later during the pupal stage. While knowledge about larval morphology of Carabidae has grown very quickly in the last decades, the pupal morphology has rarely been investigated. Also, most of the research on the morphology of adephagan pupae has dealt with the description of the habitus, and no structure has been characterised in terms of its taxo- nomic importance. The scarce interest in pupal biology may be a consequence of the difficulties encountered in obtaining and identifying pupae, as well as the perception that pupal morphology is of minor systematic importance, generally reflecting characters of the adult stage. Although pupae exhibit a number of characters shared with adults and larvae (such as those mentioned above), they also possess some unique characters. The pupae of Coleoptera are immobile and they do not feed. Therefore the main function of the pupal integument is to protect the body as it undergoes metamorphosis. All carabid pupae have specialised setae on the dorsal surface, which are unique to this life stage (von Lengerken, 1924; Jeannel, 1949). However, the distribution of these setae differs considerably among lineages within the Carabidae (see Fig. 1). Dorsal setae may be found on the head, on one of the three tergites of the thorax, on the abdominal tergites 1 to 8, on the abdominal pleurites 2 to 7, and, if everted, also on the 9th and 10th abdominal segments. During the pupal stage, which lasts about 14 to 28 days in Carabidae (Sturani, 1962), the pupa rests in a prostrate position in the pupal chamber. These setae most likely function to isolate the body from the surface of the pupal chamber. In contrast, the ventral surface and the legs are completely glabrous, presumably because they are not in contact with the ground. Therefore the distribution of the pupal setae generally allows one to infer the resting position of the pupae within the pupal chamber. However, pupae of Amphizoidae, Hygrobiidae and Dytiscidae, are exceptions to this general rule. Although dorsal setae are fully developed in these groups (Bertrand, 1972), the pupae rest in labile equilibrium on their prolonged uro- gomphi and on the setae on the head and the anterior region of the pronotum, with their dorsal surface directed upwards and not in contact with the ground. As far as we know this resting position is unique to these taxa and represents a derived condition (Ruhnau, 1986). Very few adephagan pupae have been described. The resulting lack of knowl- edge impedes detailed phylogenetic analyses based solely on pupal characters. However, some hypotheses about the phylogenetic utility of these characters can be inferred based on our present knowledge of Adephagen phylogeny. Complete dorsal pubescence is present in Dyscolus (Carabidae, Platyninae) and Dromius (Carabidae, Lebiini), and in most Hydradephaga, where it occurs on the head, as well as all segments of the thorax and abdomen (Bertrand, 1972). This pattern indi- cates that the full set of setae covering head, pro-, meso-, metanotum, abdominal tergites 1-8 (and the following tergites if everted), and pleurites 2-8, represents an ancestral (plesiomorphic) condition. Consequently, the lack of pubescence on one of these body parts represents a derived (apomorphic) condition. For example, the pupae of Clinidium have dorsal pubesence on all segments except for the head. In other groups of carabids the dorsal pubescence is lacking from additional segments. For PUPAE OF PAUSSINAE 35 Fic. 1 Cartoons of Adephagan pupae, showing the distribution of dorsal setae in various lineages. The presence of dorsal setae in various segments is indicated by the gray color. Note that the head is drawn in unnatural position for illustrative purposes. (A) Hydradephaga, Dyscolus and Dromius. (B) Clinidium. (C) Carabus. (D) Ozaenini. (E) Paussini. example, the pupae of Carabus and Cychrus lack setae on the head, mesonotum and metanotum, in addition some species in these groups lack setae on some of the abdo- minal segments (Sturani, 1962). Another important pupal character is the number and shape of the spiracles. According to Jeannel (1949), the pupae of Adephaga possess a pair of spiracles on the mesothorax, and 6 or 7 spiracles on the abdominal segments, with the spiracles 6th and 7th closed. The 8th abdominal spiracle is always missing in both pupae and adults. The mesothoracic spiracles are closed in the pupae of Hydraenidae, Amphizoidae, Hygro- biidae and Dytiscidae (Ruhnau, 1986), open in Carabidae (Sturani, 1962), including Paussinae. Within the subfamily Paussinae, pupae have been described from only 5 spe- cies, all belonging to the tribe Paussini: Paussus kannegieteri Wasmann, 1896 (Böving, 1907), P. curtisi (Westwood, 1864) (Luna de Carvalho, 1951), P. cridae (Gestro, 1916) and P. cucullatus (Westwood, 1850) (Luna de Carvalho, 1959) and Platyrhopalopsis melleii (Westwood, 1833) (Luna de Carvalho, 1977). Most of these descriptions are too superficial for well-founded morphological comparison. In this paper, we provide the first comparative study of the pupal morphology of Paussinae. In particular, pupae of the tribe Ozaenini are described and illustrated for the first time. Detailed redescriptions and figures of Paussini pupae are also provided for comparative purposes. The pupae of Paussinae are compared with those of other Carabidae and some other adephagan groups, and the morphological characters with potential phylogenetic significance are discussed. MATERIAL EXAMINED CARABIDAE, PAUSSINAE, OZAENINI 1) Pachyteles vignai Deuve, 2000: 3 pupae (1 d and 2 9 9) and 2 pupal exu- viae (1 d and 1 2). Pupae were obtained from Ist and 2nd instar larvae reared in the laboratory. The larvae were collected along a riverbank, where they had constructed 36 A. DI GIULIO ET AL. burrows in the sandy soil. Before pupation, larvae close the entrance of these burrows with ground taken from inside, forming a rounded pupal cell (Di Giulio & Vigna Taglianti, 2001). Material labelled “Ecuador, Cotopaxi. San Francisco de las Pampas, str. S. F. de las Pampas-La Union del Toachi, Km 1.300, 1550 m, 7.XII.1997, I. Tapia leg.” and deposited in Di Giulio & Vigna Taglianti’s collection (Universitä La Sapienza, Roma, Italy). 2) Pachyteles digiulioi Deuve, 2000, one 3rd instar larva dead during the pupa- tion, labelled: “Ecuador, Cotopaxi. Between Otonga and Las Pampas, about 1700 m, 1.V1.1997, A. Di Giulio & A. Tapia leg.” and deposited in Di Giulio & Vigna Taglianti’s collection (Universita La Sapienza, Roma, Italy). 3) Physea setosa Chaudoir, 1868, 2 pupae (1 & and 1 9) and 1 male pupal exuvia collected together with larvae and adults in small holes near the *mushroom chambers” within a nest of leafcutter ants Atta sexdens (Linnaeus, 1758) (van Emden, 1936). Material labelled “Mendes, 25.9.1933, Ausgrabungsbau, in Erdhöhlen zwischen den Panellas, H. Eidmann” and deposited at the Naturhistorisches Museum, Basel (Switzerland). CARABIDAE, PAUSSINAE, PAUSSINI 4) Platyrhopalopsis melleii (Westwood, 1833), 1 male pupa, labelled “India, Kerala Periyar — Lake, près de l'Hotel Aranya Nivas, 4.X1.1972, 950 m“, preserved at the Natural History Museum of Geneva (Switzerland). The pupa is one of seven ones, collected by Besuchet in the ground at the foot of an old trunk, together with 8 larvae and 80 adults; no ants were observed nearby (Luna de Carvalho, 1977), even though Platyrhopalopsis species are considered to be myrmecophilous. 5) Paussus kannegieteri Wasmann, 1896, 3 male and 2 female pupae labelled “Java, Buitenzorg, Pangerango, 15-21.111.1904, Hjalmar Jensen” and deposited in the Zoological Museum of Copenhagen (Denmark). The pupae were collected together with some larvae and many adults inside an ant nest (Böving, 1907). CARABIDAE, PLATYNINAE 6) Dyscolus megacephalus (Bates, 1891): 2 female pupae labelled “Ecuador, Cayambe, 4300 m, 01.IX.85 P. Moret”, deposited in Moret’s collection (Toulouse, France). RHYSODIDAE 7) Clinidium canaliculatum (Costa, 1839), 1 female pupa labelled “Italia, Calabria, Pollino, Colle Gaudolino, 10.VII.71, V. Vomero”, deposited in Di Giulio & Vigna Taglianti’s collection (Universita La Sapienza, Roma, Italy). METHODS All pupae were studied and drawn by using a stereomicroscope equipped with drawing tube. The description of chaetotaxy refers to only one half of the pupal body, since the positioning of the setae are nearly symmetrical on both sides of the body. One female pupa of Pachyteles vignai, was mounted on stub after critical point dehydration and gold sputtering, and examined and photographed (Figs 5-11) using a scanning electron microscope (S.E.M.) Philips XL 30 (Interdepartmental Laboratory of Electron Microscopy, L.I.M.E., University ‘Roma Tre’, Rome, Italy). PUPAE OF PAUSSINAE 37 The pupa of Platyrhopalopsis melleii was examined and photographed using a Philips E.S.E.M. (Labor für Raster-Elektronenmikroskopie, Universität Basel, Switzerland) without drying and sputtering. All pupae (except that of P. vignai mounted on a stub) are preserved in 70% ethanol. The characters of Carabus and Cychrus were obtained from Sturani (1962), those of Dromius from Casale et al. (1996), and those of Hydradephaga from Bertrand (1972). RESULTS Pachyteles vignai Deuve, 2000 Figs 2-11, 22-23 Pupa exarate, whitish, translucent; eyes and apex of mandibles brownish in more mature specimens; abdomen cylindrical (Figs 2-5). Total length about 6 mm. Microsculpture. Consisting of small subtriangular thorns regularly dispersed all over the body. Gonothecae of female corrugated (Fig. 10). Setae on notum and pleurae longitudinally corrugated (sensilla chaetica), inserted on small conical tubercles (el). Head. Bent downward; general characters as in the adult (Figs 2, 3, 5); two setae between the antennal base and the eye (Figs 2, 3) and one medial seta on the ver- tex (Fig. 3); antennothecae folded between mid legs and mesopterothecae. In newly formed pupae, the pigment of larval stemmata (one on each side) is still visible at the position of the compound eyes. Thorax. Pronotum transverse (Fig. 4), with 17-19 setae on each side of the medial groove, positioned as follows: one group of 7-8 long setae anterior, 6-7 setae along the lateral margin and one group of 4 setae, shorter than the others, in the middle of the posterior part of the discal area. Medial groove not apparent. Mesonotum strongly transverse, without setae. Mesothoracic spiracle annular. Metanotum with 5 long setae on each side, longitudinally lined on the discal area; metaptheroteca longer than mesopterotheca (Fig. 2), ending at the level of the abdominal segment IV. Podothecae smooth, with tarsi touching one another along the midline; metatarsi reaching sternite VIII (Fig. 3). Abdomen. Segments I-VIII visible in dorsal view (Fig. 4); segments IX-X inverted into the body, urogomphi absent. Tergite I with 3 long dorsal setae on each side, transversally lined; tergites II-VII with only one pair of dorsal setae on each side; tergite VIII with 7-8 setae on the posterior margin of the abdomen; pleurae of segments II-VII with 3-6 very small setae each; pleural areas of segment VIII produced in two conical protuberances, without setae (Figs 2, 22, 23). Spiracles dorsolateral (Figs 2, 4, 5), annular (Fig. 6), with a small posterior area rich of pores and with possible glan- dular function (Fig. 6, arrow); spiracles of first segment slightly wider and more cylin- drical than the following (see Fig. 6). Sternite I divided by the metacoxae in two lateral sclerites; sternites II-VII entire, 6 separate sternites distinguishable, sternites I-III not fused. Male gonotheca (Fig. 22) asymmetrical, well visible, composed of a medial swollen and multilobed area, aedeagus not clearly turned toward the left side, as in the majority of carabid pupae (Jeannel, 1941; Sturani, 1962); female gonothecae (covering the two gonapophyses) conical and symmetrical (Figs 5, 10, 23), pointed at the tip, touching one another along the midline. 38 A. DI GIULIO ET AL. Pachyteles vignai, male pupa: (2) habitus from left lateral, (3) ventral and (4) dorsal view. Scale bar = 1 mm. PUPAE OF PAUSSINAE 39 2 PA 9 7095 Fics 5-11. Pachyteles vignai, female pupa: (5) habitus, right lateral view; (6) right abdominal spiracle II with posterior glandular area (arrow); (7) claw of right metathoracic leg; (8) group of setae on abdominal tergum VIII; (9) pronotum, right lateral view; (10) microsculpture at apex of right gonoteca; (11) microsculpture on head. Scale bars: Fig. 5 = 1 mm; Figs 6, 11 = 20 um; Figs 7,8 = 100 pm; Fig. 9 = 500 um; Fig. 10 = 10 um. 40 A. DI GIULIO ET AL. A male pupa of Pachyteles digiulioi, deceased during pupation and partially still enveloped by the 3rd instar larval exuvia, shows morphological characters, including the chaetotaxy, very similar to those of Pachyteles vignai. Physea setosa Chaudoir, 1868 Figs 12-14, 24-25 The general pupal morphology and chaetotaxy of Physea setosa (Figs 12-14) and Pachyteles vignai are similar. Differences in the habitus, mostly of the prothorax, abdomen, legs and gonothecae (Figs 24, 25), reflect those of the adults. Physea setosa is one of the few myrmecophilous species of Ozaenini with adaptations for living with ants during both the adult stage (e.g. flattening of legs and body) and larval stage (e.g. physogastry of the body, cf. van Emden, 1936, Di Giulio et al., 2003). In the follow- ing, we concentrate on the differences between the two taxa. Most setae of the speci- mens examined are broken, but their position is marked by small conical tubercles. Total length about 14 mm. Head. One seta on the vertex near the antennal base; frons with 2 small swollen areas. Thorax. Pronotum strongly transverse, flattened, with 20-25 setae on each side of the medial groove, disposed as in the following: one group of 8-10 anterior setae, 7- 10 setae irregularly dispersed from the anterior corner to the lateral side and one group of 4-6 setae, medially on the posterior part. Mesonotum transverse and slender, with- out setae. Metanotum with 5 setae on each side, transversally lined on discal area; stig- ma of mesothorax well developed; metaptheroteca smaller than in Pachyteles vignai in respect to body size, ending at the level of abdominal segment III. Podothecae smooth, with pro- and mesotarsi touching one another along the midline; metatarsi reaching the sternite IV. Abdomen. Tergite I with 3 long dorsal setae on each side, transversally lined; tergites II-VII with only one pair of dorsal setae on each side; tergite VII with 7-8 setae on the posterior margin of the abdomen; pleurae of segments I-VII each with 2-6 very small setae; pleural areas of segment VIII produced in two small conical protuberances, without setae; sternites like in Pachyteles vignai. Male gonotheca (Fig. 24) asymmetrical, well visible, composed of a medial, swollen and multilobed area very similar to that of Pachyteles vignai; female gonothecae (Fig. 25) conical and symmetrical, pointed, smaller than those of Pachyteles vignai. Platyrhopalopsis melleii (Westwood, 1833) Figs 15-18 Highly specialised adult characters (e.g. flattened legs, lens-shaped antennal club, subrectangular and flattened abdomen, small prothorax) are also visible in the pu- pa (Figs 15-17). The main differences from the pupae of Ozaenini include a richer chaetotaxy of notum and pleurae, and a different position of the gonothecae, partially hidden in the large sternite VII. The number of the visible tergites and sternites is the same as in the Ozaenini. Almost all setae are broken but marked by small conical tubercles. Total length about 9 mm. PUPAE OF PAUSSINAE 4] Fics 12-14 Physea setosa, female pupa: (12) habitus from left lateral, (13) ventral and (14) dorsal view. Scale bar =2 mm. Microsculpture. Similar to that of Pachyteles vignai, with small triangular thorns (length about 3 um, distance between thorns about 15 um). 42 A. DI GIULIO ET AL. RR sl cep LE Te à Fics 15-17 Platyrhopalopsis melleii, male pupa: (15) habitus from left lateral, (16) ventral and (17) dorsal. Scale bar = 2 mm. Head. Bent downward; no setae present. Antennothecae partially covering the fore-pterothecae (as in Paussus kannegieteri Wasmann, 1896, cf. Böving, 1907); 3 sub- apical denticulations present at the posterior margin of the antennal club of the anten- notheca; in contrast to the adult, 6 or 7 antennomeres can be identified by transverse grooves, one small protuberance at the apex (Fig. 18). Thorax. Pronotum transverse, subtriangular, with more than 50 setae on each side of the medial groove on the discal area, disposed as in the following: 26-30 anterior setae, 17-18 lateral setae and 6-8 posterior setae. Medial groove not apparent. Mesonotum highly transverse, without setae; mesothoracic spiracle annular. Meta- PUPAE OF PAUSSINAE 43 Fic. 18 Platyrhopalopsis melleii: antennotheca of male pupa in dorsolateral view. Scale bar = 500 um. notum with two symmetrical, slightly swollen areas on the discal part, with 24 setae each; metaptheroteca longer than mesopterotheca, ending at the level of abdominal segment IV. Podothecae flattened, with tarsi touching one another along the midline; metatarsi reaching sternite VIII. Abdomen. Segments I-VIII visible from dorsal view; segments IX-X inverted, urogomphi absent. Tergites I-VII with 20-30 setae on each side of the midline, mostly on the discal area and on the posterolateral part; tergite VIII with more than 100 setae on each side of midline; pleurae of segments II-VII each with 10-15 small setae; pleural areas of segment VIII not clearly produced in conical protuberances. Spiracles annular, dorso-laterally positioned in a tergopleural groove. Sternites as in the Ozaenini, the basal three not fused. Male gonotheca asymmetrical, not protruding, composed of a medial swollen and multilobed area, partially hidden under the sternite VII. Paussus kannegieteri Wasmann, 1896 Figs 19-21, 26-27 Body elongate, subcylindrical, with chaetotaxy similar to that of Platyrhopa- lopsis melleii, but less rich. The setae of the available pupae are almost all broken. Total length about 6 mm. 44 A. DI GIULIO ET AL. Fics 19-21 Paussus kannegieteri, female pupa: (19) habitus from left lateral, (20) ventral and (21) dorsal view. Scale bar = 1 mm. Microsculpture. As in Platyrhopalopsis melleii. Head. Bent downward; no setae present. Antennothecae partially covering the anterior pterothecae; the 6 teeth on the posterior margin of the antennal club clearly visible; transversal sutures on the antennotheca absent; small protuberance present at apex. PUPAE OF PAUSSINAE 45 Fics 22-27 Last abdominal segments of the pupae of in ventral view: (22) Pachyteles vignai male. (23) P. vignai female. (24) Physea setosa male. (25) P. setosa female. (26) Paussus kannegieteri male. (27) Paussus kannegieteri female. Thorax. Pronotum subquadrate, divided in four parts by a distinct transverse constriction and a slight longitudinal medial groove; posterior half longer and slightly wider than the anterior; about 50 setae on each side of the medial groove, positioned as follows: about 25 anterior setae, 4-5 medial setae and 14-16 posterior setae. Mesonotum shorter than metanotum, highly transverse, without setae; mesothoracal stigma present. Metanotum with two symmetrical, slightly swollen areas on the discal part, with 7-9 setae each; metaptheroteca longer than mesopterotheca, ending at the level of abdominal segment IV. Podothecae cylindrical, slightly flattened, with tarsi touching one another along the midline; metatarsi reaching the sternite VII. 46 A. DI GIULIO ET AL. Abdomen. Segments I-VII visible from dorsal view; segments IX-X inverted, urogomphi absent. Tergites I-VII with 6-8 setae on each side of the midline, disposed as in the following: 5-7 setae lined along the posterolateral part and 1 anterior seta close to the midline; tergite VIII with 50-60 setae on each side of the midline; pleurae of segments II-VIII with decreasing number of setae, from 8 to 2 small setae each. Spiracles annular, dorso-laterally positioned in a tergopleural groove. Sternites as in Platyrhopalopsis. Male gonotheca (Fig. 26) asymmetrical, not protruding, composed of a medial swollen and multilobed area, partially hidden under sternite VII; female gonothecae (Fig. 27) conical and symmetrical, rounded at the tip. DISCUSSION Paussinae pupae have a nearly complete dorsal pubescence: all paussine species lack dorsal pubesence on the mesonotum and those of tribe Paussini (Platyrhopalopsis and Paussus) also lack pubescence on the head. It seems that the lack of the pupal mesonotal setae is a synapomorphy of the Paussinae and the lack of setae on the head of Paussini, supports the derived position of Paussini within the subfamily. The examined Ozaenini differ from the Paussini in the arrangement of the setae. In Pachyteles and Physea (Ozaenini) the setae of the metanotum and the abdominal tergites 1 to 7 are arranged in a single transverse row on each side of the midline (diagonal on metanotum). In contrast, in Platyrhopalopsis and Paussus (Paussini) the setae form an irregular group on the metanotum, and are arranged in two transverse rows on the abdominal tergites | to 7. The posterior row extends from the lateral border almost to the middle of the tergite. The internal end of the anterior row is situated also close to the midline, but it is much shorter than the posterior one. In Paussus it is on- ly represented by a single internal seta; in Platyrhopalopsis there are about 4 to 5 and they are somewhat irregularly placed. In light of the phylogenetic relationships of the Paussinae based on larval characters (Di Giulio er al., 2003), the complex setation pattern of Platyrhopalopsis and Paussus can be interpreted as a shared derived character. It represents either a synapomorphic character of the Paussini or a more derived clade within this monophyletic tribe. The morphology of pupal spiracles differs among Adephaga taxa. Whereas the mesothoracic spiracles are closed in the pupae of Haliplidae, Amphizoidae, Hygro- biidae and Dytiscidae (Ruhnau, 1986), they are well developed in Carabidae. Seven (presumably functional) abdominal spiracles of similar sizes and shapes are present in the pupae of Carabus and most Hydradephaga according to the illustrations (Sturani, 1962; Bertrand, 1972). However the Ist abdominal spiracle is larger than the following six in Clinidium canaliculatum and Dyscolus megacephalus, and the 7th abdominal spiracle is smaller and more weakly sclerotised than the other spiracles in the Paussinae. The pupal integument of all Paussinae examined have a unique microsculpture. The entire body is covered by a uniform pattern of more or less regularly spaced, small triangular thorn-like sculpticells. In Carabidae, the last abdominal segments are visible and the 9th tergite often contains remnants of the larval urogomphi, as in Carabus and Dyscolus. The shape of pupal urogomphi, as well as their setae, seem to be true remaining larval characters. In PUPAE OF PAUSSINAE 47 contrast to the other Carabidae, the last abdominal segments of Paussinae are inverted and more or less invisible. The gonothecae are almost totally visible in Ozaenini while they are partially inverted and the basal part hidden in Paussini. Another important pupal character in the Paussinae is the extent of the fusion of the abdominal segments. Luna de Carvalho (1951, 1959, 1977) stressed the importance of the number of visible abdominal sternites in paussine pupae. In contrast to four dis- tinguishable sternites in adults, there are 6 sternites visible in the pupae. However, in adult Paussinae the first 3 visible sternites are fused (sternites 2 to 4 — the first one is more or less atrophied in all Adephaga), often with the intersegmental sutures indis- tinct, so that only 4 ventral plates are seen. The sternites are still separate in the pupae, so that the pupae exhibit 6 distinct ventral plates. This condition clearly shows that the fusion of abdominal segments is an apomorphic character state of the adults, while pupae preserve an ancestral condition of six visible abdominal plates typical of Adephaga. Perhaps the most important pupal character with respect to phylogenetic impli- cations within the Paussini is the visible segmentation of the antennothecae of Platyrhopalopsis. This genus is member of the subtribe Paussina, widely considered to be the most derived clade of the Paussini (Darlington, 1950; Nagel, 1997; Di Giulio et al., 2003). The antennae of adults in this subtribe have only three segments: the scape (1st antennomere), the strongly reduced, ring-like pedicel (2nd antennomere), and a club, which is formed by the fusion of the flagellum (Nagel, 1987, 1997). The pattern of transverse constrictions in the adults of some species indicates a fusion of different groups of articles in different species. However the club of adult Platyrhopalopsis is lens-shaped without any trace of articles. In contrast, six or seven articles are apparent in the pupae, as well as a tubercle at the apex of the club, which is absent in the adults, suggesting that a number of 6-7 articles could be the ancestral state for this species. Future studies of the pupae in other Paussina species may reveal more or less antennal segments, which may provide informative characters for inferring relationships within this group. To conclude, this study suggests that pupae may contain characters useful for phylogenetic reconstruction at different levels. Some characters, such as the unique microsculpture, the inversion of the last abdominal segments and perhaps the reduction in the size of the 7th abdominal spiracle, probably represent apomorphies of the Paussinae. The extent of pubescence is interpreted as a morphocline among Adephaga, with an increased reduction in pubesence in derived taxa. The arrangement of setae and the folding of gonothecae differ between basal and derived Paussinae. 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REVUE SUISSE DE ZOOLOGIE 114 (1): 49-57; mars 2007 Chionea austriaca in caves and artificial galleries of Slovenia (Diptera, Limoniidae) Tone NOVAK!, Ignac SIVEC?, Franc JANZEKOVIC! & Erhard CHRISTIAN? l Department of Biology, Faculty of Natural Sciences and Mathematics, University of Maribor, Koroska 160, SI-2000 Maribor. E-mail: tone. novak @uni-mb.si; franc .janzekovic@uni-mb.s1 2 Slovenian Museum of Natural History, PreSernova 20, SI-1000 Ljubljana. E-mail: isivec@pms-lj.si 3 Institute of Zoology, Department of Integrative Biology and Biodiversity Research, University of Natural Resources and Applied Life Sciences, Vienna (BOKU), Gregor Mendel Str. 33, A-1180 Wien. E-mail: erhard.christian@boku.ac.at Chionea austriaca in caves and artificial galleries of Slovenia (Diptera, Limoniidae). - We sampled 54 caves and galleries in north and east Slovenia methodically for ecological data and the invertebrate fauna. The snow fly Chionea austriaca, recorded in 37% of the cavities, proved indif- ferent with regard to most of the abiotic factors, but occurred in significantly higher individual numbers in caves on north and east slopes. The subter- ranean sojourn of Ch. austriaca appears merely weather-mediated. No indications of larval development in the caves were found. Keywords: abiotic factors - speleobiology - distribution - snow fly. INTRODUCTION As their scientific and trivial names imply, snow flies — species of the genus Chionea — are usually met on the snow surface where they can easily be noticed. Nevertheless, these wing-reduced and somewhat spiderlike limoniids spend most of their adult lifetime in leaf litter, under logs and stones, in subnivean habitats or in mammal burrows. The larvae are soil-dwelling. Low catchability makes snow flies appear so rare that Byers (1983), for instance, referred to a catch of 29 specimens as a rich yield. Snow flies have been reported from altitudes up to 2700 m (3500 m in North America: Byers, 1983). Chionea (Sphaeconophilus) belgica (Becker, 1912), and Ch. (S.) lutescens Lundstrom, 1907, occur also in the colline zone and in lowlands (Grootaert, 1984; Reusch, 1988, 1997; Weber & Weidemann, 1993), whereas the bulk of the Chionea species prefer forest sites, or open habitats from the submontane zone upwards (Focarile, 1975). Chionea austriaca (Christian, 1980), originally described from a Lower Austrian cave, is distributed over western Hungary (Christian, unpubl.) and parts of Manuscript accepted 06.10.2006 50 T. NOVAK ET AL. Slovenia and Austria. At the northwestern and western limits of distribution Ch. aus- triaca touches the areas of two closely related species, namely Ch. belgica and the widespread Ch. lutescens (Christian, in press). Ecological notes on European Chionea species focus on aboveground habitats (e.g. Heim de Balsac, 1934; Nadig, 1943, 1949; Bitsch, 1955; Somme & Ostbye, 1969; Hägvar, 1971; Mendl er al., 1977; Itämies & Lindgren, 1985; Heijerman, 1987). Several authors have casually mentioned the sojourn of snow flies in caves (Strobl, 1900; Peyerimhoff, 1906; Bezzi, 1911; Czerny, 1930; Espafiol Coll, 1955, Strinati & Aellen, 1967; Turquin, 1973; Bourne, 1977, 1979; Christian, 1980; Byers, 1983; Novak & Kustor, 1983; Novak, 2005). Information about environmental conditions in cavities inhabited by snow flies, however, is anecdotal at the best. The present paper addresses the abiotic conditions at sampling sites of Ch. aus- triaca in caves and artificial galleries in Slovenia. We want to test our hypothesis that this species is only loosely aligned to macrocaverns. MATERIALS AND METHODS During the period 1977-2001, 54 caves and artificial galleries (in the following: caves) in central and northern Slovenia (mapped in Novak, 2005) were ecologically investigated in January, April, July and October according to a standardized scheme. All accessible surfaces of the cave passages like walls, ceiling, speleothems, micro- habitats below stones and wood, and water surfaces, were carefully inspected for terrestrial animals. Pitfall traps were set, on average, every 3 m starting at the entrance, and exposed for 47+1 hours. Each trap was baited with 3 g of decomposing meat attached on a wire above the trapping liquid (2 cm of fruit juice with cherry and maraschino essences, and a lacing of detergent). Substrate from the close vicinity of the trap sites was sampled for Tullgren extraction. Environmental parameters were determined chiefly according to Stewart et al. (1974) and Rowell (1997). The distance from a trap to entrance and surface, respec- tively, and the cross-section of the passage at the trap site were taken from cave maps (Novak, 2005). Illumination was measured using a Lunasix luxmeter. Substrate pH was determined with a combined electrode (91-02 Orion, USA; Orion Ionalyser 407 A) in elutriated fresh samples after 2-4 hrs stirring in an equal volume of distilled water. Substrate carbonate and organic matter contents were established for the dried and pul- verized <2 mm fraction; carbonate was quantified by Scheibler calcimetry, ignition loss after heating at 700°C for 1 h. Ground and air temperatures were measured with decimal thermometers (Intos, Croatia). Air humidity was determined with a psychro- meter according to August, ground moisture by drying at 105°C to constant weight. Airflow was detected with a hand-held anemometer (Munro IM159) or, when below 0.3 m-s"!, derived from the velocity of fog or magnesium smoke (cf. Andrieux, 1970). Cold hardiness of one male, collected on a south-facing meadow slope at Studence (46°18°53” N, 15°09’51” E, 330 m) on snow, 31.01.2006, was determined in a thermostat chamber THK/V 1-2020 (Elelstro, Slovenia). The specimen was enclosed in a glass tube with a scrap of wet paper and cooled down to lethal temperature in 12 hour treats in steps of 1°C/day, with intermediate 12 hour breaks at acclimatization temperature of +8°C. CHIONEA AUSTRIACA IN CAVES OF SLOVENIA 5] Voucher specimens of Ch. austriaca are deposited in the Slovenian Museum of Natural History (Ljubljana), in the Muséum d’histoire naturelle de la Ville de Genève, and in the Natural History Museum, Vienna. RESULTS Three of 14 European Chionea species (Oosterbroek, 2005) have been record- ed in Slovenia (Oosterbroek & Simova-Tosic, 2003, and own data; Fig. 1). Ch. (Chionea) araneoides Dalman, 1816 was found in the Krma valley (UTM: VM13); Ch. (Sphaeconophilus) alpina Bezzi, 1908 is known from Mt. Grmada (VM61, 640 m), d2r L ! Fa | Y Tu] | | i | Ta] N È T X i Use Avy e Bere | | el | 1e A| | | lei a | | | Tree | | i r | L [72 | ERA |e JE ep al | | np _|6 | | = | = | n Conta © Me JL R Se + ehe lt = “| = | È ® LIL = vale | À | + tac I M ee ee @ je \\| N, Eee A | Eee el ale (saith É @ | eo. ©. = ES ie |e ii Te =] Fic. 1 Records of snow flies in Slovenia: A Chionea araneoides, O Ch. alpina and @ Ch. austriaca. Zdenska jama cave (VL77, 460 m), Dobrepolje (VL77, 450 m), Maëkovica cave (VL47, 480 m) (Simova-Toëié & Sivec, 1977), and the alpine ice cave Ivaliceva jama in Mt. Kredarica at 2435 m a.s.l. (Novak & KuStor, 1983); the third species, Ch. (S.) austriaca was encountered in numerous caves in central and eastern Slovenia (Table 1). Here we add three snow-surface records of Ch. austriaca: Mt. Jesenik (WM13, 688 m), Studence (WM13, 330 m) and Mt. SneZnik (VL54, 1690 m, currently the southernmost locality), and a further cave record: Brezno pod Durcami in Mt. Raduha (VM84, 1840 m, currently the highest record in Slovenia), which was not included in the systematic investigations. In 20 (37%) of the 54 caves, 38 specimens (11 males, 27 females; sex ratio 29:71) of Ch. austriaca were met, at altitudes between 270 and 1515 m (Tab. 1). Tab. 2 compiles values of abiotic factors at effective trapping sites. All specimens were trapped in pitfalls exposed in January. Larvae were neither seen, nor trapped or Tullgren-extracted. Most adults occurred in the entrance zone of the caves during the winter (Fig. 2). Caves opening in northern and eastern slopes were significantly more T. NOVAK ET AL. 52 6LIOLI ‘dI s OLT «Er. SEoST ‘LE. 60097 ITINM rn 6LIOLI ‘57° PI I 00r «LT LEST '«IS.Ll9 7rINN TST 8L1007 ‘LI u O£L «EL PEST '«0E.8TI rN LOPT 8L6TTOSI ‘dI I CIS «TE OST ‘«6£.90097 OFM #8 8L6IIO8I “LI I 00S «SS. ETSI OF. 80097 = TEINM TE 6LIO8I ‘ét ‘LI I 08r SE SCI Icon ITINM LIGE 8LIOII ‘LI Yo «6. 91oST «95.89 YTINM 9¢9b 8LIO6I ‘37 I Crs «VO9TST '«SE.LEIF = ITINM 6181 6L1060° 57° PT I OTE “IR CGO? IINM SES 6L1060 ‘dI I 007 GATES LIO ZAR GIAN ZISE 8LIOSI ‘LI I 019 «ITEIoST “«69.T7E097 SINM 807€ 8LIOSI ‘37 Ò 087 «9S. LOST «Lr.8E97 = ITINM an 8LIOLO ‘dI I OLE «LT:100SI '«E0.7I9 = EOINM 6S6€ SLIOZI ‘LI 9 958 «PT EDS ‘«v 6.997 POW 881€ SLIOIL SI I Oss «TT EST "«HE.0E9IF SOINM id 6LIOIZT SE O1 I 159 «CO. PSP I “CO. L0097 O6INA LOVE 6LIOSO ‘dI p 0r6 «OSSI “SV. LTOOY IA AVA ISHE 6LIOZI ‘dI d 007 «EE SSoPT ‘ES. TE9P S6WA EITE 6LIOOI "Er ‘LI i Sig «VIL Ooh “81.81.90 ZLWA OLE 6L1090 ‘31 I 06S «PL EPP ‘LT. ETOP ELINA COP ayeq ‘pul = pag AV (4 ‘N) SaJeuIp10-07) NILO PBI AO IWIPIIUOPN}S peu NUIO[JOULUEX A BUE IDIUE]A eu nyooeNn Lid eluyn] ewel elijo) woyng ud nwojouwey A BUE ( 7 eluyn] eysaojueg eluyn eAopyef Iw99d IWTAOYI9H pod ewer npzos woadliajpeyg A vurel PAOYIHAIO © ewrel vaagisedg 1904 aygode7 nuge13 WONYSLISIG A AOI IUJOUUN) vuuel PASIAONOT vurel eyQopy nipouz, 11d yyalqo ewel vaoslepig BOIPJOA BADLIIGOYS 90d IPO, Hd vuref IQUO HAOQ A eyyeltz joyod DIVA AIS 10 ABD [APP] UONDIT[OO JO aJep pur SJENPIAIPUI JO Jaquinu = 972 ‘puy ‘(onqeuo) = 1 ‘sysTyOS = s ‘ayNeLUdd = d ‘o[q JCW = WI ‘AJUO[OP = P ‘SUOJSOLUT] = | “AJPIOWO[BUO9 = 9 :YOOIPoq = pagg (Lu) Spmnpe = J[Y ‘(| ‘SIA 99S) 9P09 PLLA ‘SOHOI[PS [RIO Ie Syeaıpur SIONI] eyrde 7 ‘19}S1391 SAVI SUDAOTS dU} ur JaquINU [esepe9 = PLD ‘5007 "NEAON UI Se Joquunu JUILINI = ON ‘VIUDAOTS JO SOABO UI DIDIAISND DaUOIYD : | AIAVL CHIONEA AUSTRIACA IN CAVES OF SLOVENIA 53 Chionea austriaca Number of individuals Distance from the entrance (m) FIG. 2 Yield of Ch. austriaca individuals relating to the distance between cave entrance and trap locality. frequently settled by Ch. austriaca as compared to other slopes (x? = 24.54; P < 0.001), and hosted significantly more individuals (X? = 17.05; P = 0.017) (Tab. 3). Preference for any kind of bedrock was not observed. In two cases (26 and 38 in Tab. 1) entrances were situated in various types of forests. By comparing the meteorological conditions in caves with vs. those without Ch. austriaca we found that in “Chionea caves” the air and ground temperatures were sig- nificantly lower, and airflow significantly higher. There were no significant differences regarding relative air humidity and substrate moisture (Tab. 4). Males and females did not differ significantly with respect to any abiotic parameter at the trapping sites (1,37 = 0.000-2.614; P = 0.11-0.98). In the cold hardiness test the single male of Ch. austriaca died at -7°C. DISCUSSION Adult Ch. austriaca were trapped in caves only during the cold season, in agree- ment with the period of their occurrence on snow. The temperature demands of Ch. austriaca are comparable to those of most other European snow flies (Wojtusiak, 1950; Bourne, 1979), and its supercooling ability — admittedly determined for one specimen only — seems to agree with the values reported for Ch. araneoides (mean supercooling point -7.5°C, range -10.5 /-4.5°C: Sömme & Östbye, 1969). A vast ecological range of Ch. austriaca is indicated by the large altitudinal range, indifference to bedrock, its occurrence in forested as well as in non-forested areas, and inside caves by tolerance 54 T. NOVAK ET AL. TABLE 2: January values of abiotic factors at effective trapping sites of Ch. austriaca, compared to the extreme values in “Chionea caves” and to the mean and extreme values of all the 54 caves (based on 605-617 sampling sites). Mean + s.d. (min-max) Min-Max in Mean + s.d. (min-max) at effective trapping sites ‘Chionea caves” in all the 54 caves Distance from entrance [m] 8+10 (0-49) 0-121 31+31 (0-121) Distance from surface [m] 9+8 (1-40) 0-70 18+14 (0-80) Passage cross-section [m2] 5+7 (0736) 0.2-80 13+31 (0.2-460) Illumination [lx] 258+169 (0--10,000) 0--100,000 194+171 (0--100,000) pH 7.9+0.8 (4.3-8.6) 4.3-8.7 8.1+0.6 (2.4-8.9) CO; [%] 21.4+24.0 (0.0-76.0) 0-96.6 31.9+29.8 (0.0-98.7) Ignition loss [%] 13.2+15.2 (0.3-52.9) 0.3-85.9 10.7+13.2 (0.3-85.9) Air temperature [°C] 1.2+3.8 (-5.0-8.9) -5.0-11.5 3.6+4.4 (-6.0-11.5) Ground temperature [°C] 1.6+3.3 (-3.0-9.0) -4.6-12.6 4.1+4.2 (-7.2-12.6) Relative air humidity [%] 91+12 (65-100) 54-100 95+9 (54-100) Substrate moisture [%] 18.4+12.0 (0.2-46.7) 0.0-73.6 20.8+14.7 (0.0-87.6) Airflow [m/s] 0.07+0.04 (0.00-2.00) 0.00-10.00 0.11+0.73 (0.00-10.00) Soil colour / modus (min-max) 10YR 5/4 (SY 3/2-7.5YR 5/6) 5YR 5/8-5Y 5/2 10YR4/4 (SYR 3/2-2.5Y 6/4) TABLE 3: Distribution of Ch. austriaca with respect to slope orientation of cave entrances. Slope Ch. austriaca Caves with All caves orientation individuals N (%) Ch. austriaca N (%) N (%) N 12 (31.6) 4 (20.0) 10 (18.5) NE 2103) 2 (10.0) 3 (5.6) E 12 (31.6) 6 (30.0) 14 (25.9) SE 3.132) 3 (15.0) 4 (7.4) S 4 (10.5) 4 (20.0) 7 (13.0) SW 0 (0.0) 0 (0.0) 4 (7.4) W 0 (0.0) 0 (0.0) 10 (18.5) NW SM) 16:0) ACH) total 38 (100) 20 (100) 54 (100) TABLE 4: Selected meteorological parameters of caves with Ch. austriaca present vs. absent. January values from all trap sites along the first 10 m (i.e., those without Chionea yield includ- ed). Significant differences asterisked *. Ch. austriaca F1, 260 JP present (N = 103) absent (N = 159) Air temperature 4.8+2.4 (0.6-9.5) 6.2+2.0 (0.3-11.4) 24.59 <0.001* Ground temperature 4.7+2.5 (-0.3-11.3) 5.7+2.0 (0.5-11.0) 12.78 <0.001* Air humidity 94+8 (69-100) 92+22 (43-100) 252 0.113 Substrate moisture 23.3+16.1 0.0-65.4 24.8+17.0 (1.1-79.2) 0.52 0.473 Airflow 0.08+0.05 (0.0-10.00) 0.06+0.04 (0.0-5.00) 4.97 0.026* of wide ranges of illumination, substrate pH, carbonate contents and ignition loss. The conspicuous prevalence of individuals in caves on north and east facing slopes contrasts with the impression that snow flies in open-air habitats do not display any slope orientation preference. Their surface activity, however, is strongly weather- CHIONEA AUSTRIACA IN CAVES OF SLOVENIA 55 dependent inasmuch heavy frost and wind drive them into more moderate hideaways. So the disproportionately high individual numbers of Ch. austriaca in caves opening out on north and east slopes is not a consequence of psychrophily, but rather the result of more frequent chill avoidance migrations there. The highest trapping yield of snow flies in the entrance zone of the caves and in passages of small cross-section supports this conclusion. Lower ground temperature and higher airflow in “Chionea caves” may be merely the consequence of the harsh climate conditions on north and east slopes. Wing reduction of snow flies has been explained as a result of regressive evo- lution due to temporary exposure to cold and windy weather (Hackman, 1964; Byers, 1961, 1983). This morphological character could lead to consider snow flies as being pre-disposed to life in subterranean habitats (cf. Vandel, 1964; Ginet & Decou, 1982). However, Ch. austriaca — like several other European congeners — inhabits macro- caverns facultatively and only temporaryly during its epigean ecophase in the cold sea- son. The fact that maximum density in caves is observed in January, when virtually all females are ready for egg-laying (Christian unpubl.), does not necessarily indicate sub- terranean oviposition. On the one hand there is no report on larval development of Chionea in caves, and on the other hand an extremely long oviposition period (from November to March) was shown for Ch. araneoides (Hägvar, 1976), so that females of Ch. austriaca — let’s assume an adult life span of two months (cf. Byers, 1983) — may lay eggs after they have quitted the cave shelter. Thus, in terms of the classical biospeleological classification (Vandel, 1964), Ch. austriaca is best included in the trogloxenes. ACKNOWLEDGEMENTS We thank Valika Kuëtor, Tomaz Proëek and Mojmir Stangelj for help with the field investigations, DuSan Devetak (Dept. Biol. Univ. Maribor) and the anonymous re- viewers for insightful remarks on the manuscript, and Michelle Gadpaille (Dept. Ang. Amer. Univ. Maribor) for linguistic improvements. The study was partly supported by the Slovene Ministry of High Education, Science and Technology within the Biodiver- sity research programme (P1-0078). REFERENCES ALLEN, S. E., GRIMSHAW, H. M., PARKINSON, J. A. & QUARMBY, C. 1974. Chemical analysis of ecological materials. Blackwell Scientific Publications, Oxford, 565 pp. ANDRIEUX, C. 1970. Contribution a l’etude du climat des cavités naturelles des massifs karstiques. Annales de Spéléologie 25: 441-559. BEZZI, M. 1911. Diptères (première série), suivi d’un Appendice sur les Diptères cavernicoles recueillis par le Dr Absolon dans les Balkans. Biospeologica, 20. Archives de Zoologie Expérimentale et Générale 8(5): 1-87. Bitscu, J. 1955. Le genre Chionea (Diptera, Tipulidae). Ecologie, systématique et morphologie. 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SOMME, L. & OstTBYE, E. 1969. Cold-hardiness in some winter active insects. Norsk Entomo- logisk Tidsskrift 16: 45-48. STRINATI, P. & AELLEN, V. 1967. Faune actuelle de deux grottes suisses de haute altitude. Rassegna Speleologica Italiana 19: 84-87. STROBL, G. 1900. Dipterenfauna von Bosnien, Hercegovina und Dalmatien. Wissenschaftliche Mittheilungen aus Bosnien und der Hercegowina, Sarajevo, 7: 552-670. TURQUIN, M.-J. 1973. La colonisation de quelques grottes du Jura par Niphadobata alpina Bezzi (Dipt. Tipulidae). International Journal of Speleology 5: 21-28. VANDEL, A. 1964. Biospéologie. Gauthier-Villars, Paris, 619 pp. WEBER, G. & WEIDEMANN, G. 1993. Niphadobata lutescens (Lundström, 1908) and N. belgica (Becker, 1912) from the Solling Mountains, Lower Saxony, FRG (Diptera, Limoniidae). Deutsche Entomologische Zeitschrift 40: 259-263. WOIJTUSIAK, H. 1950. The temperature preferendum of winter insects of the genus Boreus (Panorpatae) and Chionea (Diptera). Bulletin international, Polska Akademia Umiejetnosci, Wydziat matematyczno-przyrodniczy, Série B, 11: 125-143. u 2 LL Tr ri CRAN ue REVUE SUISSE DE ZOOLOGIE 114 (1): 59-64; mars 2007 A new genus and species of the spider family Agelenidae from western Sichuan Province, China (Arachnida: Araneae) Xiang XU!:2 & Shugiang LI!-3 ! Institute of Zoology, Chinese Academy of Sciences, Beijing 100080, China. 2 College of Life Science, Hunan Normal University, Changsha 410081, Hunan, China. 3 Corresponding author. E-mail: lisq@ioz.ac.cn A new genus and species of the spider family Agelenidae from western Sichuan Province, China (Arachnida: Araneae). - Ageleradix sichua- nensis, a new genus and new species, is reported based on specimens from western Sichuan, China. Ageleradix gen. n. is distinguished from the other genera of Agelenidae by having a special sclerite between tegulum and embolus in males, a well-developed posterior plate in the epigynum and a broad copulatory bursa in females. Two species are included in the new genus, the type species and Ageleradix cymbiforma (Wang, 1991) comb. n. Keywords: Taxonomy - Ageleradix - Agelescape - new combination. INTRODUCTION Sichuan Province is located in the Upper Yangtze Valley in the southwest part of China. It covers an area of 569,000 square kilometres and has an estimated population of over 70 million people. Lying along the transition zone between the Sichuan Basin and the Qinghai-Tibetan Plateau, western Sichuan has an unusual geography and weather conditions, which have given rise to a variety of landforms and habitats. A collection trip to western Sichuan in summer 2004 yielded a very unusual new agelenid species, which is here assigned to the new genus Ageleradix gen. n. A detailed description of this genus and its type species, Ageleradix sichuanensis sp. n., is given in the following. MATERIAL AND METHODS Measurements are given in millimetres. Eye diameters were measured at the widest part. Leg measurements are given as: Total length (femur, patella, tibia, metatarsus, tarsus). Chaetotaxy follows Thaler & Knoflach (1998): v’, v’’ position pro-, retroventral, 1’, 1’’ pro-, retrolateral, d dorsal; subscript a and 2 denote apical position and number of spines, respectively; pairs of spines are given in parentheses, Manuscript accepted 21.11.2006 60 X. XU & S.LI whorls of spines in square brackets. Terms used in the text and figure legends mainly follow Levy (1996) and Wang (1997). Abbreviations used in text and figures are: ALE - anterior lateral eye; ALS - anterior lateral spinneret; AME — anterior median eye; C — conductor; CB — copulatory bursa; D — diverticula; E — embolus; FD — fertilization duct; MA — median apophysis; MOQ — median ocular quadrangle; PA — patellar apophysis; PLE — posterior lateral eye; PLS — posterior lateral spinneret; PME — posterior median eye; PMS — posterior median spinneret; PP — posterior plate of epigynum; R — radix; RTA - retrolateral tibial apophysis; S — spermatheca; SC — scape; TP — tegular process. All type specimens are deposited in the Institute of Zoology, Chinese Aca- demy of Sciences, Beijing (IZCAS) and in the Muséum d’histoire naturelle, Genéve, Switzerland (MHNG). TAXONOMY Ageleradix gen. n. TYPE SPECIES: Ageleradix sichuanensis sp. n. SPECIES INCLUDED: Two species are here placed in this new genus: Ageleradix sichuanensis sp. n. and Ageleradix cymbiforma (Wang, 1991) comb. n. which is transferred from Agelena. ETYMOLOGY: The generic name is derived from the words “Agelena” and “radix”; the gender is feminine. DIAGNOSIS: In its general appearance (Fig. 1), this new genus resembles two other Palaearctic genera, Agelena and Agelescape, which construct funnel webs among bushes and in grass. In the shape of their epigynum, Ageleradix gen. n. females are similar to those of Agelescape by having a scape (Fig. 10), but can be distinguished by having a well-developed posterior plate in their epigynum and a broad copulatory bursa (Figs 10-11). Males can be easily distinguished from those of Agelena and Agelescape by having a radix between tegulum and embolus, by their basal median apophysis and by a strong and broad conductor (Figs 3-7). The new genus is distinct from Agelena by females having a scape extending from the anterior rim of the epigynum (Fig. 10). DESCRIPTION: Body shape, coloration and eye arrangement as in Agelena and Agelescape (Fig. 1); body covered with feather-like hairs (Fig. 2); ALE biggest, AME smallest; cheliceral groove with 3 promarginal and 3 retromarginal teeth; epigynum with a plate extending from posterior rim (Fig. 10); spermathecae widely separated from each other; copulatory bursae well-developed (Fig. 11); male palp with small retrolateral tibial apophysis and with indistinct patellar apophysis; tegulum with process; conductor very strong, slightly modified into a shallow concavity housing the distal end of the embolus; embolus originating centrally, with broad base and pointed distal end; radix developed as a partly serrated additional sclerite between tegulum and embolus; median apophysis situated basally on the palpal organ (Figs 3-9). DISTRIBUTION: Sichuan, China. AGELERADIX SICHUANENSIS FROM SICHUAN 61 Ageleradix sichuanensis sp. n. Figs 1-11 HoLoTyPE: S(IZCAS), Jintang Town, Kangding County (30.0° N, 101.9° E), Sichuan Province, China, July 19, 2004. PARATYPES: 13 (IZCAS), Yutong Town, Kangding County (30.0° N, 101.9° E), Sichuan Province, China, August 4, 1981.— 62 and 6d(MHNG), Jintang Town, Kangding County, July 18-19, 2004. — 16 (IZCAS), Changpinggou Valley, Mt Siguniang (31.1° N, 102.9° E), Xiaojin County, August 1, 2004.— 29 and 43 (IZCAS), Waba Village, Geshizha Town, Danba County (30.8° N, 101.9° E), July 30, 2004. — 19 and 26(IZCAS), Meirengu Town, Danba County, July 31, 2004. — 1? (IZCAS), Daxue Village, Geshizha Town, Danba County (30.8° N, 101.9° E), July 30, 2004. — 1 4 (IZCAS), Geshizha Town, Danba County, July 30, 2004. — SS (IZCAS), Kungyu Town (30.5° N, 102.0° E), Danba County, July 30, 2004. All type material was collected in the Sichuan Province by Dr Xiang Xu, Mr Zhengtian Zhang, Mr Yucheng Lin, and Ms Qian Wang. ETYMoLoGY: The specific name derives from the name of the province in which the types were collected. DIiaGNOSIS: This new species can be distinguished from A. cymbiforma by having a scape extending from the anterior rim of epigynum and globular sperma- thecae in females, and by the evidently different shape of the conductor and the presence of a radix in males. DESCRIPTION OF MALE: Total length 7.64-9.84. Holotype 9.27 long; carapace length 4.00, width 2.91; abdomen length 5.27, width 2.73. Two eye rows strongly procurved; in dorsal view AME in the middle of an open circle formed by the other six eyes. AME 0.12, ALE 0.19, PME 0.15, PLE 0.16; AME-AME 0.05, AME-ALE 0.07, PME-PME 0.10, PME-PLE 0.08; MOQ length 0.36, front width 0.29, back width 0.40; clypeus height 0.17. Body covered with tiny white or grey featherlike hairs (Fig. 2). Carapace brown; head region clearly narrower than thorax region in dorsal view (Fig. 1). Fovea long, longitudinal. Chelicerae red-brown, promargin and retromargin each with 3 teeth on groove. Labium wider than long. Sternum yellow- brown, its distal end slightly inserted into coxa IV. Legs thin and long. Tibiae I-IV with two rows of dorsal trichobothria; metatarsi I-IV and tarsi I-IV with single row of trichobothria dorsally. Chaetotaxy (slightly variable): Femur Id [°° (d 1°) (’ 1°’); III ULIVI div) sPatellal-IVed 14 dy. Mibia Ivy; Nena VV) EE) ile VE d'A) VA INN IST) eV) Metatarsus (VO) Ie (ve) I TIR vv) [dd ZA Ne) ve) AE Te IN Km BZ) IN Vous bar ba. |: Larsus I-11E 07 IV 1 Pavel Les TVilonsest; leg formula: IV, I, II, III. Leg measurements: I: 16.64 (4.00+5.18+4.91+2.55); II: 15.73 (3.91+4.73+4.73+2.36); II: 15.28 (3.82+4.55+4.64+2.27); IV: 20.10 (4.82+5.82 +6.55+2.91). Abdomen much longer than wide, sides darker than middle of dorsum (Fig. 1). ALS 0.62 long; basal article of PLS 0.52 long, apical article 0.62 long: AMS, PMS and PLS with conspicuous gland spigots. Male palp with indistinct patellar apophysis; conductor strong, broad, and slightly modified into a shallow concavity; embolus originating centrally, with broad base and pointed distal end; radix partly covered with denticles; tegulum with a cone-shaped process; median apophysis palmate (Figs 3-9). 62 X. XU & S.LI Fic. 1 Ageleradix sichuanensis sp.n., male from Sichuan, China. DESCRIPTION OF FEMALE: Colour and leg formula as in male. Measurements very variable. Total length 7.64-8.73. A specimen from Jintang Town, Kangding County 9.27 long; carapace length 3.64, width 2.55; abdomen length 5.09, width 3.14. AME 0.13, ALE 0.18, PME 0.14, PLE 0.16; AME-AME 0.05, AME-ALE 0.07 PME- PME 0.12, PME-PLE 0.10; MOQ length 0.39, front width 0.26, back width 0.36; clypeus heighth 0.13. Chaetotaxy (slightly variable): Femur I d (d 1’) (1° 1°’); I-IV dd (77°. Patella I-IV did, Tibia TW v”) 1” Y Ww v’) D) Rd IVA (div) (1212) (ed) (2122) Va . Metatarsus I/II (v’ v’°) (WW v’’) U’ vv]; HI [P View Vv [PI dive lh ido V1? vv2;WV [dP Po vv iv TAC OIL Vv’ v’’]. Leg measurements: I: 11.10 (3.00+3.73+2.55+1.82); II: 10.36 (2.91+3.36 +2.45+1.64); III: 10.28 (2.82+3.18+2.73+1.55); IV: 14.18 (3.64+4.27+4.18+2.09). ALS 0.60 long; basal article of PLS 0.45 long, apical article 0.65 long. Epigynal atrium large. Long scape extending over most part of atrium; distal end of scape slightly bifid and almost in contact with triangular plate extending from posterior rim of epigynum. Broad, membranous copulatory bursae bearing a pair of threadlike diverticula; diverticula with a slightly swollen part; copulatory bursae connected internally to a pair of globular spermathecae, to the outside through an atrium; fertilization duct distinct (Figs 10-11). DISTRIBUTION: Only known from the type locality. AGELERADIX SICHUANENSIS FROM SICHUAN 63 Fics 2-11 Ageleradix sichuanensis sp. n. (2) Featherlike hair. (3) Left male palp, prolateral view. (4) Same as in 3, ventral view. (5) Same, retrolateral view. (6) Conductor (separated from palpal organ), prolateral view. (7) Same, ventral view. (8) Embolus (separated from palpal organ), prolateral view. (9) Same, ventral view. (10) Epigynum, ventral view. (11) Same, dorsal view. Scale bars: 2, 6-11= 0.1 mm; 3-5= 0.5 mm. 64 X. XU & S. LI ACKNOWLEDGEMENTS The manuscript benefited greatly from comments by Dr Peter J. Schwendinger (MHNG), Dr Yuri M. Marusik (Institute for Biological Problems of the North RAS, Russia) and Dr Xinping Wang (University of Florida, USA). This study was supported by the National Natural Sciences Foundation of China (NSFC-30670239, 30370263, 30310464, 30470213, 30499341), by the National Science Fund for Fostering Talents in Basic Research (NSFC-J0030092), by the Knowledge Innovation Program of Chinese Academy of Sciences (KSCX2-YW-Z-008), by the Beijing Natural Science Foundation (6052017) and partly also by the Kadoorie Farm and Botanic Garden, Hong Kong Special Administrative Region, China. REFERENCES Levy, G. 1996. The agelenid funnel-weaver family and the spider genus Cedicus in Israel (Araneae: Agelenidae and Cybaeidae). Zoologica Scripta 25(2): 85-122. THALER, K. & KNOFLACH, B. 1998. Phyxelida anatolica Griswold, new to Cyprus (Arachnida: Araneae: Amaurobiidae: Phyxelidinae). Bulletin of the British Arachnological Society 11(1): 36-40. WANG, J. F. 1991. Six new species of the genus Agelena from China (Araneae: Agelenidae). Acta zootaxonomica sinica 16(4): 407-416. WANG, X. P. 1997. On three Agelena species from China (Araneae, Agelenidae). Bulletin of the British Arachnological Society 10(7): 253-256. REVUE SUISSE DE ZOOLOGIE 114 (1): 65-86; mars 2007 Continued massive invasion of Mysidae in the Rhine and Danube river systems, with first records of the order Mysidacea (Crustacea: Malacostraca: Peracarida) for Switzerland Karl J. WITTMANN Department of Ecotoxicology, Center of Public Health, Medical University of Vienna, Währinger Strasse 10, A-1090 Wien, Austria. E-mail: karl.wittmann@meduniwien.ac.at Continued massive invasion of Mysidae in the Rhine and Danube river systems, with first records of the order Mysidacea (Crustacea: Malacostraca: Peracarida) for Switzerland.- A survey of invasive species belonging to the Mysidae in the Rhine and Danube Rivers and adjacent freshwater systems revealed that certain species have invaded the waters of many more countries than previously known. First records of Hemimysis anomala G. O. Sars, 1907 are given for Switzerland, Slovakia, Hungary, Croatia and Serbia. The same holds true for Katamysis warpachowskyi G. O. Sars, 1893 in Croatia and Serbia, and for Limnomysis benedeni Czerniavsky, 1882 in Switzerland. New or first records are also given for L. benedeni in the systems of artificial inland waterways of northern Germany, France and Serbia. L. benedeni and H. anomala are the first representatives of the order Mysidacea (or Mysida according to certain authors) in Switzerland. A single specimen of Paramysis lacustris (Czerniavsky, 1882) serves as a first record for the upper Danube in Austria. The observed spreading of L. benedeni in inland waterways of France suggests that it probably will soon arrive at the Mediterranean coast, with unknown conse- quences for indigenous populations of the closely related species in the genus Diamysis Czerniavsky, 1882. H. anomala, a top invader of conti- nental waters in Europe, could become a potential threat at the ecosystem level if its invasion continues into large lakes, such as those connected with the Rhine system in Switzerland. Keywords: Hemimysis anomala - Limnomysis benedeni - Katamysis warpachowskyi - Paramysis lacustris - invasive aquatic species - Ponto- Caspian invaders - continental Europe - distribution - anthropogenic disper- sion - ecological impact. INTRODUCTION Up to the mid-20!h century, only few range extensions were recorded for mysids in continental waters of Europe. Behning (1938), for example, noticed that Paramysis ullskyi Czerniavsky, 1882 penetrated only a few hundred kilometres into the tributaries of the Black Sea, whereas more than 3000 km into the Volga River and its tributaries. He explained the strong differences in distribution, already observed in the 1920s, by Manuscript accepted 8.12.2006 66 K. J. WITTMANN the more developed inland navigation along the Volga; such navigation may have favoured the dispersal of this mysid and of other peracarids attached to the fouling on ship hulls. Only few later authors adopted this view, which has recently been recon- sidered by Tarasov (1996). The next major expansion event was noted in 1946 with the surprising appearance of Limnomysis benedeni Czerniavsky, 1882 in the winter har- bour of Budapest, almost 1200 km upstream of the previously known distribution limit in the Danube River (Dudich, 1947; Woynarovich, 1955). Such a strong yet rapid dis- placement of the known limit suggested anthropogenic mechanisms of dispersion. Bacescu (1966) and Mordukhai-Boltovskoi (1979) already pondered why Limnomysis dispersed to the upper reach of the Danube, while it inhabited only the lower reach of the Volga River and only a near-mouth confluent in the Don River system. After having been projected already in the 1930s, the next major change started in the late 1940s with the intentional introduction of mysids into hydropower reservoirs, lakes, and other water bodies in order to enrich the supply of food for fish. From 1948-1965 more than one hundred million mysids, together with a great variety of other invertebrates, were intentionally released into more than 200 water bodies of the former Soviet Union. Our knowledge about these introductions and their outcomes is quite obscure. This is mainly due to the inaccessibility of cryptic literature (citations in Komarova, 1991; Grigorovich et al., 2002) and inadvertent stocking (Pligin & Yemel’yanova, 1989). During the next decades some of the mysid species showed local secondary spreading from the introduction sites; a few species — essentially the four presented in Fig. 1 — transgressed the large continental watersheds and invaded vast areas of central, northern and western Europe. In the early 1980s it became clear that mysid introduc- tions could have adverse effects at the ecosystem level (Rieman & Falter, 1981; Fiirst et al., 1984; Ketelaars et al., 1999), and most of the intentional introductions were stopped. This had no direct effect on the ongoing non-intentional mechanisms of dis- persion, which culminated in 1997-1998 with the explosive range expansion of Hemimysis anomala G. O. Sars, 1907 and L. benedeni in waters of the Rhine system. The present contribution indicates further range expansions of these and additional species in large areas of continental Europe, for the first time also including Switzerland. A critical analysis of invasion histories could help to estimate the potential for future dispersal and possible threats to ecosystems and biodiversity. MATERIAL AND METHODS By convention, the river kilometres are numbered in downstream order for the Rhine and its tributaries, but in upstream order for the Danube. Measurements of abiotic parameters were taken as explained in Wittmann (2002). Salinity was calcu- lated from conductivity measurements corrected to 25°C, as a dimensionless equi- valent of overall ionic content rather than content of sea salts. Mass occurrence was defined as >1000 specimens and, as an additional prerequisite, >10 times the average yield of positive samples taken with the same method under comparable conditions. The diverse sampling methods follow those outlined in Wittmann (2002), with the exception of the following modifications introduced in 2004: the hand nets were optionally attached to a 4.6 m telescopic rod, allowing for sampling from the shore MYSID INVASION IN THE RHINE AND DANUBE 67 down to 3.5 m depth; the drift net was larger (opening 2872 cm?, mesh size 500 um for the anterior 150 cm of length, and 280 um for the posterior 100 cm) than before and kept in position 10 cm above the substrate by a steel cage. Ten bottle traps were used in 2005, designed essentially according to Odenwald er al. (2005); the main difference was that the bottles were tightly attached to 0.5 kg weights in order to throw them, filled with about 1.5 1 of water and the bait, several metres from the shore. The bait was commercial tablets for aquarist use («Wels Tabs mit Spirulina» by ‘Fa. Product Aquaristik-Terraristik’ in Austria, imported from ‘Hicari Corp.’ in Japan), composed of dried, compressed algae, insects and crustaceans. The yield obtained with the baited traps confirmed the findings of Odenwald et al. (2005) that Hemimysis anomala is more effectively trapped than other mysid species (Limnomysis benedeni and Katamysis warpachowskyi G. O. Sars, 1893). The drift net was useful for currents >0.2 m/s, especially if no or only few stagnant side arms were accessible in a particular river stretch. Drift samples yielded all three above- mentioned species more often than the remaining methods. Drift nets and bottle traps were set up over night. In 1985-2001 a total of 390 samples containing mysids were taken along the Rhine and Danube Rivers, from the upper reaches down to the deltas, as well as along the coasts of Romania and western Turkey (Black, Marmora, Aegean Seas) and in adjacent freshwaters. Part of these sampling data is available in Wittmann (1995, 2002), Wittmann et al. (1999) and Wittmann & Ariani (2000). In 2002-2005 a total of 230 samples were taken at 117 stations in the Rhine and Danube systems and in adjacent waters. Additional data were obtained from hand net samples taken by B. Csänyi during the Cousteau-Expedition 1991 at 50 stations between Danube-km 2786 and km 80. Twelve of these samples contained mysid species. Two samples with L. benedeni were studied in the collection of the Biologija Univerza Ljubljana: from the Tisa River in Serbia (26 June 1977, leg. I. B. Sket) and from the Danube at Donji Milanovac in the Carpathian breakthrough (8 Oct. 1984). Ten samples with L. benedeni and Diamysis pengoi (Czerniavsky, 1882), taken on 21 Aug.-30 Sept. 1924 by V. Pietschmann and O. Koller at Lake Patiu (46.20°N, 28.0167°E, at the River Prut, an affluent of the Danube at km 133) and at six stations from Danube-km 181 down to km 19 of the Chilia branch, were studied in the collection of the Naturhistorisches Museum Wien (Crust. Coll. nos. 5960, 5962, 13397-13408). In 1985-2004 most of the material was fixed in a solution of 4% formalin in ambient water and later transferred into an aqueous solution of 60% ethanol with 10% diethylene glycol. In 2005 most material was fixed in 96% ethanol in view of future genetic studies. Representative material was deposited in the Muséum d’histoire naturelle de la Ville de Genéve and the Naturhistorisches Museum Wien (Crust. Coll. nos. 20685-20704). RESULTS Hemimysis anomala G. O. Sars, 1907 Figs 1A, 2 The sampling campaigns in 2004-2005 yielded this species at 19 stations. These are shown in Fig. 2 together with published data from previous campaigns and litera- 68 K. J. WITTMANN pr NG A ar VA m K da Û AA KA EU A di NN AN y 3 mm B 2 A 2 D 2 c FIG. 1 The currently most expansive mysid species in continental waters of Europe: Hemimysis ano- mala (A), Paramysis lacustris (B), Katamysis warpachowskyi (C) and Limnomysis benedeni (D). Adult females from the Danube River in Vienna (A, C, D) and from the Danube Delta (B). Adults of the four species are macroscopically distinguished by body proportions, and by the size and form of eyestalks and cornea. A, C and D taken from slightly schematized drawings in Wittmann (2002). ture data. A number of first records were noted in 2005: H. anomala was found south- wards of the already published range limit in the Rhine system, namely in Strasbourg (km 295) and Basel (km 170). The finding at Strasbourg is represented by a total of seven specimens trapped in five out of seven bottles exposed over night (24/25 Oct. 2005) on the right river bank, in the Port de la Redonne and at the main harbour entrance (48.59372°N, 7.80210°E). The new southern limit of H. anomala in the Rhine system is now marked by the harbour of Kleinhiiningen (km 170), where a total of five specimens were found in three out of six baited bottle traps exposed over night (47.58757°N, 7.59340°E; 25/26 Oct. 2005; first record for Switzerland). A single specimen of H. anomala, trapped at Danube-km 1865 at the entrance of the dock harbour of Bratislava, represents the first record of this species for Slovakia (48.12443°N, 17.15068°E; 16 Sept. 2005). A number of new records fill the large gap MYSID INVASION IN THE RHINE AND DANUBE 69 along the middle course of the Danube River: the first record for Hungary was taken at the right river bank at km 1578 in Dunaüjväros, where 40 specimens were collected by drift net (46.95704°N, 18.95602°E; 28/29 July 2005). A single specimen of H. ano- mala collected by drift net at the right river bank at Danube-km 1333 in Vukovar represents the first record for Croatia (45.35380°N, 19.00383°E; 29/30 July 2005). The first record for Serbia was taken at the right river bank at km 1059 in the impoundment basin in Veliko GradiSte, where 1349 specimens were collected by drift net (44.76710°N, 21.51488°E; 30/31 July 2005). This station lies 19 km above the entrance into the Carpathian breakthrough of the river. Mass occurrences were found at certain stations in turbid waters with varying ionic content, i.e. in the impoundment basin (371 y S/cm) of the Danube River at km 1059 (14 days after the peak of a moderate flood event), in the Mittellandkanal (km 4) at 1600-1630 uS/cm and in the winter harbour of Linz (Danube-km 2132) at 296 uS/cm. Remarkably, the first two of these three stations simultaneously showed a mass occurrence of Limnomysis benedeni (see below). The H. anomala mass occur- rences in the Mittellandkanal and at Linz were local. Inspections of the same Linz harbour site, this time operating the same hand net from a swimming platform in the same way, yielded no Hemimysis on 18 June 2004, 1167 specimens on 15 July 2005 and again zero on 14 Aug. 2005. The rich yield was made three days after the peak of a strong flood event, the zero yields during periods of normal water levels. L. benedeni collected together with H. anomala with the same nets showed a completely different phenology: 16, 294, and 652 specimens, respectively. In 2004-2005 H. anomala was found only in strongly anthropogenic habitats. The measurements of abiotic factors are integrated in Table I. The 19 positive sampling stations showed massive structures for shore or river bank stabilization, such as large stones, boulders, concrete or steel. Additional structures included vegetation, such as algal cover on hard substrate or loose stands of submerged macrophytes. Hand net samples taken during the day contained Hemimysis at only six stations where the animals were found between large stones or boulders in <1.5 m depth (<3.5 m exam- ined). A slightly higher abundance in shallow water was revealed by 36 positive bottle traps exposed over night at ten stations in 0.6-3.5 m depth. Limnomysis benedeni Czerniavsky, 1882 Figs 1D, 2 The records of this species yielded by the Cousteau-Expedition 1991 and own excursions in 1985-2005 in the Rhine and Danube systems, and in waters of western Turkey, are given in Fig. 2, together with available literature data. The Cousteau- Expedition captured this species on the banks of the Danube River at kms 2112, 1533, 1070, 954, 877, 851, 624, 506, 496 and 296. Own sampling in 2004-05 yielded this species at 92 stations. The following first records were taken in 2005: the species was found southwards of the already published range limit on the Rhine River and in its parallel, artificial side arm Grand Canal d’ Alsace, at four stations between Breisach (km 225) and Basel (km 170). Hand net collecting in the harbour of Kleinhiiningen in Basel (47.58740°N, 7.59142°E) yielded the first record of the order Mysidacea for Switzerland. Hundreds of Limnomysis specimens were found there during two inspec- tions (10 July and 26 Oct. 2005), with maximum densities in 0.2-1.0 m depth among 70 K. J. WITTMANN omala x y! FEU North Sea Limnomysis benedeni ~~ Pa LS > HR Pj North Sea I Vistula Black Sea A 1900 - 1940 O 1941 - 1984 27 è erre, 01 © 1985 - 2001 $ i i) Marmara 2) Sakarya D 3 Mediterranean as è LN _ Cat O 2002 - 2005 TS = 4 en Er FiG. 2 Distribution of Hemimysis anomala and Limnomysis benedeni in the Rhine-Danube system and other waters. Overlapping symbols are arranged with the older records on top, so that the MYSID INVASION IN THE RHINE AND DANUBE Wil the filiform, green algae covering vertical concrete and steel walls. Eight stations fur- ther upstream in the river and in Lake Constance (Bodensee) contained no mysids. Also, a westward range expansion in and from the Rhine system was noted by findings in the Mosel River at Riol (49.79697°N, 6.79727°E), Canal de la Marne au Rhin at Strasbourg (48.59133°N, 7.78302°E), Canal du Rhöne au Rhin at Niffer (47.71270°N, 7.50190°E) and at Mulhouse (47.75442°N, 7.34880°E). L. benedeni was found for the first time in navigation channels of NW Germany: Mittellandkanal at km 4 and km 35, and Dortmund-Ems-Kanal at km 48. The only mysid species found in the Ems-Jade- Kanal was the marine euryhaline species Praunus flexuosus (O. F. Miiller) in the brackish reach (S = 26-31) at km 70-71. Regarding the Danube system, first records of L. benedeni are here given for the Drava River at km 20 in Croatia, for the Tisa River 9 km above mouth into the Danube River (collection material from 1977) in Serbia, and for the DTD-Channel-System that connects (among others) the Tisa and Danube in Serbia (record in the Veliki Kanal at Srbobran, 45.54040°N, 19.78738°E). Mass occurrences were found at certain stations in turbid waters with varying ionic content, i.e. in the Mittellandkanal (km 4) at 1600-1630 uS/cm, in the Neusiedler See at 3580 uS/cm, in Lake Balaton (811 y S/cm) and in the impoundment basin (351- 371 uS/cm) of the Danube River at km 1059 (14 days after the peak of a moderate flood event). The first three of these mass occurrences were local, while the impound- ment basin was not tested in this regard. In 2004-2005 L. benedeni was found in a broad range of habitats from near- natural to strongly anthropogenic. The measurements of abiotic factors are integrated in Table I. Maximum densities of Limnomysis were observed among vegetation, with a preference for stands in only 0.3-1.0 m depth; nonetheless, L. benedeni was also present down to the greatest depth sampled (4 m). Very high densities were found among brush-like plant structures, such as cover of long, filiform algae on hard sub- strates or fine roots of Salix trees, or in dense stands of submerged macrophytes or in flooded terrestrial weeds. Densities were generally lower on and in the space between stones and over soft substrate. A few animals even lived on bare walls of wood, concrete or steel. As observed by snorkelling during the day, the animals swam a few cm from the substrate or were in direct contact with it. Hand net collecting and horizontal surface tows from boats showed that part of the population kept a greater distance from the substrate during the night, with a few specimens swimming up to the surface. expansion history can be traced from the symbols indicating different periods. White lines indi- cate man-made canals, black lines rivers. Original records (in part covered by superimposed symbols) are from waters of the Rhine-Danube system, channel systems in France and north- western Germany, and from waters of western Turkey. Literature records are from Salemaa and Hietalahti (1993), Eggers er al. (1999), Kelleher er al. (1999), Wittmann er al. (1999), Tittizer et al. (2000), Borcherding (2001), Haesloop (2001), Grigorovich et al. (2002), Rehage & Terlutter (2002), Wittmann (2002), Zettler (2002), Rudolph & Zettler (2003), Lundberg & Svensson (2004), Arbaciauskas (2005), Horecky et al. (2005), Janas & Wysocki (2005), Michels (2005), Müller et al. (2005), Dumont (2006), Holdich et al. (2006) and additional literature cited in these publications. 72 K. J. WITTMANN Katamysis warpachowskyi G. O. Sars, 1893 Figs 1C, 3 The sampling campaigns in 2004-2005 yielded this species at 21 stations. These are given in Fig. 3 together with published data from previous campaigns and litera- ture data. This species was found only in the Danube system, where the following new records were noted: on 16 June 2004, one adult female was taken with hand net close to the fish-ladders, immediately below the impoundment weir of Greifenstein (48.3495°N, 16.2465°E) at river-km 1949 in Austria. This finding shifted the known distributional limit of K. warpachowskyi by only 13 km further upstream. This was within the same impoundment basin in which the species had already been detected three years earlier (Wittmann, 2002). A number of new records fill the large gap along most of the middle course of the Danube: the first record for Croatia was nine spe- cimens of K. warpachowskyi sampled by hand net in the yachting harbour of Vukovar at river-km 1333 (45.35313°N, 19.00408°E; 29 July 2005). The first records for Serbia were one specimen taken with the hand net at the right river bank near Veliko Golubinje (44.46675°N, 22.17268°E) in the Carpathian breakthrough at km 986, followed by eight specimens taken with the drift net in the impoundment basin at Veliko GradiSte at km 1059, and then 46 specimens with the hand net in the winter harbour of Kovin at km 1109 (all three samples on 30/31 July 2005). In 2004-2005 K. warpachowskyi was found in a broad range of habitats from near-natural to strongly anthropogenic. The measurements of abiotic factors are inte- grated in Table I. Maximum densities of Katamysis were observed on boulders, large stones and on coarse to soft substrates with plant debris, with a preference for stands in =1.5 m depth, but also present in 0.2-1.5 m. This species was also found between submerged macrophytes, between flooded terrestrial weeds and in accumulations of stones and shells. In most hand net samples, K. warpachowskyi occurred together with Limnomysis benedeni and the snail Viviparus acerosus (Bourguignat, 1862). Generally, the drift net and bottle traps yielded only small numbers of Katamysis. No mass oc- currences were recorded for this species. Paramysis spp. Figs 1B, 3 The records of P. lacustris (Czerniavsky, 1882) yielded by the Cousteau- Expedition 1991 and by own excursions in 1985-2005, in the Danube system, are given in Fig. 3, together with available literature data. The Cousteau-Expedition collected Paramysis species only in the lower reach of the Danube River. A total of 13 specimens of P. lacustris were taken at km 624, km 595, km 489 and km 296. Only fragments of two individuals of P. intermedia Czerniavsky, 1882 were sampled at km 624 and km 595. The sampling campaigns of 1985-1998 yielded P. lacustris in the Predelta (Lake Sinoe), in the main river branches of the Danube and in a great number of lakes, side arms and canals of the Danube Delta (325-3020 uS/cm). The excursion to western FIG. 3 Distribution of Katamysis warpachowskyi and Paramysis lacustris in the tributary system of the Black Sea and other waters. Overlapping symbols are arranged with the older records on top, so that the expansion history can be traced from the symbols indicating different periods. White MYSID INVASION IN THE RHINE AND DANUBE 73 ey sh < EN Azov Sea / di grato u Black Sea Di 35° > mel Li nu Mi N \ Balaton -f A 1900 - 1940 O 1941 - 1984 © 1985 - 2001 © 2002 - 2005 lines indicate man-made canals, black lines rivers. Original records (in part covered by super- imposed symbols) are from waters of the Danube system and of western Turkey. Literature records are from Bäcescu (1940, 1954), Mordukhai-Boltovskoi (1964, 1979), Pligin and Yemel’yanova (1989), Komarova (1991), Tarasov (1996), Daneliya (2001, 2003), Arbaciauskas (2002), Grigorovich et al. (2002), Wittmann (2002), Gruszka et al. (2003), Audzijonyte (2006) and additional literature cited in these publications. 74 K. J. WITTMANN Turkey in July 1988 showed the presence of this species in a total of five lakes (Fig. 3) near the coasts of the Marmora Sea (Uluabat Gölü, Kus Gölü) and the Black Sea (Durusu Gölü, Sapanca Gölü, Akgöl). All these lakes contained clearly freshwater in the conductivity range of 270-500 uS/cm at surface temperatures of 27-30°C. One immature female of P. lacustris surprisingly appeared in a hand net sample taken by snorkelling in stands of Myriophyllum in 1.5 m depth in the almost isolated (see ‘Discussion’) backwater Alte Donau in Vienna (at height of Danube-km 1930; 48 .24213°N, 16.42132°E; 10 June 2004). Despite great efforts, no further specimens belonging to this species could be found in waters along the upper and middle reaches of the Danube in 2004-2005. DISCUSSION RANGE EXPANSION HISTORY OF HEMIMYSIS ANOMALA: This species has populated vast areas of continental Europe, originating from an exclusively Ponto-Caspian range in the early 20th century. Indigenous populations were found in the salinity range of 0-18 (Table I), living over hard substrate or algal vegetation along the shores of the Caspian Lake, and the Black and Azov Seas. Before human interventions, the species was known as an essentially estuarine or ‘marine’ (in terms of NW Black Sea salini- ties) animal, penetrating not more than 40-60 km into the lower reaches of rivers. Bacescu et al. (1971) pointed it out as the only mysid species of Caspian origin occur- ring in the open Black Sea. Before 1998 the records of H. anomala in the Danubian range were restricted to the delta up to 34 km distance from the Black Sea (Bäcescu, 1954; Popescu, 1963; most later records are merely citations of these two papers). This was essentially confirmed by own sampling in the delta, where H. anomala was rare and only taken in 1995 at mile 13.6 of the Sulina branch, and in 1998 at km 68 of the Sfintu Gheorghe branch (Wittmann er al., 1999). Due to artificial deepening of the Sulina branch, a distinct influence of saline waters invading up to km 80 inland from the sea were reported during the drought season in August-September (Bondar, 1983). Thus the ‘original’ Danubian range of this species fitted well to the distribution of an essentially estuarine species (Bäcescu, 1954) that only marginally penetrates into freshwater. H. anomala started its range expansion in 1957 as an invader of European brackish and freshwaters by intentional introduction into an impoundment basin of the Dnieper River in the course of fisheries management (Pligin & Yemel’yanova, 1989). The main aim was that this photophobic species would enrich the spectrum of fish food in deep impoundment basins. This actually worked: Hemimysis established populations in up to 20-50 m depth (Zhuravel, 1960). This approach was pursued in other reser- voirs and lakes of the former Soviet Union (Grigorovich er al., 2002). From impound- ment basins in tributaries of the Baltic Sea the species spread into brackish coastal waters of Lithuania, Finland, Sweden and Poland (Razinkovas, 1996; Arbaciauskas, 2002; Lundberg & Svensson, 2004; Janas & Wysocki, 2005). From there it invaded more western Baltic tributaries, together with the River Odra (Gruszka et al., 2003), the system of navigation channels in northern Germany (Eggers er al., 1999; Rehage & Terlutter, 2002; Zettler, 2002; Rudolph & Zettler, 2003; Miiller et al., 2005) and, again westwards, tributaries of the North Sea (Elbe and Weser systems; Haesloop, MYSID INVASION IN THE RHINE AND DANUBE 75 2001; Horecky er al., 2005). Additional, so far unpublished records from waters connected with the channel system in north-eastern Germany were kindly communi- cated by E. Rudolph (Havel): Spree River at the village of Radinkendorf, 52.21133°N, 14.27235°E, 24 Dec. 2004, leg. Wulf (Lamitsch), and Havel River at the village of Bahnitz, 52.49717°N, 12.40410°E, 2 Sept. and 1 Nov. 2005, leg. Rudolph. In 1997-1998 H. anomala showed an explosive range expansion by appearing in vast areas of the Rhine-Main-Danube system, ranging from brackish and freshwaters in the Rhine Delta, to the middle Rhine, the rivers Main and Neckar, and the upper Danube. Already in 1999 this species was reported in the Westerschelde estuary in Belgium (Verslycke et al., 2000). Shortly thereafter, in 2000, it appeared at two stations in the middle reach of the Meuse River (Usseglio-Polatera & Beisel, 2003). In June 2005 (four months before my findings reported above) it appeared for the first time in the north-east of France (Dumont, 2006). Due to an unknown introduction, the species surprisingly appeared in 2005 in waters of the Trent River in the English Midlands (Holdich et al., 2006). Even if certain populations may have been overlooked in earlier years, the speed of range expansion clearly points to anthropogenic dispersion by navigation, as also pointed out below for Limnomysis benedeni. The extent and speed of the invasions by both species were probably favoured by the opening of the Main-Danube-Channel in 1992. From the currently available data for H. anomala one cannot definitely conclude whether the invaders of 1997-1998 originated from Baltic populations and transgressed this channel from north to south, or if they came from the Black Sea (Danube Delta) and transgressed the channel from south to north (see discussions in Kelleher er al., 1999; Wittmann et al., 1999; Bij de Vaate et al., 2002; Müller er al., 2005). Multiple immigrations cannot be excluded (Miiller et al., 2005). Future genetic studies could be helpful to clarify such questions. With the anthropogenic appearance in the upper Danube in 1998 (Wittmann er al., 1999) a very large gap of records between km 1933 and km 68 became obvious, covering the entire middle and the lower reaches of the river, and part of the delta. A total of 37 hand net and drift net samples taken by me in 2001-2004 at 20 stations between km 1931 and km 1722 yielded Hemimysis only at km 1931. In accordance with this, KoSel er al. (2003) reported only L. benedeni and Katamysis warpachowskyi but no H. anomala for the fauna of Slovakia. Possibly there was a disjunction in the Danubian distribution of H. anomala over a few years. A substantial part of the gap was closed in 2005 by the above-presented first records for Slovakia, Hungary, Croatia and Serbia. As in L. benedeni, a further upstream (southwards) expansion was noted in 2005 when H. anomala was recorded from the French (Dumont, 2006) and the Swiss reach of the Rhine for the first time. Most of these findings were made in harbours, pointing to a possible anthropogenic dispersion by navigation. The present paper is the first to demonstrate the mass occurrence of H. anomala in rivers. The first hypothesis is that these mass occurrences were local and short-lived, possibly caused by flood events that had displaced the mysids by passive drift and/or by active avoidance of strong currents, so that the animals accumulated in more calm zones (harbour at Danube-km 2132, or deep and wide impoundment basin at km 1059). This will be the subject of future research. 76 K. J. WITTMANN RANGE EXPANSION HISTORY OF LIMNOMYSIS BENEDENI: As in the preceding species, L. benedeni also populated vast areas in continental Europe, starting from an essentially Ponto-Caspian distribution in the early 20th century. Indigenous popu- lations were found in the salinity range of 0-14 (Ovéarenko et al., 2006; Table I), with the greatest population densities in the oligohaline range (0.5-5), but the greatest number of populations in freshwater. Mass occurrences were only found at pH > 7.7. A favourable development in alkaline waters is also indicated by findings of Szalontai et al. (2003) that the oxygen consumption of juvenile Limnomysis is lower at pH 8.4 than at pH 5.4. The animals mainly dwelled among vegetation in stagnant and slow- flowing waters in the tributary systems of the Caspian Lake and the Black and Azov Seas. Own sampling in July 1988 revealed several populations in streams and lakes belonging to the drainage systems of the Marmora Sea and the Black Sea in western Turkey (Fig. 2). In most large Ponto-Caspian river systems this species penetrated several hundred kilometres beyond the oligohaline reach of the mouth area. The Danubian distribution once extended from the saline mixing zone in the Black Sea up to river-km 460. In any case, the primary distribution did not extend into the former cataract stretch between km 941 and km 1040 in the Carpathian breakthrough. L. benedeni started its range expansion as an invader of European freshwaters in 1946, when it surprisingly appeared in the winter harbour of Budapest at Danube- km 1644 (the erroneous indication in Dudich, 1947: Tab. 1, was corrected by Woynarovich, 1955; now there is a bridge at this point, today’s harbour entrance is at km 1642). During the following five decades L. benedeni expanded its distribution in the Danube upstream to the beginning of the Main-Danube-Channel at Kelheim in a series of small (1-173 km) steps. By incorporating additional literature and data from collections, the list given by Wittmann er al. (1999) is here updated. Fourteen stations are currently known: km 1683 (year 1949), km 1787 (<1953), km =1872 (<1954), km 1911 (1973), km 1919 (1982), km 1920 (1983), km 2093 (1986), km 2112 (1991), km 2132 (1992), km 2214 (1993), km 2228 (1994), km =2320 (1995), km 2376 (1997), km 2410 (1998). Limnomysis transgressed the Main-Danube-Channel probably shortly be- fore 1997 and thereafter colonized vast areas of the Rhine system, probably in down- stream direction, in about 1997-1998. Already in 1998 a further upstream (southwards) expansion was noted by the appearance of Limnomysis in the French reach of the Rhine (Wittmann & Ariani, 2000). Most of the observed upstream expansions probably reflect passive transport by ships. Specimens of this species were found on the outside hull of ships, and also inside, particularly in rest water (bilge water) and in cooling- water filters (Reinhold & Tittizer, 1998; Wittmann et al., 1999; Bij de Vaate ef al., 2002). The new records, presented above for the Mosel River, the upper Rhine and the navigation channels in France, all came from navigable courses and therefore poten- tially reflect passive transport by navigation. L. benedeni was intentionally introduced mainly in the 1950-1960s in eastern Europe, in order to enrich the supply of food for fish. These operations started in 1948 with the release of mysids into a hydropower reservoir of the Dnieper River, followed during the next decades by a great number of other waters in the former Soviet Union (Grigorovich et al., 2002), particularly in Lithuania (Arbaciauskas, 2002), and in Hungary (Lake Balaton; Woynärovich, 1955). Introduction of Limnomysis into Lake MYSID INVASION IN THE RHINE AND DANUBE Vil Aral (Kazakhstan and Uzbekistan; Mordukhai-Boltovskoi, 1979) was possibly due to inadvertent stocking (Aladin er al., 2003). From introduction sites along tributaries of the Baltic Sea L. benedeni spread into coastal waters (Olenin & Leppäkoski, 1999; Arbaciauskas, 2002) and other tributaries, including the waters of the Odra system in Poland (Michels, 2005). The new records presented above for navigation channels in north-western Germany may originate from a (north-) eastwards expansion of popu- lations from the Rhine system, although a westwards expansion from Baltic tributaries appears possible as well. RANGE EXPANSION HISTORY OF KATAMYSIS WARPACHOWSKYI: In most large Ponto- Caspian river systems Katamysis originally penetrated up to 200 km into rivers of the Black Sea, or 120 km into those of the Caspian Lake. Black Sea populations were found in a salinity range of only 0-5, with the optimum in freshwater; Caspian popu- lations showed a broader range (0-14; Table I). A similar difference was also observed in Paramysis lacustris (Table I). In 2000 K. warpachowskyi surprisingly appeared in waters of the Don system (Fig. 3), from where it had never been reported before. According to Daneliya (2001), this may indicate a recent immigration from the Volga River via the Volga-Don navigation channel, which was completed in 1951-1952. Deliberate introductions were of little importance for the distribution of this species. In 1956 it was released into the Dubossary reservoir (Dediu, 1966), upstream of its pre- viously known distribution range in the Dniester River. For more details on the distri- bution, ecology, bionomics and possible modes of dispersion of K. warpachowskyi, see Wittmann (2002). In 2001 this species was surprisingly detected at km 1936, more than 1700 km upstream of its previously known limit of distribution in the Danube (Wittmann, 2002). So far it has not transgressed the limits of the watersheds belonging to its exclusively Ponto-Caspian range. With the new record at km 1949 in 2004 the upper limit shifted by only 13 km and thus remained within the same impoundment basin. This was not surprising, because the comparatively rheophilic K. warpachowskyi is considered to be capable of swimming against the water currents observed along the banks of this im- poundment basin under normal hydrological conditions. Within the same month (Oct. 2001) an already differentiated distribution was found along the Austrian, Slovakian and Hungarian reach, down to km 1769 (Wittmann, 2002). This campaign was continued in 2004 with sampling down to Danube-km 1722, in the side arm Kis Duna (47.76335°N, 18.69988°E; 23 June 2004) near its flow back into the main river, and in 2005 down to km 986 in the Carpathian breakthrough, yielding the first records for Croatia and Serbia. In no case was a lower limit documented by negative downstream samples, because the species was always present at the lowest station examined. Extrapolating this, it seems likely that before 2005 a continuous population range already existed from kms 1936-1949 down to the delta. The absence of this species in samples taken at 26 stations in the Hungarian and Austrian reach in 1997-1998, as well as in samples of preceding campaigns (Nesemann et al. 1995; Wittmann, 1995), including the Cousteau-Expedition 1991, suggest that the middle and upper reaches of the Danube were colonized within a few years before 2001. 78 K. J. WITTMANN TABLE I. Hydrological and physicochemical parameters at sampling stations of four Ponto- Caspian mysid species?. Samples selected / parameter Hemimysis anomala Limnomysis benedeni m+S.D5 range n m + S.D.P range n All samples: Depth (m) 4.037 + 5.422 0-60 103 1.452 + 1.166 0-10 479 Water current (m/s) 0.152 + 0.222 0.0-0.81 98 0.106 + 0.194 0.0-1.5 443 Temperature (°C) 17.21 + 447 2-28 78 19.38 + 4.79 0-31 353 Salinity 2.140 + 4.209 0.1-18.0 87 0.594+1.659 0.0-14.0 345 Conductivity (uS/cm) 3792 + 6947 279-29200 87 1215+2774 195-22300 345 pH 7.866+0.503 6.35-8.65 63 7.970 +0.599 5.54-9.57 267 Carbonate hardness (°d) 8.642 + 0.975 6-12 60 9.209 + 3.455 3-30 191 Oxygen content (mg/l) 719741422 3.99-10.80 63 7.922 +1994 3.75-18.10 263 Turbidity (NTU)® 28.61 + 26.29 5-137 63 34.17 + 41.44 1-274 266 All regions, drift samples excluded: Water current (m/s) 0.031 + 0.078 0.0-0.35 69 0.073 + 0.148 0.0-1.0 406 All sample types, samples in the Caspian Lake excluded: Salinity 1.475 + 3.317 0.1-18.0 82 0.480 +1.274 0.0-14.0 340 Conductivity (uS/cm) 2697 + 5486 279-29200 82 1025 + 2139 195-22300 340 Samples selected / parameter Katamysis warpachowskyi Paramysis lacustris m+S.D.b range n m+S.D.b range n All samples: Depth (m) 2.011 + 1.570 0-10 84 3.908 + 4.896 0-48 122 Water current (m/s) 0.146 + 0.283 0.0-1.5 74 0.061 + 0.122 0.0-0.5 83 Temperature (°C) 17.72+4.61 0-27 TAI 19.82 + 5.54 0-30 70 Salinity 0.919 + 2.915 0.1-13.8 74 1.909 + 3.269 0.0-13.8 82 Conductivity (4 S/cm) 1738 +4828 279-22873 74 3451 +5450 270-22873 82 pH 7.693 +0.729 6.13-9.35 54 8.344 #0.625 7.42-9.42 24 Carbonate hardness (°d) 9.294 + 3.437 5-25 Sil 9.889 + 2.374 8-16 18 Oxygen content (mg/l) 8.359. 2.1.9727 .3.99160.99 54; 7.401 #1.979 5.01-12.17 23 Turbidity (NTU)° 30.02 + 42.81 1-272 54 43.30 + 49.97 1-194 23 All regions, drift samples excluded: Water current (m/s) 0.122 + 0.282 0.0-1.5 67 0.051 + 0.109 0.0-0.5 80 All sample types, samples in the Caspian Lake excluded: Salinity 0.159 + 0.413 0.1-5.0 68 1.179 + 1.956 0.0-8.0 TS Conductivity (x S/cm) 475 + 721 279-8800 68 2242 + 3338 270-13800 75 a data sources as in Figs 1,2 € nephelometric turbidity units b mean + standard deviation RANGE EXPANSION HISTORY OF PARAMYSIS SPP.: In the large Ponto-Caspian river systems P. lacustris originally penetrated about 200-600 km into the Danube, Dnieper, Don and Ural Rivers, and about 900 km into the Volga. Almost the same penetration distances were found for P. intermedia, with the remarkable difference that it reached as far as about 2000 km from the Caspian Lake into the Kama River, a northern tribu- tary of the Volga. In the Caspian Lake P. lacustris showed a more continuous distri- bution in estuarine waters to freshwaters along the west coast (Fig. 3), while the dis- tribution in the more saline Black Sea showed a distinct maximum at coastal stretches influenced by the freshwater input of large rivers. As shown in the ‘Results’, P. lacus- tris was found in freshwater lakes of western Turkey, where it locally occurs together MYSID INVASION IN THE RHINE AND DANUBE 79 with Limnomysis benedeni (Figs 2, 3). It is remarkable that P. lacustris was also found in Lake Beizehir (Bacescu 1948, 1966) in the Anatolian highland. The salinity range of P. lacustris was 0-14 (Table I); the range was 0-12 for P. intermedia. In 1936 the upper limit of the known distribution of P. lacustris and P. inter- media in the Danube system was in the Orlea swamps (Lake Potelu) that drain into the river at km 644 (Bacescu, 1940). In 1991 this limit again coincided for both species and was still close to the previous position, downstream of the mouth of the Iskar River at km 624, as derived from the material of the Cousteau-Expedition. In 2004 the single specimen of P. lacustris (see ‘Results’) surprisingly was recorded at the height of Danube-km 1930 in an almost isolated backwater. This back- water is connected with an artificial side arm of the river solely by an underground pipe and pumping station for stabilization of the water level. This was 1286 km upstream from the nearest documented population (Lake Potelu), or 620 km upstream from the nearest confluence with the drainage system of an introduction site (Lake Balaton). Due to its relatively isolated position, this single specimen does not provide definite evidence of a viable population of P. lacustris unless further material is found, and is therefore indicated with a question mark in Fig. 3. This species, along with P. inter- media, already had disappointed previous expectations of existing populations: accor- ding to Bacescu (1966 and pers. comm.) and Bacescu et al. (1971), Paramysis stocks were taken from Lake Oltina (drainage into Danube at km 338) and transplanted into Lake Balaton (drainage at km 1497) by Hungarian fishery biologists. A first attempt was made with P. intermedia in 1955 and a second one with both species in 1964 (P. lacustris indicated as P. kowalewsky in Bacescu, 1966). Despite long-term monitoring by the biological station at Tihany (e.g., Ponyi et al., 1971) and the detailed study of Muské & Leitold (2003) on the distribution of malacostracans in Lake Balaton, no Paramysis species were ever reported from this lake. Own sampling in 1983, 1997, 1998 and 2005 always yielded Limnomysis benedeni, but no Paramysis species. The same result was achieved by three hand net samples taken by L. Forré and H. Nesemann at different stations in Lake Balaton in 1990-1991 (Nesemann et al., 1995). Both species did not survive the first winter after introduction into the Rybinsk reservoir in the most northern reach of the Volga, 2900 km from the Caspian Lake (Slynko et al., 2002). By contrast, introduction of P. intermedia into hydropower reservoirs in the middle reach of the Volga in 1957-1966 was successful (Borodich & Havlena, 1973). Extensive successful introductions of P. lacustris and P. intermedia were carried out in vast areas of the Ukraine, Crimea Peninsula, Uzbekistan, Kazakhstan and Kirgizstan, particularly to the lakes Aral, Balkhash and Issyk-Kul (Mordukhai-Boltovskoi, 1979; Komarova, 1991; Grigorovich et al., 2002; Aladin et al., 2003). Following release into water reservoirs and lakes draining into the Baltic Sea, P. lacustris established itself in a great number of waters of Lithuania (Arbaciauskas, 2002). It was also introduced in reservoirs of the Western Dvina (= Daugava), from where it subsequently spread into coastal lakes of Latvia (Gasiunas, 1972); additional introductions include the Volkhov reservoir that drains into Lake Lagoda (Zhuravel, 1969). A secondary spread, subsequent to introduction, was reported for P. lacustris populations in the Neman, Western Dvina, Dnieper and Volga Rivers. In the 1960s this species appeared in the Gulf of Finland (Jansson, 1994; not shown in Fig. 3 due to missing details on location). 80 K. J. WITTMANN MODES AND FACTORS OF DISPERSION: Van der Velde et al. (2000) listed the success factors for crustacean invaders, most of which were also relevant for mysids. Unintentional introductions of mysids were mainly referred to navigation, particularly transport in ballast water, in cooling-water filters and on the outside wall of ships; construction of waterways and inadvertent stocking were also reported to be of major importance. Jazdzewski & Konopacka (2002) argued that the massive invasion of Ponto- Caspian species in central to western Europe may have been facilitated by the in- creasing ionic content in large rivers caused by industrial und agricultural pollution during recent decades. This may be relevant for the invasions by the more (Hemimysis anomala) or less (Limnomysis benedeni) halophilic mysids, but needs to be differen- tiated for the Rhine and Danube Rivers: after several decades of increase, the ionic content distinctly decreased in these river systems over the last 1-2 decades (Weilguni & Humpesch, 1999; Van der Velde et al., 2000). According to Kelleher et al. (2000) and Van der Velde et al. (2000), the improved water quality of the Rhine may have facilitated the establishment of certain invaders. This is supported by own observations in the Danube, where the appearance of H. anomala in 1998 and Katamysis warpa- chowskyi in 2001 preceded the re-establishment of the stenoecious, indigenous snail Theodoxus danubialis (C. Pfeiffer, 1828) in 2004 (9 Sept., six snails sampled with drift net (!), together with L. benedeni; at river-km 1933 in Vienna), after four decades without positive records of the snail in the Austrian reach of the main river. In the 1930s this snail was common along river banks in Vienna, became rare in the 1950s, and was declared as already extinct for Austria by Reischiitz (1981), but has persisted in a relict population in the Leitha River, a tributary of the Danube (Frank, 1982). When sampling for neozoans in 2005, hundreds of Theodoxus were seen on stones and concrete walls along the banks of the Danube in Vienna. The Austrian reach of the Danube has been subject to intensive faunistic studies since the 1930s (e.g., Vornatscher, 1938; Moog et al., 1994), clearly confirming that all currently observed mysid species are neozoans rather than re-established indigenous faunal elements, such as the Theodoxus snails. Simberloff & Von Holle (1999) and Ricciardi (2001) explained part of the success of Ponto-Caspian invaders by “invasional meltdown”, i.e. that preceding in- vasions could facilitate subsequent invasions. Accordingly, the zebra mussel Dreissena polymorpha (Pallas, 1771) as a primary invader provided substrate, food and shelter for subsequent invaders. In the case of mysids, such interactions may be complex and require detailed future studies: K. warpachowskyi was repeatedly found in accumu- lations of stones and shells and thus may profit from empty Dreissena shells as shelter; adult H. anomala are strong predators of small crustaceans and may profit from the mass occurrence of the invasive amphipod Chelicocorophium curvispinum (G.O. Sars, 1895) as food in the Rhine and Danube systems. On the other hand, L. benedeni, K. warpachowskyi and the juveniles of H. anomala are mostly micro-herbivores: their food supply may be reduced by strongly enhanced plankton clearance rates (Ojaveer et al., 2002) due to invasions by D. polymorpha. The mysid invaders H. anomala, L. benedeni and K. warpachowskyi have in common that they are euryhaline and often found in great densities in harbours MYSID INVASION IN THE RHINE AND DANUBE 81 (Wittmann, 2002). Ports are considered to be distribution hubs of invertebrates (Ricciardi & Rasmussen, 1998). The strong increase of traffic along European water- ways during the last decades could partially explain the observed acceleration of mysid invasions. EFFECTS AT THE ECOSYSTEM LEVEL: Detrimental effects from introductions or invasions of Mysidae species in continental waters have been reported for a number of species, particularly Hemimysis anomala, Mysis diluviana Audzijonyte & Väinölä, 2005, M. relicta Lovén, 1862, Neomysis mercedis Holmes, 1896 and Paramysis lacus- tris. Damage at the ecosystem level was mainly related to overgrazing of zooplankton and subsequent effects (Rieman & Falter, 1981; Fürst er al., 1984; Koksvik et al., 1991; Ketelaars er al., 1999; Haskell-Presenter, 2004) and to out-competing of native species by invaders (Arbaciauskas, 2002, 2005; San Francisco Bay Institute, 2004). In parti- cular the top invader H. anomala is the focus of great concern due to its high expansion potential in combination with potential mass occurrences and due to its role as a pred- ator of zooplankton. This species is essentially omnivorous with increasing percentage of zooplankton consumption with increasing body size (Borcherding er al., 2006). One important prerequisite for mass occurrences is deeper water, because the photophobic mysids are pelagic during the night and inhabit the dark zone or the bottom during the day, where they are more protected from predation by visually oriented fish. Limnomysis benedeni and Katamysis warpachowskyi are essentially micro- herbivorous filter feeders with a small portion of their diet composed of invertebrates. This leaves them a more marginal role in the food web as compared to the two Mysis species and to H. anomala. L. benedeni and K. warpachowskyi are also more benthic than these species. Accordingly, Wittmann & Ariani (2000) and Wittmann (2002) did not expect strong detrimental effects by L. benedeni and K. warpachowskyi. No strong impact on the zooplankton community was observed after the invasion of L. benedeni in the backwater Alte Donau in Vienna (Wittmann & Ariani, 2000). After a first occur- rence in 1993 the population densities of Limnomysis peaked in 1996-2000, as revealed by hand net collecting along the shore. This peak was followed by a marked reduction in 2002-05, concurrent with increased water transparency in the course of successful restoration measures. The above-cited data suggest that mass occurrence is an important prerequisite for potential environmental damage by mysids. The mass occurrences here reported for H. anomala and L. benedeni were local and/or short-lived. This was not directly observed for the occurrence in drift samples, but mass occurrences would have been short-lived effects of flood events there as well (discussed above). FUTURE PROSPECTS: Upon first detection of Limnomysis benedeni in the French reach of the River Rhine, Wittmann & Ariani (2000) speculated that the range expansion of this species could continue into the system of navigation channels in France, from where the mysid could spread along the Rhöne River down to the Mediterranean coast. With the above-presented new records from several navigation channels, the first part of this prediction has become true earlier than expected. If its range expansion continues at the same speed, Limnomysis could reach the Mediterranean coast within a few years, with unknown consequences. The indigenous 82 K. J. WITTMANN populations of the closely related genus Diamysis Czerniavsky, 1882, represented by a number of species in brackish to freshwater bodies all around the coasts of the Mediterranean, could be particularly affected. If the diffusion of Hemimysis anomala along the Rhine system continues, it may enter Lake Constance (Bodensee) and, in the long term, also other deep lakes in Switzerland and neighbouring countries. This could represent a threat to the lake ecosystems in that it leads to overgrazing of zooplankton. Similarly, the Ponto-Caspian gammarid amphipod Dikerogammarus villosus (Sowinsky, 1894), a recent invader of Lake Geneva (Bollache, 2004), is expected to spread into other lakes and to seriously threaten the biodiversity in European freshwater ecosystems. ACKNOWLEDGEMENTS The author is grateful to I. B. Sket (Ljubljana) for the loan of collection material, to H. Nesemann (Vienna) and B. Csanyi (Budapest) for access to material of the Cousteau-Expedition of 1991, and to E. Rudolph (Havel) for communicating unpublished records of Hemimysis anomala in north-eastern Germany. LITERATURE CITED ALADIN, N.V., PLOTNIKOV, I. S. & FILIPPOV, A. A. 2003. Opportunistic settlers in the Aral Sea. 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The Phlyctimantis and Kassina of Cameroon (Amphibia, Anura, Hyper- oliidae). - The Cameroonian batrachofauna includes one species of Phlyctimantis and five species of Kassina, which are reviewed in this paper. Phlyctimantis have been found in only five 10 minutes side squares; the discussion of the status of the southernmost population shows that all known Cameroonian populations must be referred to P. boulengeri. In the genus Kassina, K. maculosa and K. decorata were tentatively synonymised by authors. Although they share a great number of mor- phological characters, they exhibit constant differences in cephalic and dorsal maculation and occupy different distribution areas. Thus, distinction at specific level appears to be the most appropriate status for these taxa. A new species, Kassina wazae, is described; it superficially resembles K. senegalensis but differs by several distinctive features, as vocal sac, lateral head maculation, presence of a small tarsal tubercle. For each species, many original data on distribution, ecology and phonocenosis are given. Finally, attention is drawn on the unexpected lack of a true forest-species of Kassina in the Cameroonian fauna. Keywords: Africa - Cameroon - batrachology - systematics - ecology. INTRODUCTION Parmi les rainettes africaines (famille des Hyperoliidae) une vingtaine d’especes se distinguent par les caracteres de leur tétard et de l’appareil vocal des males, sans équivalents dans le reste de la famille. Cet ensemble est actuellement divisé en cing genres très proches, dont le principal est Kassina. La plupart des espèces ne corres- pondent pas à la notion habituelle de « rainette » car elles ont la particularité de vivre sur le sol, ou en tout cas de peu s’élever sur la végétation, et cela méme lorsqu’elles sont forestières. Toutefois, dans le genre Phlyctimantis, le comportement et l’habitus sont ceux des rainettes: on peut admettre que ce genre est resté proche de la souche d’oü sont dérivés les autres Hypéroliens « kassinoides » (Laurent, 1976; Drewes, 1984). Au Cameroun, les représentants des genres Phlyctimantis et Kassina sont peu nombreux, mais restent pourtant assez mal connus. Il a donc semblé utile de rassembler ici des données inédites concernant leur systématique, leur chorologie et leur étho- écologie. MATÉRIEL ET MÉTHODES Les spécimens sur lesquels est basé ce travail ont été récoltés la nuit, en général par repérage auditif des mâles en cours d’activité vocale. Dans un but pratique, toutes les données relatives à l’ensemble du matériel étudié ont été regroupées dans un même tableau synoptique (Tab. 1): — nombre de mâles (M) et de femelles (F), — numéro de Manuscrit accepté 20.10.2006 88 J.-L. AMIET TABLEAU 1. Principales données relatives au matériel étudié (juvéniles non compris). Précisions dans le texte sous « Matériel et méthodes ». Espèce Nombre Numéro Localité Coordonnées Alti- Date Popu- de de de la tude de lations spécimens collection maille capture Ph. boulengeri Ouest 4M 78.052-055 Fainschang 9°20°-30° x 5°30°-40° =200 14.IV.78 Sud 4M 87.001-004 Tom 10°30’-40’ x 2°20’-30’ =450 02.1.87 K. maculosa Centre 4M 70.739-742 Olembé 11°40’-50’ x 4°10’-20’ =460 17.IX.70 3M 71.482-484 Foulasi (Obala) 11°30°-40° x 4°00’-10’ =540 19.IV.71 3M 71.1051-1053 Elomzok 11°30’-40’ x 4°10’-20’ =500 13.IX.71 6M,1F 71.1073-1077; 71.1143-1144 Elomzok / Minkama 11°30’-40’ x 4°10’-20’ =500 13.IX.71 2M 71.1068-1069 Binguela 10°20’-30’ x 3°40’-50’ =700 14.IX.71 4M 72.655-658 Foulasi (Obala) 11°30’-40° = 4°00’-10’ =540 12.IX.72 2M 75.303-304 Elomzok 11°30’-40’ x 4°10’-20’ =500 15.X.75 2M 83.013-014 Yambassa / Bombato 11°10°-20° x 4°20’-30’ =400 29.KX.83 Ouest 2F 70.432-433 Nsoung 9°40°-50° x 4°50°-5° = 1400 19.11.70 4M 71.737-738; 71.910-911 Dschang 10°00’-10° x 5°20’-30’ = 1400 24.V.71 1M 79.048 Niamboya (Yimbéré) 11°30’-40’ x 6°10’-20’ 760 24.V1.71 Mt Cameroun 2M,1F 81.146-148 Bonianango 9°10°-20° x 4°00°-10° =550 11.VII81 Adamaoua 2M 81.088-089 Hourso Manang 13°10°-20° x 7°20’-30’ 900 20.V.81 1M 81.109 Ndjaouro Ndekaou 13°00’-10’ x 7°20’-30’ 900 20.V.81 3M 82.095-097 Mbella Assom 12°50’-13° x 6°30’-40’ =900 20.V.82 K. decorata Ouest 10M,1F 71.683-692; 71.801 Santchou 9°50’-10° x 5°107-20? 7 2700 2A 1M 71.1120 Santchou 9°50’-10° x 5°102-207 2700 22-7 2M 72.301-302 Lac Manengouba 9°40°-50° x 5°00’-10’ =2000 29.11.72 Adamaoua 12M 71.957-964; 71.1005-1008 Sadolkoulaye 13°50°-14° x 7°10’-20° = 1200 29.VI.71 1M 71.1011 Ngaoundéré Vina 13°40’-50’ x 7°10°-20° =1100 02.VII.71 6M,1F 73.245-249; 73.306-307 Sadolkoulaye 13°50’-14° x 7°10’-20’ = 1200 28.1V.73 1M,2F 73.397-399 Wakwa 13°30’-40’ x 7°10°-20° = 1200 23.V1.73 16 M,2F 81.229-246 Sadolkoulaye 132507-142 272102207 = 1200 179281 PHLYCTIMANTIS ET KASSINA DU CAMEROUN 89 TABLEAU 1 (suite). Espèce Nombre Numéro Localité Coordonnées Alti- Date Popu- de de de la tude de lations spécimens collection maille capture K. senegalensis Sud 1M 70.132 Nanga Eboko 12°20°-30° x 4°40°-50° =600 VIIL.70 1M 70.732 Elomzok / Minkama 11°30°-40° x 4°10°-20° =500 05.X.70 1M 71.1054 Elomzok 11°30°-40° x 4°10°-20° =500 13.IX.71 1M 71.1078 Elomzok / Minkama 11°30°-40 x 4°10°-20° +500 13.IX.71 1M 71.1134 Elomzok / Minkama 11°30°-40’ x 4°10°-20° =500 04.X.71 1M,1F 173.244; 73.300 Sadolkoulaye 13°50’-14° x 7°10°-20° = 1200 28.1V.73 3M 77.083-085 Elomzok 11°30°-40° x 4°10°-20° =500 07.X.77 1M 81.067 Belel 14°20°-30° x 7°00°-10° = 1540 16.V.81 1M 81.068 Sadolkoulaye 13°50°-14° x 7°10°-20° = 1200 17.V.81 3M 81.110-112 Ndjaouro Ndekaou 13°00°-10° x 7°20’-30’ 900 20.V.81 IE 82.198 Lipel 12°50’-13° x 6°30’-40’ =900 20.V.82 1M 83.015 Yambassa / Bombato 11°10°-20° x 4°20°-30° =400 29.[X.83 1M 84.001 Ebebda 11°10°-20° x 4°20°-30° 380 21.11.84 1M 86.025 Makouopnsap 11°00°-10° x 5°10°-20° =650 18.IIL.86 Nord 2M 71.940;71.1002 Réserve Bénoué 13°40’-50’ x 8°00°-10° =400 28.VI.71 9M 72.532-540 Badjouma 13°30’-40’ x 9°20°-30° =250 20.VII.72 1M 72.562 Dogba 14°10°-20° x 10°40°-50° = 400 12.VII.72 166 72.546 Koza (Jiller) 13°50’-14° x 10°50’-11°= 450 15.VII.72 MSIE e712 5971-598 Mokolo 13°50’-14° x 10°50°-11°= 700 15.V11.72 1M 73.326 Garoua / Pitoa 13°20’-30° x 9°20’-30’ =180 01.VII.73 SMF ES 379-378 Salak 14°10’-20’ x 10°20’-30’ = 420 03.VII.73 5M 75.364-368 Mora / Waza 14°20’-30’ x 11°10’-20° = 300 14.VII.75 1M 81.107 Karna 13°30’-40’ x 7°40°-50° 500 21.V.81 K. wazae 6M 75.358-363 Mora / Waza 14°20’-30’ x 11°10’-20’ = 300 14.V11.75 K. cassinoides 3M 71.941-942; 71.1003 Réserve Bénoué 13°40’-50’ x 8°00°-10° =400 28.VI.71 4M 73 321-324 Garoua / Pitoa 13°20’-30’ x 9°20’-30’ =180 01.VII.73 1M,1F 73.451-452 Mbé 13°30’-40’ x 7°50’-8° 580 13.VIL.73 2M 75.176-377 Boki 13°30’40’ x 8°40°-50° 320 17.VIL.75 collection de l’auteur, — localité d’origine, telle qu’elle est reportée sur les étiquettes de collection (en général, village proche du point de capture), — coordonnées géo- graphiques de la maille de dix minutes de côté où se situe le point de capture, — altitude, — date de capture. 90 Valeurs de L chez les Phlyctimantis et Kassina du Cameroun. Rectangles: mäles; cercles: femelles. Abréviations: mac: K. maculosa; dec: K. decorata; cassin: K. cassinoides; sen: K. sene- galensis; bou: P. boulengeri, MC: Mont Cameroun: Ad: Adamaoua; C: centre; N: nord; S: sud: J.-L. AMIET 490 480 470 460 450 440 430 420 410 400 390 380 370 360 350 340 330 320 NI 310 R 300 = 290 280 270 260 250 240 230 220 210 200 FIG. 1 W: ouest. PHLYCTIMANTIS ET KASSINA DU CAMEROUN 9] Pour tenir compte de la variation géographique qui affecte la plupart des espèces traitées, les données des tableaux I et II et de la figure 1 sont réparties en grands en- sembles de populations (par exemple, « decorata Adamaoua » ou « senegalensis Sud ») qui sont définis dans les parties relatives à chaque taxon. Le matériel comprend 168 individus adultes. Ce chiffre est relativement faible car, pour diverses raisons, les régions de savanes ont été relativement peu prospectées, alors que les Kassina sont surtout savanicoles. Cet inconvénient devrait être compensé par le fait que tous les spécimens, sauf trois, proviennent de captures personnelles et que l’environnement dans lequel ils ont été trouvés est donc connu avec une précision satisfaisante. Les mensurations ont été faites au pied à coulisse, en dixièmes de mm. La longueur (L) a été mesurée du bout du museau à l’entrejambe (et non à l’anus), l’ani- mal étant maintenu bien à plat (il est souvent incurvé vers le bas). Les autres données morphométriques ont été rapportées à L et exprimées en pourcentage: — largeur de la tête au niveau des tympans (T), — longueur de l’avant-bras, du coude à la base du pouce (AB), — longueur de la cuisse, de l’entrejambe au genou (C), — longueur de la jambe, du genou au talon (J), — longueur du pied, du talon à l’extrémité de l’orteil IV (P). Dans tous les cas où un matériel suffisant était disponible, les valeurs de C et J se sont révélées égales ou, en proportion semblable, légèrement supérieures l’une à l’autre. C’est donc le rapport C+J / L qui a été retenu. Il n’a pas été tenu compte des données morphométriques céphaliques, qui n’ont montré qu’un intérêt diagnostique limité chez les espèces étudiées. Le tableau II synthetise les principaux résultats des 947 mensu- rations effectuées et des 611 rapports calculés. Dans le genre Kassina, certains éléments de la maculation — parfois restés méconnus —, sont très significatifs au plan taxonomique. Une place importante leur sera accordée, aussi bien dans le texte que dans les illustrations. Tous les dessins ont été faits à la chambre claire, les spécimens étant en milieu liquide. Ce dernier point est très important car, à sec, le contour des macules peut être difficile à percevoir; c’est la seule méthode qui permette le « décryptage » de la maculation chez certaines espèces si les spécimens ont été fixés en livrée nocturne, sombre et peu contrastée (voir plus loin). De nombreux dessins de maculation ont été donnés car c’est le meilleur moyen d’en montrer à la fois la variation et les traits fondamentaux. Pour faciliter la comparaison entre les différentes espèces, la figure 1 synthétise toutes les données relatives à la taille (valeurs de L). Dans le même but, les dessins de pieds et de mains ont été regroupés dans la figure 2. Les distributions sont représentées sur des cartes quadrillées à 10 minutes sexa- gésimales. On trouvera plus de précisions sur les méthodes de cartographie utilisées pour l’étude de la batrachofaune camerounaise dans un travail antérieur consacré à ce sujet (Amiet, 1983). L’orthographe des noms de lieux est celle figurant sur les feuilles au 1 / 200 000 de la couverture cartographique du Cameroun. Dans les parties relatives à la distribution et à l’écologie des espèces, les données phytogéographiques et clima- tiques sont empruntées aux travaux fondamentaux de Letouzey (1968, 1985) pour les premières, et de Suchel (1972, 1988) pour les secondes. Les appels d'individus appartenant à tous les taxons ont été enregistrés mais, mon matériel bio-acoustique n’étant pas exploitable pour le moment, les vocalisations 9 J.-L. AMIET K. decorata ta K. maculosa (Adamaoua) Ph. boulengeri K. cassinoides K. senegalensis K. wazae FIG. 2 Face plantaire du pied (tarse non compris) et face palmaire de la main chez les Phlyctimantis et Kassina du Cameroun. Figures 4 la méme échelle, sauf P. boulengeri, réduit de moitié. Remarquer la disposition des orteils V et IV, ainsi que l’absence de tubercules sous-articulaires proximaux sous les mémes orteils, chez K. maculosa et K. decorata. PHLYCTIMANTIS ET KASSINA DU CAMEROUN 93 supra-tympanique loreale wes i LY 4 nasale à oe SEU __— sous-oculaire f posterieure sous-oculaire anterieure Fic. 3 Représentation schématique des macules céphaliques latérales dans le genre Kassina. Voir aussi les Figs. 9, 12 et 13. ne seront pas utilisées dans un but taxonomique. Elles serviront néanmoins pour pré- ciser les périodes d’activité vocale ainsi que les répartitions. Dans ce dernier cas, tous les enregistrements susceptibles de prêter à confusion ont été écartés. Le présent travail n’a pas pour objectif de présenter des monographies des dif- férentes espèces concernées. Pour les mieux connues, les descriptions se limiteront aux points les plus remarquables. Pour alléger le texte et les tableaux, les noms des auteurs et les dates de description des espèces n’ont pas été mentionnés. On les trouvera ci-après, genres et espèces étant classés par ordre alphabétique: Afrixalus fulvovittatus (Cope, 1861 « 1860 »), À. paradorsalis Perret, 1960, À. vittiger (Peters, 1876), A. weidholzi (Mertens, 1938), Amnirana albolabris (Hallowell, 1856), Astylosternus diadematus Werner, 1898, Bufo maculatus Hallowell, 1855 « 1854 », B. regularis Reuss, 1834, B. steindachneri Pfeffer, 1893, B. xeros Tandy, Tandy, Keith & Duff Mac Key, 1976, Chiromantis rufescens (Günther, 1869 « 1868 »), Hemisus guineensis Cope, 1865, Hilde- brandtia ornata (Peters, 1878), Hoplobatrachus occipitalis (Günther, 1859 « 1858 »), Hydro- phylax galamensis (Duméril & Bibron, 1841), Hyperolius adametzi Ahl, 1931, H. adspersus Peters, 1877, H. balfouri viridistriatus Monard, 1951, H. bolifambae Mertens, 1938, H. bopeleti Amiet, 1979, H. cinnamomeoventris Bocage, 1866, H. concolor (Hallowell, 1844), A. igbet- tensis Schigtz, 1863, H. kuligae Mertens, 1940, H. pardalis Laurent, 1947, H. platyceps (Boulenger, 1900), H. sylvaticus nigeriensis Schigtz, 1967, H. tuberculatus (Mocquard, 1897), HA. viridiflavus aureus Perret, 1966, H. viridiflavus pallidus Mertens, 1940, Kassina arboricola Perret, 1985, K. cassinoides (Boulenger, 1903), K. cochranae (Loveridge, 1941), K. decorata (Angel, 1940), K. fusca Schigtz, 1967, K. lamottei Schigtz, 1967, K. maculosa (Sternfeld, 1917), K. schioetzi Rödel et al., 2002, K. senegalensis (Duméril & Bibron, 1841), Leptopelis aubryi (Duméril, 1856), L. calcaratus (Boulenger, 1906), L. nordequatorialis Perret, 1966, L. viridis (Giinther, 1869 « 1868 »), Phlyctimantis boulengeri Perret, 1966, P. leonardi (Boulenger, 1906), Phrynobatrachus auritus Boulenger, 1900, P. natalensis (Smith, 1849), P. cf. werneri (Nieden, 1910), Phrynomantis microps Peters, 1875, Ptychadena aequiplicata (Werner, 1898), P. bibroni (Hallowell, 1845), P. floweri (Boulengeri, 1917), P. oxyrhynchus (Smith, 1849), P. perreti Guibé & Lamotte, 1958, P. pumilio (Boulenger, 1920), P. schubotzi (Sternfeld, 1917), P. stenocephala (Boulenger, 1901), P. straeleni (Inger, 1968), P. trinodis (Boettger, 1881). P. trinodis (Boettger, 1881). REMARQUES SUR QUELQUES CARACTERES D’INTERET TAXONOMIQUE Il est d’abord nécessaire de s’arréter sur certaines particularités plus ou moins négligées dans les descriptions antérieures, et dont il sera fait état 4 plusieurs reprises par la suite. J.-L. 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W 9 IN EI IN ZI AT WTI AI WEI ‘NEI W 9 HTWS HI'WT So I 97 90 G 61 ST A @ 9] 91 IE CE SE PSE SEE 97€ ree ELE TSE Ise Ste SSE GEE ce Tre OTE STE (019 8°87 67€ STE 6I£ CEE OSE EIE dt VE S'9€ 8°6€ Tre 69€ SIE € Ch CBE EHE ELE se CILS Nt Wr YHI’WOI W 9 HE WET ATWLI AS'N9 AI WMEI AI W9 WI AT NS CHA 9°07 el PST 9'TI USE] STE Sl L's EST rl € SI rel SELr SILr esp TOE PIE SSPE 817 Orie €°767 C'ELT L'8TE 8LE (44 097 LIV ETE cs C8T OIT 1270 OST 09T 00€ p9oc ‘Al Cor L8V OOS SCE SVE €07 087 ve CEE SOT OVE 06€ IN Wr AIN OI N 9 HE NET ATWLI AS WIE AI WEI AI W9 WI ATWS dI MC On omy B 2 le) ce = 3 À à E BS = GR ees BS S ® Sì à où vò ons $ = ons = a ae vo S SI = SS SI S E S S SS S S à S 3 S S S $ $ es : i a = = 5 Ca n “od A}-11899 ‘auu9 A0WI ‘ofewruruu IMOLA ‘9JELUIXEUI INIA ‘spansaw (A) SA][OwI9] 39 (W) sPfew suswmpds op 91quiou :seq ua Ney op ‘SINAJUA op sLIgs onbeyo nog *« SOpoygUI Jd [ILIAC] » OA :SUONBIA9IQY ‘9Ipn}o [OLA ne soAne]oi sonbinowoydiow saouuoq ‘7 NVAIAVL PHLYCTIMANTIS ET KASSINA DU CAMEROUN 95 K. maculosa K. senegalensis K. cassinoides Fic. 4 Aspect de la région gulaire chez les males de Phlyctimantis et Kassina. Sac vocal de type 1: P. boulengeri, K. maculosa et K. senegalensis. Sac vocal de type 2: K. cassinoides (voir aussi, pour ce type, la Fig. 13). LIVREES SOMBRES ET LIVREES CLAIRES Chez les Phlyctimantis et Kassina, contrairement aux Hyperolius, il n’y a ni phases ni morphes mais, comme chez eux, il y a une forte variabilité de la livrée en fonction des conditions ambiantes. Bien que ces variations soient, globalement, nycthémérales, on peut de facon plus concrete parler de « livrée sombre » et de « livrée claire ». La premiére est marquée par un assombrissement général de la pigmentation, entrainant une forte atténuation des contrastes; elle se développe la nuit, mais aussi le jour chez des animaux maintenus a l’obscurité et dans une ambiance humide. La livrée claire s’accompagne d’un renforcement du contraste entre les macules et la coloration de fond; ce contraste atteint son maximum si l’animal est soumis ä un fort éclairement, ce qui ne doit pas étre souvent le cas dans les conditions naturelles, la phase de repos 96 J.-L. AMIET ne paraissant pas se dérouler a découvert. Heureusement, si l’observation des macules peut étre difficile en livrée sombre (cf. ci-dessus: « Matériel et méthodes »), ni la forme ni la disposition de celles-ci ne sont affectées. Fait 4 premiere vue inattendu, chez les Kassina et les Phlyctimantis la face ventrale, y compris la région gulaire des miles, s’éclaircit ou s’assombrit en même temps que la face dorsale. Il résulte de ce qui précède que des spécimens de collection pourront donner une impression fort différente suivant qu’ils ont été sacrifiés en livrée diurne ou nocturne et que, en les comparant, il faut toujours avoir ce phénomène présent à l’esprit. MACULATION CEPHALIQUE LATERALE Chez les Kassina, les côtés de la tête portent des macules de forme et de taille variables mais de situation constante: — macules supra-tympaniques, — macules sous- oculaires postérieures, — macules sous-oculaires antérieures, — macules loréales, — macules nasales (Fig. 3). La fusion, la disparition ou l’extension de certaines de ces macules conduisent a des « patterns » spécifiques, fait qui parait avoir échappé jus- qu’ici aux descripteurs (voir Figs. 9, 12 et 13). APPAREIL VOCAL Drewes (1984) a décrit en detail l'appareil vocal des Kassina et genres voisins, et en a montré la profonde originalité. Parmi les espèces représentées au Cameroun, il a étudié essentiellement K. senegalensis. Cette espèce offre en effet un « modèle » largement répandu. Il en existe un autre, un peu différent, propre à deux des espèces de la faune camerounaise. Les caractéristiques principales de ces deux types sont les suivantes (Fig. 4 et Pl. II, Fig. e). — Type 1. La partie médiane épaissie du tégument gulaire, ou glande gulaire, est bien développée, non déprimée ni sollonnée a sa surface, et présente un contour rectangulaire, elliptique ou plus ou moins arrondi. De chaque côté se trouvent des fossettes fortement pigmentées, au fond desquelles se distinguent des replis constitués par un tégument plus mince, non granu- leux, de coloration noirätre. Quand le sac vocal est gonflé, ce tégument se déplisse et dessine deux aires très foncées contrastant avec la couleur plus claire du reste du sac (RISI Fig. d): Chez les especes camerounaises, ce type est représenté chez Phlyctimantis boulengeri, Kassina maculosa, K. decorata et K. senegalensis. — Type 2. La glande gulaire est relativement petite, rétrécie vers l’arriere, plus ou moins concave et/ou sillonnée. Il n’y a pas de dépressions de part et d’autre de la glande gulaire, mais le tégu- ment, épais, forme des plis très contournés, séparés par de profonds sillons. Ce système complexe de plis permet un fort accroissement de la surface du tégument quand le sac vocal se gonfle. Ce type est représenté chez une espéce nouvelle décrite plus loin ainsi que chez K. cassinoides, dont la glande gulaire est particulièrement réduite et mal différenciée. La disparition des aires noires du sac vocal chez ces espéces affaiblit les interprétations (trés anthropocentriques) de la « fonction » de ces macules, évoquées par Drewes (1984). PHLYCTIMANTIS ET KASSINA DU CAMEROUN 97 Dans les deux cas précédents, comme l’a bien vu ce dernier (op. cit.), la partie du tégument affectée par le gonflement du sac vocal s’étend largement vers l’arrière. La limite postérieure de la partie gonflable apparait sous la forme d’un pli ou d’un bourrelet pectoral, allant de la base d’un bras à l’autre. AUTRES CARACTERES SEXUELS SECONDAIRES Sur la face interne de l’avant-bras, les males de Kassina sont munis d’un empä- tement glandulaire, parfois légèrement saillant, qui s’étend presque jusqu’au poignet. Chez les Phlyctimantis cette zone glandulaire est reportée sur les doigts I, II et III. Dans les deux cas, ces formations se reconnaissent par leur absence de pigmentation. Un caractère propre aux femelles des Kassina, et généralement omis par les auteurs, réside dans l’aspect du tégument ventral. Chez toutes les espèces, il est lisse sous la gorge, mais cette texture peut s’étendre plus ou moins loin vers l’arrière: région pectorale chez K. maculosa et K. decorata, deux tiers antérieurs de la région abdo- minale chez K. senegalensis, et presque les trois quarts chez K. cassinoides. Il faut remarquer que ce caractère se retrouve chez les mâles de deux espèces d’Afrique de l’Ouest, K. schioetzi et K. cochranae, dont le centre de la région pectorale est lisse (Rödel er al., 2002). Phlyctimantis boulengeri Perret, 1986 HISTORIQUE Contrairement a l’Afrique de l’Ouest et à l’est de la République démocratique du Congo, les localités camerounaises oü des représentants du genre Phlyctimantis ont été trouvés sont peu nombreuses et dispersées. Cette constatation parait pouvoir étre étendue aux pays limitrophes. Dans une publication de 1978, suite à la redécouverte de P. leonardi a Fainchang, dans la région de Mamfe, j’ai fait le point sur les localités signalées aupa- ravant au Cameroun: plusieurs citations, entre 1911 et 1940, concernent un étroit secteur côtier à la base du versant sud du Mont Cameroun; une autre citation, la pre- miere pour le territoire camerounais (1908), concerne une localité proche de Kribi, Londji, elle aussi en situation littorale. A l’inverse des précédentes, la localité de Fainchang se situe largement à l’intérieur des terres (environ 180 km de la côte), mais a basse altitude, dans un secteur se rattachant, au point de vue climatique et phyto- géographique, a la Plaine littorale. Grâce au petit matériel frais (5 mâles) recueilli à Fainchang, un complément de description a pu étre donné, avec en particulier une comparaison entre les caractéres chromatiques des spécimens camerounais (en livrée diurne et en livrée nocturne) et ceux décrits par Schigtz (1967) a partir de son matériel de Côte d’ Ivoire (Amiet, 1978). Quelques années plus tard, Perret (1986) a publié une mise au point sur le genre Phlyctimantis dans laquelle il établit que les syntypes ayant servi a la description origi- nale de P. leonardi par Boulenger (1906) forment une série heterogene. S’appuyant sur un matériel de provenances diverses (de la Côte d’Ivoire au Gabon), il scinde l’espèce de Boulenger en deux: — l’une, P. leonardi, définie d’après du matériel gabonais et sup- posée exister jusqu’en RDC, — l’autre, P. boulengeri n. sp., avec un des spécimens de Fainchang comme holotype et répandue depuis le Cameroun jusqu’a la Cöte d’Ivoire. Les descriptions de Perret font ressortir deux caractéres discriminants principaux: — la 98 J.-L. AMIET taille, de 1 cm en moyenne plus grande chez leonardi que chez boulengeri, — une « peau dorsale finement verruqueuse » chez boulengeri, alors que le tégument dorsal est « lisse ou très finement chagriné » chez leonardi; accessoirement, P. boulengeri est distingué aussi par sa maculation jaune plus apparente et son tympan mieux marqué. En 1987, j'ai trouvé un site de reproduction de Phlyctimantis a Tom, village proche de la frontiere équato-guinéenne. Cette localité a le grand intérét d’étre plus proche du Gabon que les localités connues jusqu’alors: la terra typica de P. leonardi, Ndjolé, est à environ 260 km à vol d’oiseau, soit une centaine de km de moins que Fainchang. Une autre station de Phlyctimantis a été trouvée en 1991 par Ch. P. Blanc dans la Plaine littorale, au NE de Campo, a une cinquantaine de km au sud de l’ancienne localité de Londji. La présence de Phlyctimantis pres de la frontiere sud du Cameroun pouvait laisser supposer que les deux espèces reconnues par Perret seraient présentes sur le territoire. Comme on va le voir, la comparaison des spécimens de Fainchang et de Tom ne confirme pas cette hypothèse. COMPARAISON DES SPÉCIMENS DE FAINCHANG ET DE TOM Le matériel utilisé est modeste, 4 mâles pour chacune des localités, mais 1°) ces individus ont été capturés en pleine activité vocale et ont donc des caractères sexuels secondaires pleinement développés, 2°) ils constituent des échantillons homogènes car, dans chaque localité, ils ont été pris au même endroit et la même nuit. Dans ce qui suit, l’accent sera mis sur les caractères qui, d’après Perret (op. cit.), permettent de séparer boulengeri de leonardi. Taille, proportions, habitus: comme le montrent la figure 1 et le tableau II, il n’y a pas de différence de taille notable entre les individus des deux échantillons. Il en est de même pour les proportions des différents segments des membres. La morphologie céphalique, la palmure, les disques digitaux ne fournissent pas non plus de caractères distinctifs. Pourtant, les individus de Fainchang ont un aspect plus robuste, alors que ceux de Tom paraissent plus graciles, mais il est difficile d’accorder à cette différence d’apparence une valeur discriminante de niveau spécifique. Tegument: dans ma description des spécimens de Fainchang j’ai, pour la pre- miere fois, signalé l’existence d’un relief tégumentaire: « Le tégument dorsal est apparemment lisse, garni en fait d’infimes verrucosités éparses ». Le terme « verru- cosités » me parait a present imprécis. Le tégument dorsal forme de petites excrois- sances mousses, arrondies ou plus ou moins étirées, irregulierement dispersées et peu apparentes (elles ne se voient plus sur les spécimens fixés), qui peuvent effectivement être qualifiées de verrues. Mais, en plus de ces petites verrues, parfois sur elles mais aussi entre elles, il y a d’infimes spinules coniques, cornées, blanchätres, telles qu’on peut en observer par exemple chez les Hyperolius du groupe d’H. nasutus (Amiet, 2005). Ces aspérités sont présentes sur toute la face supérieure du corps, de la tête et des membres, et méme, encore plus petites, sur les reliefs du tégument gulaire. Chez les spécimens de Tom, le tégument dorsal parait plus lisse, les petites verrues se localisant sur le dessus des membres. Les aspérités cornées, en revanche, sont aussi nombreuses que chez les individus de Fainchang. PHLYCTIMANTIS ET KASSINA DU CAMEROUN 99 Dents vomeriennes: elles sont bien développées (encore plus que chez Kassina cassinoides) chez les individus des deux localites. Caractères sexuels secondaires: (1) La glande gulaire, assez petite (de l’ordre de 6 à 8 mm de large), a un contour pentagonal, hexagonal ou subcirculaire; postérieu- rement, elle forme un repli qui paraît libre mais est en fait relié par un raphé au reste du tégument gulaire. Le disque est flanqué de deux dépressions où apparaissent, sous forme de gros plis, les aires noires rétractées du sac vocal (et non « les sacs vocaux », comme je l’ai écrit par erreur dans mon article de 1978). Chez certains individus, il y a un pli pectoral assez peu marqué (Fig. 4). (2) Contrairement aux Kassina, les aires glandulaires des membres antérieurs ne sont pas situées sur les avant-bras mais sur le face supérieure des doigts I, II et, de façon plus limitée, III. (3) Faute de spécimens femelles, il n’est pas possible de dire si les petites aspérités cornées tégumentaires sont propres aux mâles, mais cela est probable. Cette hypothèse est confortée par le fait qu’elles manquent chez trois juvéniles de Tom, et qu’elles sont peu développées chez le plus petit des mâles adultes de la même localité. Aucun des caractères précédents ne diffère de façon notable entre les individus des deux localités. Pigmentation: à première vue, les spécimens de Fainchang s’opposent par leur teinte très foncée, sur les faces dorsale et ventrale, à ceux de Tom, beaucoup plus clairs. Quoique très apparente, cette différence n’a pas de valeur discriminante : elle est due seulement au fait que les premiers ont été fixés alors qu’ils étaient en livrée nocturne, alors que les seconds l’ont été en livrée diurne. En réalité, il n’y a pas de différence substantielle entre la coloration des individus de Fainchang et de Tom. Dans les deux cas, il n’y a plus aucune trace d’une maculation dorsale « structurée », mais simple- ment quelques petites taches foncées irrégulièrement dispersées et, chez certains indi- vidus, des « ombres » correspondant à des macules loréales et supra-tympaniques. Les bandes jaunes et noires alternées des membres postérieurs montrent, dans les deux échantillons, une variabilité individuelle qui empêche de les utiliser comme caractères distinctifs. DISTRIBUTION, ÉCOLOGIE Sur la carte Fig. 5 ont été reportées les localités camerounaises, anciennes et récentes, où des Phlyctimantis ont été signalés; de plus, toutes les mailles ou j’ai relevé au moins cinq espèces d’Anoures (souvent beaucoup plus, jusqu’à la cinquantaine) ont été barrées d’une ligne diagonale, ce qui permet de mieux apprécier la distribution en fonction de la « pression d’investigation » sur les différentes régions du territoire. Deux faits en ressortent: — 1°) la grande dispersion des sites peuplés par des Phlyctimantis, qui ne paraît pas pouvoir être imputée seulement à une insuffisance des prospections; — 2°) leur localisation à basse altitude, dans la Plaine littorale ou à ses abords immé- diats, contrastant avec leur absence sur le Plateau sud-camerounais (à peu près cir- conscrit par la courbe de niveau de 600 m). Les localités ou la présence de Phlyctimantis a été relevée ont en commun: — une température moyenne annuelle élevée, de l’ordre de 24° a 26°5 C, en relation avec leur faible altitude; — une importante quantité de précipitations, de 2 m par an à 9 m (au pied du Mt Cameroun); — une végétation climacique correspondant a la forét ombrophile de type atlantique (ou biafréen). Ces facteurs peuvent conditionner l’aire d’ensemble sur le territoire, mais n’expliquent pas sa discontinuité. 100 J.-L. AMIET Ane Fic. 5 Carte du Cameroun au sud du 8ème parallèle montrant les localités où P. boulengeri a été observé. 1: Isongo et Mubenge (= Mowange, = Mubange); 2: Fainchang (= Fineschang, = Fainschang); 3: Londji (= Longji); 4: Campo; 5: Tom. En 6 a été ajoutée la localité équato-guinéenne de Punta Frailes (= Punta Europa). Pour 1, 3 et 6, les récolteurs et auteurs sont mentionnés dans Amiet, 1978. Dans la localité de Fainchang, le 14 avril 1978, une douzaine de mâles étaient actifs dans une raphiale inondée après de fortes pluies. Ils étaient accompagnés de cen- taines d’H. bolifambae, de nombreux H. concolor et Afrixalus paradorsalis et de quelques Hyperolius sylvaticus nigeriensis. Le site de reproduction de Tom était dif- férent: un grand étang découvert, entouré de buissons sur lesquels étaient perchés les mâles, estimés à une cinquantaine. Bien que la station ait été visitée au tout début de janvier, donc en saison sèche, l’activité vocale était intense et il n’a pas été possible de faire un relevé des espèces compagnes, difficiles à repérer dans la « masse sonore » produite par les Phlyctimantis. Dans les environs proches pouvaient être entendus Afrixalus fulvovittatus, Hyperolius platyceps, H. pardalis et H. tuberculatus. Dans les deux stations, la nature des sites de reproduction et les espèces compagnes témoignent du caractère parasylvicole des Phlyctimantis, ce qui confirme les observations de Largen & Dowsett-Lemaire (1991) au Congo et de Rödel & Ernst (2001) en Côte d'Ivoire. Il est cependant surprenant que, contrairement aux autres espèces parasylvi- coles, très « expansionnistes », les populations de Phlyctimantis restent aussi localisées PHLYCTIMANTIS ET KASSINA DU CAMEROUN 101 et disséminées!. Cette situation est d’autant plus paradoxale que — 1°) l’aire de répar- tition du couple P. leonardi — P. boulengeri est immense, allant du centre de la cuvette congolaise au Libéria; — 2°) dans leurs rares stations camerounaises, les populations paraissent florissantes à en juger par le nombre de mâles actifs. Pour parachever ce tableau, il faut souligner que les sites de reproduction ne montrent aucune particularité notable: dans la région de Fainchang, une dizaine de formations marécageuses par- tageant les mémes caractéristiques et présentant des peuplements identiques ont été prospectées, mais une seule hebergeait des Phlyctimantis. DISCUSSION La comparaison qui précède a montré qu’il n’y a pas de différence notable entre les mâles des populations de Fainchang et de Tom. Leur taille relativement faible et leurs aspérités tégumentaires conduisent, si l’on suit les critères distinctifs définis par Perret (1986), à les placer sous le nom de P. boulengeri. Cependant, cette attribution n’est pas entièrement satisfaisante pour les raisons suivantes. 1°) A Tchissanga, dans le bassin du Kouilou (Congo), Largen et Dowsett- Lemaire ont trouvé une importante population d’un Phlyctimantis qu’ils ont rapporté à P. leonardi. Ils indiquent toutefois que « The relative small size of these males (snout- vent length 45-50 mm) does, however, cast some doubt on Perret’s contention that leonardi is a significantly larger species than boulengeri » (Largen & Dowsett- Lemaire, 1991). De plus, comme me l’a indiqué M. Largen (comm. épist.), les 8 mâles de cette localité « have the dorsum densely beset with minute asperities... ». 2°) Cet autre critère distinctif que constituent les aspérités tégumentaires devrait de toute façon être précisé. Il faudrait en particulier savoir si elles existent chez les femelles de P. boulengeri, ce qui n’est pas explicitement indiqué dans la description. Au cas où il s’agirait d’un caractère propre aux mâles, ne disparait-il pas en dehors des périodes de reproduction ? Sans réponse a ces questions, le critére tégumentaire n apparait pas totalement fiable. 3°) Entre les localités de Tom (P. boulengeri) et de Ndjolé (terra typica de P. leonardi), il n’y a pas d’obstacle géographique ou écologique majeur, ni de limite biogéographique reconnue, qui puissent expliquer le remplacement d’une espèce par une autre. Il est possible que des localités intermédiaires abritent des populations de Phlyctimantis et des recherches complémentaires seraient nécessaires pour comprendre les rapports spatiaux entre P. leonardi et P. boulengeri dans l’intervalle d’environ 260 km qui sépare Tom de Ndjolé. 4°) Les grands mâles de Bélinga (Gabon) sur lesquels Perret a fondé sa redescription de P. leonardi proviennent d’une localité plus « continentale » que celles de Tom, Ndjolé et Tchissanga. Au Congo, P. leonardi a été cité de la Sangha (Guibé, 1946, in Largen & Dowsett-Lemaire, 1991). En RDC, la même espèce, d’après Laurent (1976), s’avance jusqu’au cœur de la cuvette congolaise. On peut se demander si ces ! Un cas un peu similaire est celui d’Hyperolius bopeleti, qui paraît propre aux formations forestières dégradées ou à couvert discontinu et ne peuple pourtant qu’une étroite bande côtière: cf. Amiet (1979). 102 J.-L. AMIET populations sont bien conspécifiques des populations établies dans les régions plus littorales. Il subsiste donc un certain nombre d’incertitudes et d’ interrogations au sujet des Phlyctimantis d’ Afrique centrale. Elles confèrent un caractère provisoire à |’ attribution des populations camerounaises a P. boulengeri. Le probleme de Kassina maculosa (Sternfeld, 1917) et de X. decorata (Angel, 1940) HISTORIQUE K. maculosa a été décrite en 1917 par Sternfeld, sous le nom de Megalixalus maculosus, sur deux spécimens provenant de l’Ubangi, dans l’actuelle République démocratique du Congo. En 1940, Angel a décrit, sur une unique femelle provenant du cratere du Mt Manengouba, au Cameroun, un Megalixalus decoratus dont Laurent & Combaz (1950), après examen du type, ont montré l’appartenance au genre Kassina. Ni Angel ni Laurent et Combaz n’ont tenté un rapprochement entre les deux espèces?. Perret (1966) semble être le premier a l’avoir envisagé. Pour designer ses spécimens du Cameroun, il utilise le nom « decorata » et écrit: « Ne pouvant apporter la preuve de la synonymie de decorata avec maculosa (Sternfeld) je conserve pour l’instant l’espece d’ ANGEL du Cameroun mais je soupçonne l’existence d’une seule espèce 2. Peu après, Schigtz (1967), évoquant K. decorata et K. maculosa, mentionne que « Perret (in litt.) has informed me that he regards the two last mentionned species as only subspecifically distinct from each other ». Dans son travail sur les rainettes d’Afrique (1999), il reprend cette opinion: « there do not seem to be any convincing arguments for separating K. decorata and K. maculosa at the specific level... ». Dans leur révision très documentée des « spotted Kassina » d’Afrique de l’Ouest, Rödel, Grafe, Rudolf & Ernst (2002) ont été amenés a discuter le cas de ces espèces, apparentées aux trois espèces qu'ils reconnaissent dans leur territoire d’étude. Ils écrivent que « According to Perret (1966) and Schigtz (1999) K. decorata (Angel, 1940) is a synonym of K. maculosa (Sternfeld, 1917) », substituant ainsi une forme affirmative à la tournure dubitative des auteurs cités. Néanmoins, ils indiquent ensuite qu'ils utilisent les deux noms « because they have been used in the literature, and because our acoustic analyses showed remarkable variation ». En fait, dans la suite du travail, il n’y a pas de comparaison entre les appels des deux espèces. Les auteurs donnent les résultats de l’analyse des appels de neuf K. decorata du Cameroun (enregistrés par Françoise Dowsett-Lemaire) et relèvent que, sur plusieurs points, ils different de ceux des trois espèces faisant l’objet de leur révision; en ce qui concerne K. maculosa, ils évoquent seulement le sonagramme d’un individu du Cameroun publié par Schigtz (1999), en mentionnant qu’il ressemble à l’appel de K. decorata mais que, faute d’échelle sur l’axe y, une comparaison avec leurs analyses n’est pas possible. En ce qui concerne l’origine des données bio-acoustiques relatives à K. macu- losa et K. decorata, les précisions suivantes doivent être apportées. 1°) Le sonagramme 2 Laurent et Combaz citent « M. maculatus Sternfeld » pour préciser que cette espèce a été « reléguée dans la synonymie de K. senegalensis » par Mertens (1941). On peut supposer que « maculatus » correspond ici à un lapsus pour « maculosus ». PHLYCTIMANTIS ET KASSINA DU CAMEROUN 103 de K. maculosa publié par Schigtz (1999) a été obtenu a partir d’un de mes enre- gistrements, réalisé a Obala, au nord de Yaoundé. 2°) Les enregistrements de K. deco- rata analysés par Rödel er al. (2002) ont été effectués par Françoise Dowsett-Lemaire pres du Lac Edib, dans l’ouest du Cameroun, et attribués a K. decorata par C. Wilde (F. Dowsett-Lemaire, comm. pers.). Il faut relever que, à l’époque ot ont été faites ces identifications, une forte incertitude régnait quant a la séparation des taxons et a leur dénomination. Elles sont cependant probablement valables car, après avoir procédé a |’ étude de mon matériel préservé, il m’apparait qu’il y a au Cameroun deux entités distinctes, reconnaissables uniquement par leur maculation: chez l’une, celle-ci correspond à ce que Angel a décrit chez K. decorata, alors que chez l’autre elle correspond à la description de K. maculosa. Il a été impossible de trouver d’autres caractères morpho- logiques nets pour les séparer (les appels n’ont pas paru discriminants sur le terrain). Dans ce qui suit, avant d’en discuter le statut taxonomique, ces Kassina seront désignées comme « Kassina type decorata » et « Kassina type maculosa ». DESCRIPTION Caractères communs aux deux types: tégument dorsal lisse sur la tête et une partie de la région médio-dorsale, avec des granulations, souvent étirées longitudinale- ment, à l’arrière et sur les côtés du dos ; ces excroissances, peu denses et peu saillantes, sont plus ou moins marquées, peut-être en fonction des conditions ambiantes lors de la fixation (et peut-être aussi du mode de préservation). Tégument de la région loréale, des côtés de la tête et des flancs sub-lisse, passant par de fines granulations au tégu- ment ventral. Celui-ci fortement réticulé-granuleux, les granulations de contour poly- gonal et en général plus longues que larges; pas de zone lisse médio-pectorale (différence avec K. cochranae). Chez les femelles, la gorge et la région pectorale sont lisses. Tégument des membres lisse, sauf sous les cuisses, où il est granuleux. Membres antérieurs remarquablement grêles et longs (plus que chez les autres Kassina et, de façon générale, que chez les autres Hypéroliens); doigts minces et déliés, terminés par des pelotes non dilatées ou dilatées, mais en ce cas plus longues que larges et sans sillon périphérique; tubercules sous-articulaires très saillants; des excroissances surnuméraires sous les doigts; aucune palmure interdigitale (Fig. 2). Pied (Fig. 2) long et étroit, avec l’orteil V étroitement accolé à l’orteil IV, jusqu’à hauteur du tubercule sous-articulaire distal; un rudiment de palmure entre les orteils III et IV3; pelotes adhésives non ou modérément dilatées aux orteils IV et V, sans sillon périphérique; tubercules sous-articulaires bien saillants mais, sauf excep- tion, pas de tubercule proximal aux orteils V et IV, et donc un seul tubercule visible à l’orteil V et deux à l’orteil IV. La disparition des tubercules sous-articulaires basaux et la fusion partielle des deux orteils externes ne semblent pas avoir été signalés jusqu'ici chez ces Kassina. Entre les tubercules sous-articulaires, de longues excroissances d’aspect similaire. Sac vocal des mâles de type 1: épaississement médian (glande gulaire) lisse, plus long que large, à bords rectilignes et parallèles; de part et d’autre, fossettes de rétraction du sac bien apparentes, pigmentées de noir; limite postérieure de la partie 3 C’est apparemment par erreur que Sternfeld (917) écrit « Zehen mit halber Schwimmhaut » (« orteils palmés jusqu’à mi-hauteur »), ce qui avait déjà intrigué Mertens (1940). 104 J.-L. AMIET Fic. 6 Maculation du dos, du bras et de la cuisse chez K. maculosa. Dessins a la chambre clair limités au contour pour les macules dorsales. Les numéros de collection permettent de retrouver l’ori- gine géographique des spécimens en se reportant au tableau 1. PHLYCTIMANTIS ET KASSINA DU CAMEROUN 105 FIG. 7 Maculation du dos, du bras et de la cuisse chez K. decorata. Dessins a la chambre claire limités au contour pour les macules dorsales. Les spécimens des rangées supérieure et moyenne pro- viennent de la Plaine des Mbos (Santchou, cf. Tab. 1), ceux de la rangée inférieure sont des juvéniles du cratére du Manengouba (non référencés dans le Tab. 1). gonflable généralement marquée par un pli interbrachial. Face interne des avant-bras des mâles avec une zone glandulaire étirée, légèrement saillante. Livrée dorsale constituée de macules foncées sur un fond plus clair, les macules étant toujours entourées d’un fin liséré jaunâtre ou blanchätre. Il y a une forte difference entre la livrée sombre, de tonalité générale brun-noirätre, où les macules et 106 J.-L. AMIET 71.1011 73.399 FIG. 8 Maculation du dos, du bras et de la cuisse chez K. decorata, populations de |’ Adamaoua (voir Tab. 1 pour les localités et dates de capture des spécimens). Dessins à la chambre claire limités au contour pour les macules dorsales. leur liséré sont à peine discernables, et la livrée claire, montrant un fort contraste entre les macules et le fond, le liséré très apparent donnant alors une impression de taches ocellées. Cette différence entre les deux livrées est beaucoup plus marquée que chez les autres Kassina locales (voir plus haut pour les conditions d’observation de la maculation sur des spécimens à livrée sombre). Chez les deux types, toutes les macules céphaliques latérales sont présentes, les sous-oculaires non fusionnées, la nasale et la loréale fusionnées ou non (Fig. 9). Pas de taches paires occipitales (différence avec K. arboricola). PHLYCTIMANTIS ET KASSINA DU CAMEROUN 107 Face ventrale en général pigmentée seulement sous la gorge des mâles, sous la forme d’un assombrissement presque uniforme ou de mouchetures plus ou moins concentrées en macules diffuses. Le reste de la face ventrale, clair (blanchatre en alcool), peut étre voilé, en livrée sombre, par un fin réseau de chromatophores enserrant les granulations tégumentaires. Membres plus ou moins régulièrement barrés de sombre sur fond clair. Les régions inguinales et axillaires et les parties des membres postérieurs non visibles lorsque ceux-ci sont repliés sont marbrées de jaune vif et de noir sur le vivant, de blanc et de noir sur les spécimens préservés en alcool. Livrée de type maculosa: il peut y avoir, dans la région occipitale, une macule étirée longitudinalement, mais la majorité des macules ont un contour plutöt elliptique et sont moins de deux fois plus longues que larges. Leur nombre et leur taille varient beaucoup, y compris à l’intérieur d’une même population et il est difficile d’y déceler une répartition en rangées longitudinales (Fig. 6 et PI. I, Fig. b). La macule supra- tympanique ne se prolonge pas en arrière du tympan (Fig. 9). Sauf quelques exceptions, la maculation des cuisses est constituée de bandes transverses foncées alternant avec des bandes claires. Livrée de type decorata: plusieurs macules dorsales sont très étirées longitudi- nalement. Elles ont tendance à former des alignements: deux latéro-dorsaux et une bande médio-dorsale qui, assez souvent, se dédouble dans sa moitié postérieure (Fig. 7 et 8 et PI. I, Figs. a, c et d). Dans les cas extrêmes, la livrée peut évoquer celle de Kassina « à bandes » comme K. senegalensis. La maculation céphalique offre un bon caractère distinctif: en général, la macule supra-tympanique se poursuit vers le bas en contournant le tympan (Fig. 9). La maculation des cuisses est souvent irrégulière et tend à se disposer en marbrures étirées longitudinalement. DISTRIBUTION, VARIABILITE Kassina type maculosa: son aire camerounaise prolonge vers l’ouest celle qu’elle occupe dans le nord de la RDC, d’où proviennent les types de K. maculosa (cf. carte in Schigtz, 1999). Dans l’état actuel des connaissances, elle atteint a l’ouest du Cameroun les régions de Dschang et de Kumbo (si, comme il est probable, les trois males de Kakar signalés par Mertens, 1940, se rapportent a la méme forme) mais ne pénetre pas au Nigeria. Comme, sur ce territoire, aucune Kassina similaire n’est connue, il y a entre les populations camerounaises les plus occidentales et les espéces apparentées d’Afrique de l’Ouest, une énorme lacune de distribution, de l’ordre de 1200 km, difficilement explicable. La limite nord de répartition au Cameroun coincide probablement avec la retombée septentrionale du Plateau de 1’ Adamaoua, tandis que vers le sud elle colonise les régions les plus « secondarisées » de la zone forestiére (voir ci-après). L’absence de pointages dans un vaste secteur du Cameroun, en gros la moitié est du Plateau sud-camerounais, est sûrement imputable à une insuffisance des prospections. Dans son aire camerounaise, l’espèce varie peu, sauf en ce qui concerne la taille, apparemment plus faible dans les populations de I’Adamaoua. Une exception doit être faite pour les populations du Mont Cameroun. L'existence d’une Kassina y est 108 J.-L. AMIET 81.148 71.1068 K. maculosa K. decorata Adamaoua Fic. 9 Exemples de maculation céphalique laterale chez K. maculosa et K. decorata. Remarquer, chez cette derniere, la forme caractéristique de la macule supra-tympanique. connue depuis longtemps car Köhler en avait obtenu une femelle et un juvénile au- dessus de Mubenge, versant sud, vers 920 m (spécimens d’abord identifiés comme K. senegalensis par Mertens, 1940). La femelle et les deux mâles que j’ai trouvés a Bonianango, versant est, vers 620 m, proviennent de la seule petite population repérée, malgré plus d’une vingtaine de sorties de nuit, sur le pourtour du Mont Cameroun. Ces spécimens se distinguent nettement par: — leur grande taille (Tab. 2 et Fig. 1), — leur habitus élancé (mais les dessins de la Fig. 6 représentent des individus peut-être émaciés après une période de captivité), — l’élongation des pieds (valeurs de P/L supérieures à celles de toutes les autres Kassina mesurées, et dépassées seulement par un spécimen de Phlyctimantis!), — la pigmentation plus dense de la gorge (grosses taches ocellées chez les mâles, quelques macules arrondies chez la femelle), — la coloration plus vive du pied (Pl. I, Fig. b). Ces individus font penser à K. arboricola (cf. photos in Perret, 1985, et Rödel er al., 2002) mais s’en distinguent par la pigmen- tation foncée moins étendue sur la face ventrale des mâles et, surtout, par l’absence de la double tache occipitale caractéristique de cette espèce. PHLYCTIMANTIS ET KASSINA DU CAMEROUN 109 Kassina type decorata: elle n’est connue que de deux secteurs distants d’environ 360 km (Fig. 10): - Mont Manengouba et Plaine des Mbos, a l’ouest du Cameroun, — region de Ngaoundéré, sur le Plateau de 1’ Adamaoua. Il est possible que son aire de répartition soit à la fois plus étendue et moins largement bipolaire, mais on peut d’ores et déjà admettre qu’elle est beaucoup plus restreinte que celle de Kassina type macu- losa. Les caractéristiques morphologiques des populations du Mont Manengouba et de la Plaine des Mbos, spatialement très proches et même peut-être en continuité, sont semblables. En revanche, les populations de l’Adamaoua sont nettement différentes par : — leur nanisme (chez le plus grand individu de l’Adamaoua, la taille est inférieure à celle du plus petit individu de l’Ouest: Fig. 1 et Tab. 2), — leurs membres postérieurs proportionnellement plus courts (Tab. 2), — les pelotes des doigts et des orteils non dilatées. Les deux premiers caractères se retrouvent, mais nettement moins marqués, chez les Kassina de type maculosa de l’Adamaoua. ÉCOLOGIE, CYCLE D'ACTIVITÉ VOCALE, PHONOCENOSES Kassina type maculosa: son habitat optimal paraît correspondre aux savanes humides de type guinéen, parsemées de buissons et de boqueteaux et parcourues de galeries forestières. Elle peut y être localement abondante, comme l’ont montré les prospections dans le secteur Obala - Bafia, au nord et au nord-ouest de Yaoundé. Sur l’Adamaoua et sur les reliefs de la Dorsale camerounaise, où la végétation herbacée est encore plus largement dominante (savanes soudano-guinéennes plus ou moins arborées de l’Adamaoua, secteurs densément cultivés et déboisés des pays bamoun et bamiléké), Kassina type maculosa peut atteindre environ 1500 m d’altitude, voire 2000 m si les spécimens de Kakar se rapportent à ce type. Elle peut aussi, plus localement, se comporter en espèce parasylvicole. Elle pénètre en effet dans la zone forestière, où elle « descend » plus ou moins vers le sud comme le font d’autres espèces primitivement savanicoles telles que Bufo regularis ou Hoplobatrachus occipitalis. Dans ses stations intra-forestières, elle n’occupe que des zones à végétation très « secondarisée » et, à en juger par le faible nombre de mâles actifs, elle n’y est représentée que par de petites populations dispersées. Sur les sites de reproduction, les mâles peuvent émettre leurs appels depuis des buissons, où ils sont capables de s’élever jusqu’à environ 1,5 m en « empoignant » les rameaux, mais aussi depuis des touffes de graminées; dans ce dernier cas, ils peuvent grimper sur les chaumes et les feuilles, ou simplement se poster sur la partie supérieure des touradons. L'activité vocale des mâles débute aux premières pluies, quand les collections d’eau commencent à se remplir. Suivant leur situation géographique, les populations sont soumises à un climat à une ou deux saisons pluvieuses. Le premier cas correspond, en gros, à l’Adamaoua et à la plus grande partie de la Dorsale camerounaise, alors que le second correspond au Plateau sud-camerounais: il y a alors deux périodes d’activité vocale, l’une en fin mars - avril, l’autre en septembre - octobre. La comparaison de relevés effectués dans 11 sites à végétation graminéenne dominante (région d’Obala, Plateau bamiléké et Adamaoua) et dans 5 sites à végétation secondaire de la zone forestière (région de Yaoundé - Mbalmayo) donne un aperçu des phonocénoses auxquelles participe K. type maculosa. Dans le premier cas, les espèces 110 J.-L. AMIET PSE SECTE TEE SATTE / K. maculosa DE la I 4 Sen nee m7 LS BA OZ Sem AL TY N K. decorata È RI Dar i 7 lateau de l'A HF P / Fic. 10 Carte du Cameroun au sud du 8ème parallèle montrant la répartition des mailles où ont été repérées K. maculosa et K. decorata. La limite forêt — savane est symbolisée par la ligne avec triangles (pointés vers la savane). Les chiffres correspondent aux localités de Perret (1966): Bangwa (1) et Sangmélima (2). vocalement actives auxquelles elle est le plus souvent associée (40% des sites ou plus) sont: Afrixalus fulvovittatus (100% des sites), Hyperolius balfouri viridistriatus (91%), Kassina senegalensis (63,6%), Phrynobatrachus natalensis, Hoplobatrachus occipi- talis, Hyperolius tuberculatus (54,5%), Leptopelis nordequatorialis, Hyperolius igbet- tensis, Bufo regularis, B. maculatus (45,5%). Cette combinaison d’espèces est tout à fait caractéristique des savanes humides guinéennes et soudano-guinéennes. Dans les sites intra-forestiers, les espèces-compagnes les plus fréquentes constituent un bon échantillonnage du peuplement parasylvicole: Afrixalus fulvovittatus (100% des sites), Hyperolius tuberculatus, H. adspersus, Amnirana albolabris, Ptychadena perreti (60%), Bufo maculatus, Chiromantis rufescens, Hyperolius cinnamomeoventris, H. platyceps, H. pardalis, Afrixalus paradorsalis (40%). Les individus trouvés à la base du versant est du mont Cameroun partageaient les abords d’un ruisseau coupé par un barrage avec un mélange d’espèces comprenant des parasylvicoles typiques (Leptopelis aubryi, Afrixalus paradorsalis, Hyperolius concolor, H. kuligae, H. bolifambae) mais aussi des sylvicoles qui, dans cette région très arrosée, subsistent dans les brousses luxuriantes remplaçant la forêt (Astylosternus diadematus, Leptopelis calcaratus, Ptychadena aequiplicata, Phrynobatrachus auritus). PHLYCTIMANTIS ET KASSINA DU CAMEROUN 111 Kassina type decorata: les trois secteurs où ont été trouvées des Kassina de type decorata offrent des conditions d’habitat différentes et seront donc traités séparément — Mont Manengouba: c’est un volcan inactif dont le fond est une plaine her- beuse, large de 3 km dans son plus grand diamètre et située à environ 2000 m d’alti- tude. Les points d’eau sont constitués par deux lacs occupant des cratéres secondaires et par une zone marécageuse pouvant se transformer en étang au plus fort de la saison pluvieuse. A l’exception des parois rocheuses, boisées, les pentes environnantes sont couvertes de prairies paturées par des troupeaux de zébus et des chevaux. Je n’ai pas pu faire de sorties de nuit dans ce site mais trois sorties diurnes (29.III.72, 17-19.1V.76 et 22-23 III.81) m'ont permis d’y trouver: Bufo sp. (apparenté a B. villiersi?), Amni- rana albolabris (tétards de grande taille, à l’époque péchés pour la consommation locale!), Phrynobatrachus cf. werneri, Hyperolius adametzi et, a chacune des sorties, des Kassina, en général juvéniles, abritées sous des pierres. A partir des points d’eau du fond du cratére, ces Kassina peuvent s’élever sur les pentes dominantes, au moins jusque vers 2200 m d’altitude. — Plaine des Mbos: il s’agit d’un fossé d’effondrement, formant une vaste plaine à fond plat entre le Plateau bamiléké au nord et le mont Manengouba au sud. Son alti- tude est de 700 à 720 m. La végétation est constituée par une mosaïque complexe de savanes humides et de massifs boisés, a caractére souvent marécageux, et de cultures vivrieres. La batrachofaune est peu diversifiée, constituée pour l’essentiel de quelques espèces euryéciques et parasylvicoles, qui peuvent être représentées par des popu- lations extrêmement abondantes. Dans la nuit du 24 mai 1971, des dizaines de mâles formaient un chœur continu dans une prairie inondée. Beaucoup chantaient en partie immergés, prenant appui sur des mottes de terre ou des débris végétaux (comportement qui n’a Jamais été observé chez K. type maculosa). Quatre ans plus tard, à un jour près (25.V.75), seuls quelques appels ont été entendus. — Adamaoua oriental: l’essentiel du matériel recueilli dans l’est de I’ Adamaoua provient de la localité de Sadolkoulaye, située à 35 km à l’est de Ngaoundéré. Le site a été visité, de nuit, les 29.VI.71,5.VII.71,28.IV.73 et 17.V.81. Sur un substrat cuirassé absolument plan est établie une prairie qui, en saison des pluies, se transforme en un petit lac herbeux peu profond. Des milliers de Batraciens s’y reproduisent et, au plus fort de la période d’activité vocale (mai-juin) y créent un vacarme assourdissant. Les Kassina sont représentées par K. senegalensis (grande forme de |’ Adamaoua, voir ci- après) et par une Kassina naine à pattern decorata. Cette dernière, extrêmement abon- dante, peut faire des chœurs ininterrompus, comme dans la Plaine des Mbos. Les especes participant a la phonocénose sont Hoplobatrachus occipitalis, Ptychadena pumilio, P. straeleni, Hyperolius balfouri viridistriatus (lui aussi de petite taille), H. igbettensis et Afrixalus fulvovittatus. De vastes prairies marécageuses d’aspect semblable existent plus prés de la ville de Ngaoundéré. Au sud de celle-ci, près de Wakwa, la même Kassina naine a été trouvée, en compagnie ou a proximité de Ptychadena pumilio, Hyperolius b. viridi- striatus, H. viridiflavus aureus, Afrixalus fulvovittatus et A. weidholzi. 112 J.-L. AMIET DISCUSSION Il faut d’abord relever que la livrée de type maculosa correspond bien à la figure que donne Sternfeld (1917) dans sa description de Megalixalus maculosus, de méme qu’il y a une parfaite concordance entre la livrée de type decorata et la description trés précise faite par Angel (1940) de la livrée de son Megalixalus decoratus. Sans se référer directement a la description d’Angel, mais a celle de Perret (1966), Mertens (1968) évoque aussi une bande vertébrale chez ses 3 ex. de Kassina du lac Manengouba, qu’il nomme cependant K. maculosa. En l’absence de caractères morphologiques discriminants autres que la macu- lation, l’interprétation taxonomique des Kassina de type maculosa et de type decorata pose un probleme difficile, apparemment sans équivalent dans la batrachofaune came- rounaise. Pour tenter de le résoudre, les données biogéographiques suivantes doivent être prises en compte. 1°) Méme si des recherches ultérieures montrent que la forme a pattern decora- ta est plus largement répandue que nous ne le savons à présent, il y a de très fortes probabilités pour qu’elle soit absente au moins dans la moitié ouest du Plateau sud- camerounais, dont la batrachofaune est bien connue. Les aréotypes des deux formes sont donc différents. 2°) Néanmoins, les aires de répartition se recoupent en partie, dans l’Ouest et 1’ Adamaoua, où sont localisées les populations de type decorata. 3°) Les populations des deux formes sont étroitement contigués mais, dans l’état actuel des connaissances, ne s’interpénètrent pas. La situation, plus précisément, est la suivante. (1) Dans l’Ouest, si on admet que les populations de type decorata du mont Manengouba et de la Plaine des Mbos représentent un « foyer » unique, on constate que des individus parfaitement caractérisés de la forme maculosa sont présents 4 km au sud (Nsoung) et 20 km au nord (bord du Plateau bamiléké) et « en- cadrent » donc ce foyer de la forme decorata. (2) Sur 1’ Adamaoua, il y a environ 40 km a vol d’oiseau entre les populations de type decorata de Wakwa et celles de type maculosa de Hourso Manang. Il est difficile d’imaginer que la forme decorata représente une simple morphe, qui resterait localisée en quelques points d’une aire par ailleurs occupée par la seule morphe maculosa. Le statut de sous-espèce ne paraît pas non plus satisfaisant en raison de la proximité spatiale des deux formes et de l’intrication de leurs aires de répartition. Les données biogéographiques suggérent une « imperméabilité génétique » entre les Kassina des deux types, situation a laquelle correspondrait plutöt une séparation de niveau spécifique. Deux problémes connexes doivent aussi étre pris en considération. 1°) Que représentent les populations de Kassina à pattern decorata de 1’ Adamaoua par rapport à celles de l’Ouest, séparées par environ 360 km à vol d’oiseau? La découverte de populations géographiquement intermédiaires pourrait montrer qu’elles correspondent aux extrémités d’un cline. Dans le cas d’une bipolarité effective, deux possibilités devraient étre envisagées: (1) disjonction d’une aire auparavant continue occupée par une Kassina de type decorata, ou (2) acquisition indépendante d’un pattern identique (convergence) par des populations différant par leur taille et les proportions de leurs membres postérieurs. Un statut subspécifique conviendrait pour la premiere éventualité PHLYCTIMANTIS ET KASSINA DU CAMEROUN 113 alors que la seconde impliquerait une distinction de niveau spécifique. 2°) Un probleme similaire se pose pour les Kassina de type maculosa du Mont Cameroun. Le hiatus de distribution est moindre, de l’ordre de 115 km, mais les nombreuses prospections effectuées dans la région intermédiaire montrent qu’elles constituent très probablement un isolat. Des recherches complémentaires seraient cependant nécessaires avant de décider si elles représentent une sous-espèce, voire une espèce, endémique du Mont Cameroun. La discussion qui précède montre la complexité du problème posé par ces Kassina, et importance du travail de terrain qui reste à accomplir pour mieux en pré- ciser les données... sans être sûr de le résoudre, car 1l est possible que les méthodes morphologiques traditionnelles trouvent ici leurs limites. Le maintien du statu quo représente une solution trop « réunisseuse » (il est difficile d’admettre que les individus du Mont Cameroun, mesurant près de 4 cm, et ceux de la région de Ngaoundéré, dont la taille est de l’ordre de 2,5 cm, soient placés sous un même nom spécifique). Celle consistant à distinguer deux espèces divisées chacune en deux sous-espèces, voire à distinguer quatre espèces, est insuffisamment étayée dans l’état actuel des connais- sances. Pour le moment, la solution la plus équilibrée paraît donc être de considérer les Kassina à pattern decorata et les Kassina à pattern maculosa comme deux espèces distinctes. On peut en donner les diagnoses suivantes. K. maculosa: la plupart des macules dorsales sont sub-circulaires ou elliptiques et ne forment pas d’alignements nets; la macule supra-tympanique ne se recourbe pas vers le bas; la maculation des cuisses est généralement transverse. K. decorata: les macules dorsales sont au moins deux fois plus longues que larges, disposées suivant trois lignes principales, la médiane formant très souvent une bande continue; macule supra-tympanique incurvée vers le bas; marbrures des cuisses en général longitudinales. SPÉCIMENS DE RÉFÉRENCE Cinq spécimens de K. maculosa sont déposés au Muséum d’Histoire naturelle de Genève sous les numéros MHNG 2680.45 à 2680.49; 5 spécimens de K. decorata « ouest » sont déposés dans la même institution sous les numéros 2680.35 à 2680.39 et 5 de K. decorata « Adamaoua » sous les numéros 2680.40 à 2680.44. Kassina senegalensis (Duméril & Bibron, 1841) HISTORIQUE K. senegalensis, espèce-type du genre, a d’abord été trouvée près du Lac Galam, au Sénégal. Les recherches ultérieures ont montré qu’il s’agit d’un des Amphibiens savanicoles les plus largement répandus puisque son aire de répartition atteint la Somalie et l’Afrique australe (cf. carte in Schigtz, 1999). Toutefois, comme le souligne Schigtz (ibid.), il est possible que plusieurs espèces différentes, actuel- lement confondues sous K. senegalensis, peuplent certaines parties de cette aire immense. En tout cas, les populations du Cameroun ne diffèrent pas ostensiblement de celles d’ Afrique de l’Ouest (Schigtz, 1967; Rödel, 2000) ou de celles du nord-est de la RDC (Inger, 1968), que ce soit par leur aspect ou par leurs vocalisations. 114 J.-L. AMIET ci {| x a N ET Lo 3 NEP Soe Gi * ee e w cs a 2 a SI 1536 Hd = . TRS FIG. 11 Maculation dorsale chez K. senegalensis, populations méridionales (rangée supérieure) et septentrionales (rangée moyenne), et chez K. wazae (rangée inférieure). Pour les localités et dates de capture des spécimens figurés, se reporter aux numéros correspondants dans le tableau 1. Remarquer le contour finement dentelé des macules chez K. wazae. PHLYCTIMANTIS ET KASSINA DU CAMEROUN 115 712.335 K. senegalensis 1523711) K. cassinoides Fic. 12 Exemples de maculation céphalique latérale chez K. senegalensis (remarquer les sous-oculaires, fusionnées ou non) et chez K. cassinoides (grande extension de la supra-tympanique et fusion de toutes les macules). COMPLEMENTS DE DESCRIPTION Cette espéce est trop connue pour qu’une redescription soit nécessaire. Seuls les points suivants doivent être mentionnés. — 1°) Les macules sous-oculaires peuvent être soit séparées (63 % des individus étudiés), soit fusionnées (37 %); la macule loréale est fusionnée à la macule nasale; entre les premières et les secondes, un espace de la même teinte que le fond, de forme plus ou moins quadrangulaire, est généralement très apparent (Fig. 12). — 2°) L'appareil vocal externe, bien étudié par Drewes (1984), est du type 1: glande gulaire très développée (c’est, relativement, la plus grande chez les Kassina camerounaises), de forme rectangulaire ou elliptique, fossettes de rétraction des aires noires du sac vocal bien visibles, un gros pli interbrachial (Fig. 4 et Pl. II, Fig. e). — 3°) Chez la femelle, comme l’ont signalé Schmidt & Inger (1959), la face ven- trale est presque lisse dans ses trois-quarts antérieurs. — 4°) D’après Perret (1966) les dents vomériennes sont présentes. Dans mon matériel, elles ne sont bien développées que chez 4 individus sur 45 (8,8 %), absentes chez 30 / 45 (66,6 %) et très petites, masquées par la muqueuse buccale, chez 11 / 45 (24,4 %). DISTRIBUTION, VARIABILITÉ La distribution au Cameroun de K. senegalensis (carte Fig. 14) offre un raccourci de ce qu’elle est à l’échelle de l’Afrique: elle s’étend vers le sud jusqu’à la limite de la zone forestière, dont elle contourne au plus près les sinuosités. Vers le nord, elle atteint le domaine sahélo-soudanien. Dans cette bande, large d’environ 900 km, l’espèce montre une certaine variabilité, qui a conduit à séparer ici des populations « nord » et « sud » (Tab. 1 et 2, 116 J.-L. AMIET Planche I Kassina decorata et K. maculosa. (a) K. decorata Ouest, Santchou, 24.V.71. (b) K. maculosa Mt Cameroun, Bonianango, 11.VII.81. (c) K. decorata Adamaoua, Sadolkoulaye, 28.1V.73. (d) K. decorata Adamaoua, Ngaoundéré Vina, 02.VII.71. En a, c et d, livrée claire (diurne), en b, livrée de transition. Remarquer comment les orteils V et IV de l’individu figuré en d se replient ensemble et indépendamment des autres orteils sur une encoche d’une penne de fougere. Fig. 1 et carte Fig. 14), en prenant le rebord septentrional du Plateau de 1’ Adamaoua comme limite, mais il s’agit la d’un choix surtout pratique. Populations méridionales: la maculation dorsale est très développée, avec une large bande médio-dorsale continue, deux bandes latéro-dorsales plus ou moins discontinues et de grosses macules sub-circulaires dans la région postérieure: il peut y avoir un alignement de macules entre la bande médiane et les latéro-dorsales, esquissant deux bandes supplémentaires; ce type de maculation est illustré par la rangée supérieure de la Fig. 11. La taille moyenne est relativement grande, en parti- culier chez les individus du Plateau de 1’Adamaoua, auxquels correspondent, sur la Fig. 1, cing des six valeurs de L les plus élevées (PI. II, Fig. b). PHLYCTIMANTIS ET KASSINA DU CAMEROUN 117 Planche II Kassina senegalensis, K. wazae et K. cassinoides. (a) K. senegalensis Nord, Salak, 03.VII.73. (b) K. senegalensis Sud, Sadolkoulaye, 28.1V.73. (c) K. wazae, holotype, Mora / Waza, 14.VII.75. (d) K. senegalensis Sud, sac vocal gonflé, Minkama, 07.X.77. (e) de gauche à droite, face ventrale de K. wazae, K. senegalensis et K. cassinoides, Mora / Waza, 14.VII.75. Remarquer les macules sous-oculaires fusionnées en b, les denticulations irréguliéres du pourtour des macules et l’absence de sous-oculaires en c. En e, sac vocal de type 1 chez K. senegalensis (au centre), et de type 2 chez K. wazae (a gauche) et K. cassinoides (a droite). 118 J.-L. AMIET Populations septentrionales: à partir du piedmont de l’ Adamaoua, la taille et la maculation ont tendance a se réduire en allant vers le nord. Au sud de la Bénoué, on trouve encore des individus bien maculés (comme le spécimen de gauche, ligne du milieu, de la Fig. 11) mélés a des individus a bandes rétrécies. Dans la région de Garoua et de Mora-Waza, ils sont nettement plus petits et chez tous les bandes sont encore plus minces; les bandes latéro-dorsales sont les premiéres affectées par la réduction de la maculation; dans les cas extrémes, seule subsiste la bande médiane, plus ou moins discontinue (PI. II, Fig. a). De plus, les membres postérieurs sont pro- portionnellement plus courts dans les populations septentrionales (Tab. 2). Une comparaison portant seulement sur les populations les plus méridionales et les plus septentrionales pourrait suggérer une distinction subspécifique. Mais celle-ci rendrait mal compte du fait qu’il n’y a pas de gradient régulier, pas plus qu’une dis- continuité franche, dans le passage d’un type à l’autre. D’autre part, il faudrait savoir si une situation semblable se retrouve dans d’autres territoires, ce que la littérature ne dit pas. Il vaut mieux donc en rester au seul statut spécifique, en admettant une assez forte variation géographique des populations. ÉCOLOGIE Au Cameroun, plusieurs espèces d’Anoures, savanicoles à l’origine, ont pénétré plus ou moins profondément dans la zone forestière, en profitant des milieux herbacés de substitution dus à l’extension des surfaces cultivées et déboisées (voir plus haut, X. maculosa). Au contraire, K. senegalensis fait partie d’un groupe d’espèces qui sont restées cantonnées aux formations de savane, fussent-elles au contact immédiat de la forêt (Prychadena stenocephala et bibroni, Phrynobatrachus natalensis, Hyperolius igbettensis, Hemisus guineensis). Cette incapacité paraît assez paradoxale en regard de l’immense répartition de l’espèce au Cameroun et, plus généralement, en Afrique sub- saharienne. Du sud au nord de son aire camerounaise, K. senegalensis peuple en effet toute une succession de zones bioclimatiques allant des savanes humides de type guinéen, à vocation forestière dans les conditions climatiques actuelles, aux steppes sahélo- soudaniennes, et il est même probable que, encore plus au nord, elle atteigne le secteur sahélo-saharien aux abords du Lac Tchad. Elle est ainsi soumise à des régimes de pré- cipitations très différents : deux saisons pluvieuses et environ 1500 mm de pluie par an au sud, puis une saison des pluies unique de 7 mois, avec un surcroît de précipitations dû à l’altitude sur le Plateau de l’ Adamaoua (environ 1700 mm / an), et ensuite un gradient climatique régulier marqué par l’allongement de la saison sèche (8 à 9 mois dans la région de Mora) et la diminution des précipitations (environ 750 mm dans le même secteur). L’adaptabilité de cette espèce se manifeste aussi au niveau des températures: de 28°C de moyenne annuelle dans la plaine du Logone, à environ 20° dans la plus haute station relevée (1540 m, près de Bélel). Dans toute son aire de répartition, K. senegalensis est très souvent associée à d’autres Kassina: K. maculosa jusqu’à la bordure nord de l’ Adamaoua, puis, succes- sivement, les deux espèces suivantes. On trouvera, dans les parties relatives à ces espèces, des indications sur les diverses phonocénoses auxquelles participe K. sene- galensis. PHLYCTIMANTIS ET KASSINA DU CAMEROUN 119 Kassina wazae sp.n. TYPES Holotype: male capturé entre Mora et Waza, Cameroun, le 14.VII.75; numéro de collection Amiet: 75.378; déposé au Musée d’histoire naturelle de Genève sous le numéro MHNG 2680.50. Paratypes: cing autres males, méme localité et méme date; deux sont déposés au Musée d’histoire naturelle de Genève sous les numéros MHNG 2680.51 et 2680.52: autres paratypes actuellement dans la collection de l’auteur (numéros 75.361, 75.362 el 5-503)! DIAGNOSE Kassina ressemblant beaucoup par sa taille et son aspect a K. senegalensis de la « forme sud », mais s’en distinguant par: — son sac vocal de type 2, à glande gulaire petite et a bords convergents vers l’arriere (forme sub-triangulaire ou trapézoidale), — la présence d’un petit tubercule tarsien, — l’absence de macules sous-oculaires, — l’aspect finement déchiqueté sur les bords de toutes les macules, — les vocalisations. ETYMOLOGIE De la localité de Waza, sur la limite ouest du Parc national de Waza, proche de l’endroit où l’espèce a été trouvée. DESCRIPTION Principaux caractères de l’holotype: mensurations en mm: L: 35,5; T: 11,9; AB: PACA CETTE" P: 17. Proportions,en Yo: T/L: 33,5; AB/L: 20: C+J/L: 63.6; PIE: 47,9. Glande gulaire avec un sillon longitudinal dans sa partie postérieure (artefact probable). Maculation de la face dorsale: voir Fig. 11; maculation céphalique latérale: voir Fig. 12; livrée in vivo (livrée claire): voir Pl. II, Fig. c. Habitus, tégument, membres: espèce de taille moyenne (les 6 spécimens collec- tés mesurent entre 32,3 et 35,5 mm) a habitus allongé, semblable a celui des plus grandes K. senegalensis. Vu en plan, le museau paraît tronqué à son extrémité, alors que son contour est plutöt arqué chez K. senegalensis. Tympan un peu plus haut que large, presque vertical (à peine visible de dessus), situé juste derrière |’ ceil. Pas de dents vomériennes. Tégument dorsal d’apparence lisse sur toute sa surface; tégument ventral fortement et régulièrement granuleux (au moins chez les mâles), y compris sous les cuisses. Membres antérieurs relativement courts pour un représentant du genre Kassina (moyenne AB/L = 21 %). Main (Fig. 2) à doigts courts et épais, se rétrécissant progressivement vers l’extrémité, leurs pelotes non dilatées, plus étroites que les phalanges qui les portent; tubercules sous-articulaires larges et saillants; des excrois- sances supplémentaires modérément marquées sous certaines phalanges; aucune trace de palmure. Membres postérieurs épais et relativement plus courts que chez toutes les autres especes camerounaises de Kassina, sauf la forme septentrionale de K. senegalensis (moyenne C+J/L = 64,5 %, P/L = 49 %). Pied (Fig. 2) large et épais; orteils se rétré- cissant assez régulièrement depuis leur base, leur pelote terminale plus étroite que les 120 J.-L. AMIET Fic. 13 Caracteres diagnostiques de Kassina wazae n. sp. (a) Region gulaire (sac vocal de type 2). (b) Maculation céphalique laterale (disparition des sous-oculaires). (c) Empätement glandulaire de l’avant-bras. (d) Tubercule tarsien sous deux angles d’observation différents. (e) Pour compa- raison, tarse chez K. senegalensis (pas de tubercule tarsien). Voir aussi Fig. 11 (maculation dorsale). phalanges qui les précèdent (difference avec K. senegalensis et K. cassinoides); tuber- cules sous-articulaires tous présents, larges, bien saillants; un gros tubercule méta- tarsien interne, large, et un net tubercule métatarsien interne, petit et conique; une petite excroissance souple sur le tarse, en retrait du métatarsien interne (Fig. 13 d), assimilable à un tubercule tarsien (caractère propre à cette seule espèce au Cameroun, existant aussi chez K. fusca); un rudiment de palmure, épaissi, a peine différencié entre les orteils I et IT et II et III. Caractéres sexuels secondaires: région gulaire des males caractéristique (Fig. 13 a et Pl. II, Fig. e): — partie médiane épaissie du sac vocal (= glande gulaire) rela- tivement petite, ses bords épais et tendant a converger vers l’arriere, d’où un contour en triangle a sommet tronqué (trapézoidal chez un des six individus), sa surface plutöt concave; — pas de fossettes de rétraction apparentes mais, sur le pourtour de la partie PHLYCTIMANTIS ET KASSINA DU CAMEROUN 121 mediane, de forts plis circonvolués délimités par de profonds sillons. Cette structure de la région gulaire, assez proche de celle de K. cassinoides, correspond au type 2 défini plus haut. Un bourrelet interbrachial. Sur la face interne des bras, une zone glandulaire plus ou moins saillante, non pigmentée, s’élargissant vers le poignet (Fig. 13 c). Livrée: livrée dorsale (Fig. 11 et Pl. II, Fig. c) de même type que chez K. sene- galensis, et plus précisément les populations méridionales de celle-ci, car la macula- tion est large et très apparente: une bande médio-dorsale continue débute entre les yeux et se poursuit jusqu’a la région sacrée en s’élargissant d’avant en arriere (chez un individu, elle montre un début de dédoublement postérieur); de part et d’autre, de grandes macules beaucoup plus longues que larges forment deux alignement latéro- dorsaux débutant par les macules supra-oculaires; dans la région tout a fait postérieure, il peut y avoir de grosses macules sub-circulaires; contrairement a ce qui s’observe chez les K. senegalensis les plus maculées, il n’y a pas de rangées intermédiaires para- vertébrales. Latéralement, un caractére important de la maculation réside dans l’absence des macules sous-oculaires (Fig. 12 b), aussi bien antérieure que postérieure (ces macules sont présentes, séparées ou fusionnées, chez K. senegalensis). Membres peu ornés: 0 ou 1 macule sur le bras, 1 ou 2 sur l’avant-bras, 1 ou 2 sur la cuisse, 2 sur la jambe, 2 ou 3, trés petites, sur la tranche externe du pied. Comme chez K. senegalensis et K. cassinoides, il n’y a pas de coloration jaune a la base des membres. Une particularité notable, bien qu’elle ne soit pas apparente au premier coup d’ceil, est que toutes les macules sont très finement et irrégulièrement dentelées sur leur pourtour, qui a de ce fait un aspect déchiqueté. Sur le vif, en livrée diurne contrastée, les macules ont une teinte cuivreuse contrastant élégamment avec le blanc argenté de la teinte de fond. La face ventrale est incolore, sauf la région gulaire qui est assombrie, d’un gris plus ou moins foncé (caractére probablement propre aux males). DISTRIBUTION ET ECOLOGIE L’unique localité où K. wazae a été repérée, à environ 40 km au NE de Mora, se situe à la limite des steppes à épineux et des prairies inondables sahélo-soudaniennes (ou « yaérés »). Dans une grande plaine sans arbres, en contrebas de la route surélevée, plusieurs mares étaient, le 14 juillet 1975, déjà bien remplies: il avait plu au début du mois puis, assez abondamment, le 11 et le 12. La phonocénose accompagnant K. wazae ne comprenait plus guère d’espèces précoces, mais des espèces relativement tardives, appelant de préférence à proximité de collections d’eau assez profondes: Bufo stein- dachneri, Hydrophylax galamensis et Hyperolius viridiflavus pallidus. La forme septentrionale de K. senegalensis était représentée par des mâles beaucoup plus nombreux que ceux de K. wazae. Des l’arrivée sur le site, la présence de cette dernière a été détectée par ses appels, des « hou » brefs, assez étouffés, bien distincts des « oui » modulés de K. sene- galensis. Au moment de ma visite, les males de K. wazae se tenaient plus loin de l’eau que ceux de K. senegalensis. 122 J.-L. AMIET / K. senegalensis sud re N K. senegalensis nord e K. cassinoides O K. wazae 11° 10° ge 8° 78 6° inst 7 | | 5° gO! o = LI 4° 3° 14 13° 14° D 15° 16° Fic. 14 Carte du Cameroun montrant la répartition des mailles ot ont été repérées K. senegalensis (populations « sud » et « nord »), K. cassinoides et K. wazae. La limite forét — savane est sym- bolisée par la ligne avec triangles (pointés vers la savane) et le rebord septentrional du Plateau de l’Adamaoua par la courbe de niveau de 900 m. PHLYCTIMANTIS ET KASSINA DU CAMEROUN 123 DISCUSSION Comme le releve Schigtz (1999), de nombreuses « formes » de statut taxo- nomique incertain sont rattachées a K. senegalensis. L’espece décrite ci-dessus en est apparemment proche mais plusieurs bons caractères permettent de l’en séparer et le fait que les deux soient localement syntopiques confirmerait, si besoin était, qu’elles sont spécifiquement distinctes. K. wazae est peut-être une espèce adaptée à la zone sahélo- soudanienne, et donc à de longues saisons sèches, de l’ordre de neuf mois là où elle a été trouvée. Bien que ses appels soient aisément repérables, elle n’a pas été notée plus au sud, où elle paraît être remplacée par K. cassinoides. La batrachofaune de l’extrême nord du Cameroun, en gros entre les 11ème et 13ème parallèles, reste quasiment inconnue. On peut supposer que la découverte de K. wazae ne représente pas un cas exceptionnel et que la prospection de la zone des yaérés et des confins du Lac Tchad permettrait de trouver d’autres espèces nouvelles pour la science ou, au moins, pour le territoire camerounais. Kassina cassinoides (Boulenger, 1903) COMPLÉMENTS DE DESCRIPTION Cette Kassina est la plus grande du Cameroun (cf. Tab. 2 et Fig. 1). Il n’y a aucune différence entre les spécimens camerounais et ceux d’Afrique de l’Ouest, tels qu'ils ont été décrits et figures par les auteurs qui ont pu les observer sur le terrain (Schigtz, 1967, 1999; Rödel, 2000; Böhme, 2005). Il est donc superflu d’en donner une description et seules seront relevées ici les particularités suivantes. Dents vomériennes: chez 9 des 11 spécimens étudiés, elles sont bien déve- loppées; chez les deux autres, elles sont petites et plus ou moins masquées par la muqueuse buccale, mais décelables en passant un objet dur à leur emplacement. Maculation céphalique latérale: elle est caractérisée par la fusion des sous- oculaires en une macule linéaire unique, aux bords plus ou moins nets, reliant la loréale et la nasale, fusionnées, à la macule supra-tympanique très étendue et recouvrant le tympan (Fig. 12); cet ensemble, qui réunit toutes les macules latérales de la tête, est au point de départ des bandes latérales et latéro-dorsales. Disques digitaux: Schigtz (1967) et Rödel (2000) relèvent la présence de disques de la largeur des tubercules sous-articulaires à l’extrémité des doigts et des orteils. Ce caractère se retrouve chez les spécimens camerounais (Fig. 2). Il peut paraître surprenant chez une espèce savanicole et apparemment terrestre mais Rödel (op. cit.) indique qu’il a observé des mâles appelant depuis des arbres à des hauteurs de plusieurs mètres. Même si ce comportement n’est pas général (il pourrait dépendre du type de formation végétale localement fréquenté par l’espèce), il permet de mieux comprendre, du point de vue fonctionnel, l’existence de ces disques. Région gulaire des mâles: K. cassinoides partage avec l’espèce précédente la particularité d’avoir un sac vocal de type 2. D’après Schigtz (1967), « The “protective flap”, characteristic of the other members of the genus, is missing or very poorly developed ». Chez les individus du Cameroun la partie médiane du sac vocal (glande 124 J.-L. AMIET gulaire) est effectivement plus petite que chez les autres especes et, surtout, son pour- tour forme des bourrelets recoupés par des sillons: l’aspect qui en résulte est très semblable à celui du reste du tégument gulaire, très fortement plissé, ce qui peut donner l’impression que la partie médiane manque. De plus, il n’y a pas d’aires noirätres à la surface du sac vocal, donc pas de fossettes de part et d’autre de la glande gulaire. Une autre particularité est que, chez les 10 mâles examinés, il n’y a pas de repli inter- brachial, bien que le sac vocal s’étende loin vers l’arrière (Pl. II, Fig. e). Tegument ventral de la femelle: il est lisse sur presque toute sa surface. Seule la partie postérieure de l’abdomen montre des granulations, ou plutôt des ridules trans- verses. DISTRIBUTION, ÉCOLOGIE, PHONOCÉNOSES Ses appels caractéristiques et audibles de loin facilitent le repérage de K. cassi- noides. L'espèce n’a été rencontrée que dans la partie nord du Cameroun, où elle a été relevée dans 6 mailles de 10 minutes de côté (Fig. 14). Vers le sud, son aire de distri- bution s’arrête juste au pied du versant septentrional de l’Adamaoua (Amiet, 1972). D'autres espèces savanicoles butent aussi sur cet obstacle, qui paraît jouer le rôle de « filtre thermique »: Phrynomantis microps, Hydrophylax galamensis, Ptychadena schubotzi. Vers le nord, K. cassinoides paraît absente dans les régions de Maroua et de Mora-Waza, où de nombreuses espèces d’Anoures ont pourtant été notées dans les diverses stations prospectées. Si elle était confirmée, cette différence pourrait être d’origine climatique. La région où K. cassinoides a été trouvée est en effet soumise a un climat soudanien de basse altitude, avec 900 à 1500 mm de précipitations annuelles et une saison sèche de moins de 7 mois; sa végétation est constituée de savanes boisées de type méso-soudanien ou soudano-sahélien. Celle où elle paraît absente connaît des conditions climatiques plus rigoureuses, avec moins de 900 mm de précipitations et plus de 7 mois de saison sèche, ce qui se traduit par un appauvrissement de la végé- tation: steppes à épineux, ou prairies inondables dans la vallée du Logone. Sur les sites de reproduction, les mâles de K. cassinoides ne sont jamais très nombreux, contrairement à ceux de K. senegalensis, les effectifs pouvant s’estimer en unités pour la première et en dizaines pour la seconde. En 1981, près de Mbé, au pied de l’Adamaoua, l’espèce était déjà bien active le 21 mai, probablement à la suite de pluies précoces, mais la période optimale d’activité se situe plutôt entre la mi-juin et la mi-juillet. Les autres espèces vocalement actives en même temps que K. cassinoides ont été relevées dans les six stations où l’espèce a été entendue. Celles qui ont été notées dans au moins la moitié des relevés sont les suivantes: K. senegalensis et Ptychadena bibroni (6/6), Hoplobatrachus occipitalis (5/6), Phrynomantis microps, Phrynobatrachus natalensis et Leptopelis viridis (4/6), Prychadena schubotzi, Hydro- phylax galamensis et Afrixalus weidholzi (3/6). D'autres espèces peuvent définir des variantes géographiques de cette phonocénose. Dans la partie sud, plus arrosée par suite de la proximité de 1’ Adamaoua, s'ajoutent: Afrixalus fulvovittatus, Hyperolius balfouri viridistriatus, Hyperolius igbet- tensis et Ptychadena oxyrhynchus, alors que dans la partie nord ces espèces sont rem- placées par Bufo xeros, Ptychadena trinodis, Ptychadena floweri et Hildebrandtia ornata (Afrixalus vittiger existe aussi dans ce secteur mais n’a pas été relevé dans les mémes biotopes que K. cassinoides). PHLYCTIMANTIS ET KASSINA DU CAMEROUN 93 CONCLUSION Cing espèces de Kassina peuvent être à présent reconnues au Cameroun. Une est nouvelle pour la science et connue d’une seule localité. Deux autres, morpho- logiquement très proches, avaient été confondues auparavant. Quant au genre Phlyctimantis, il n’est représenté que par une seule espèce. Ce bilan est relativement modeste en regard de la richesse spécifique globale de la batrachofaune camerounaise (environ 200 espèces) Un fait surprenant est l’absence d’une espèce de Kassina sylvicole, comparable aux K. lamottei ou K. arboricola de l’Afrique de l’Ouest. Il semble improbable qu’une telle espèce ait pu échapper aux investigations des chercheurs qui ont prospecté la zone forestière du Cameroun. Je n’en ai pas non plus trouvé trace malgré environ 1000 sorties de nuit effectuées dans tout le sud du pays. | En revanche, il est très possible que K. fusca, ou un taxon proche, soit repré- sentée au Nord-Cameroun, car la faune batrachologique de cette région est relati- vement plus mal connue que celle de la zone forestière ou de la Dorsale. REMERCIEMENTS Plusieurs collègues m’ont apporté leur aide pour ce travail. J.-L. Perret et T. Frétey, avec leur obligeance habituelle, m'ont fait bénéficier de leurs connaissances bibliographiques et m’ont communiqué références et documents. F. Dowsett-Lemaire, Ch. P. Blanc et M. Largen m'ont fourni d’utiles précisions sur des enregistrements sonores ou des spécimens qu’ils avaient obtenus. Qu'ils trouvent ici l’expression de ma reconnaissance pour leur amicale collaboration. REFERENCES AMIET, J.-L. 1972. Compte rendu d’une mission batrachologique dans le Nord-Cameroun. Annales de la Faculté des Sciences du Cameroun 12: 63-78. AMIET, J.-L. 1978. Les Amphibiens Anoures de la région de Mamfé (Cameroun). Annales de la Faculté des Sciences de Yaoundé 25: 189-219. AMIET, J.-L. 1979. Description de l’adulte et de la larve d’Hyperolius bopeleti n. sp. (Amphibia Anura, Hyperoliidae). Annales de la Faculté des Sciences de Yaoundé 26: 113-124. AMIET, J.-L. 1983. Un essai de cartographie des Anoures du Cameroun. Alytes 2 (4): 124-146. AMIET, J.-L. 2005. Les Hyperolius camerounais du groupe d’H. nasutus (Amphibia, Anura, Hyperoliidae). Revue suisse de Zoologie 112 (1): 271-310. ANGEL, F. 1940. Description de trois Amphibiens nouveaux du Cameroun, matériaux de la Mission P. Lepesme, R. Paulian et A. Villiers (2° Note). Bulletin du Museum, 2° série 12 (6): 238-243. BÖHME, W. 2005. Presence of Kassina cassinoides (Boulenger, 1903) in Senegal. Herpetozoa 18: 177-178. BOULENGER, G. A. 1906. Report on the Batrachians collected by the late L. Fea in West Africa. Annali dell Museo Civico di Storia Naturale di Genova 2: 157-172. DREWES, R. C. 1984. A Phylogenetic Analysis of the Hyperoliidae (Anura) : Treefrogs of Africa, Madagascar and the Seychelles Islands. Occasional Papers of the California Academy of Sciences 139: 1-70. INGER, R. F. 1968. Amphibia. Exploration du Parc National de la Garamba 52: 1-90. LARGEN, M. J. & DOWSETT-LEMAIRE, F. 1991. Amphibians (Anura) from the Kouilou River basin, République du Comgo. Tauraco Research Report 4: 145-168. 126 J.-L. AMIET LAURENT, R. F. 1976. Les genres Cryptothylax, Phlyctimantis et Kassina au Zaire. Annales du Musee Royal d’Afrique centrale 213: 1-67. LAURENT, R. F. & CoMBAZ, J. 1950. Sur l’attribution générique de certains Batraciens appar- tenant ä la sous-famille des Hyperoliinae. Revue de Zoologie et Botanique africaines 43(3): 269-280. LETOUZEY, R. 1968. Etude phytogéographique du Cameroun. Encyclopédie Biologique, LXIX. P. Lechevalier, Paris, 508 pp. LETOUZEY, R. 1986. Notice de la carte phytogéographique du Cameroun au 1 / 500 000 (1985). Institut de la Carte internationale de la Végétation, Toulouse, 240 pp. MERTENS, R. 1940. Amphibien aus Kamerun gesammelt von M. Köhler und Dr. H. Graf. Senckenbergiana 22 (3-4): 103-135. MERTENS, R. 1968. Zur Kenntnis der Herpetofauna von Kamerun und Fernando Poo. Bonner Zoologische Beiträge 19 (1-2): 69-84. PERRET, J.-L. 1966. Les Amphibiens du Cameroun. Zoologische Jahrbücher Abteilung für Systematik 8: 289-464. PERRET, J.-L. 1985. Description of Kassina arboricola n. sp. (Amphibia, Hyperoliidae) from the Ivory Coast and Ghana. South African Journal of Science 81: 196-199. PERRET, J.-L. 1986. Considérations sur le genre Phlyctimantis Laurent et Combaz (Anura, Hyperoliidae). Bulletin de la Societe neuchäteloise des Sciences naturelles 109: 19-26. RÖDEL, M.-O. 2000. Herpetofauna of West Africa. Vol 1. Amphibians of the West African Savanna. Chimaira, Frankfurt-am-Main, 335 pp. RÖDEL, M.-O. & ERNST, R. 2001. Redescription of the tadpole of Phlyctimantis boulengeri Perret, 1986 (Anura, Hyperoliidae) with preliminary comments on the biology of the species. Alytes 18 (3-4): 178-186. RODEL, M.-O., GRAFE, T. U., RUDOLF, V. H. W. & ERNST, R. 2002. A review of West African Spotted Kassina, Including a Description of Kassina schioetzi sp. nov. (Amphibia: Anura: Hyperoliidae). Copeia 2002 (3): 800-814. SCHIGTZ, A. 1967. The treefrogs (Rhacophoridae) of West Africa. Spolia Zoologica Musei Hauniensis 25: 1-346. SCHIOTZ, A. 1999. Treefrogs of Africa. Chimaira, Frankfurt am Main, 350 pp. STERNFELD, R. 1917. Reptilia und Amphibia. Hamburgische Wissenchaftliche Stiftung Ergeb- nisse der Zweiten Deutschen Zentral Afrika Expedition 1910-1911, 1: 407-510. SCHMIDT, K. P. & INGER, R. F. 1959. Amphibians. Exploration du Parc National de I’ Upemba 562264 pps OPE SUCHEL, J.-B. 1972. La répartition des pluies et les régimes pluviométriques au Cameroun. Travaux et documents de géographie tropicale, 5. Université fédérale du Cameroun et C.N.R.S., 287 pp. SUCHEL, J.-B. 1988. Les climats du Cameroun. Thése de Doctorat d’Etat, Université de Saint- Etienne, t. I-II: 1188 pp., et t. IV, atlas. REVUE SUISSE DE ZOOLOGIE 114 (1): 127-139; mars 2007 Redescription of Protaphorura octopunctata (Tullberg, 1876) and Protaphorura quadriocellata (Gisin, 1947) with description of two new related species from Siberia and Europe (Collembola: Onychiuridae) Romuald J. POMORSKI! & Ighor KAPRUS’2 ! Department of Biodiversity and Evolutionary Taxonomy, Zoological Institute, Wroctaw University, Przybyszewskiego 63/77, PL-51-148 Wroctaw, Poland. E-mail: onychus @biol.uni.wroc.pl 2 State Museum of Natural History, Ukrainian National Academy of Sciences, Teatral’na St. 18, UA-79008 L’ viv, Ukraine. | E-mail: kaprus-i@museum.lviv.net Redescription of Protaphorura octopunctata (Tullberg, 1876) and Pro- taphorura quadriocellata (Gisin, 1947) with description of two new related species from Siberia and Europe (Collembola: Onychiuridae). - Protaphorura octopunctata (Tullberg, 1876) and Protaphorura quadrio- cellata (Gisin, 1947) are redescribed based on type specimens and new materials. Protaphorura flavorufula (Martynova, 1976) is a junior syno- nym of P. octopunctata. Protaphorura tolae Sp. n. and Protaphorura saltu- aria sp. n. two new, closely related species from Yakutia and East Carpathians are described. An identification key to all Palearctic Prota- phorura species with 4 and more pseudocelli at base of antenna is pro- vided. Keywords: Collembola - Onychiuridae - Protaphorura - taxonomy - Europe - Siberia. INTRODUCTION Protaphorura octopunctata (Tullberg, 1876) was described on the basis of one specimen collected in the delta of the Yenisei river near “Dudino” (at present Dudinka near Norilsk [69°24’N 86°08’E]). According to the information of Dr Kronestedt (Stockholm) there is no such specimen in the collections of the Swedish Museum of Natural History in Stockholm and in the Museum of Evolution, Uppsala University (Tullberg worked at Uppsala University). Only specimens collected during the “Yenissey-Expedition, 1876” have survived until today, labelled “Tschulkowa, Lipura octopunctata, Sibirien” (Chulkovo [62°45’N 88°25’E], situated on the left bank of the Yenisei river, about 45 km to the north of the Sukhaya Bakhta river mouth and Bakhta settlement) and determined by H. Schött. Manuscript accepted 22.11.2006 128 R. J. POMORSKI & I. KAPRUS A short Latin diagnosis and three drawings (Tullberg, 1876) point out the following characters of P. octopunctata: - postantennal organ consists of 30 simple vesicles, arranged perpendicularly to the long axis of the organ; - 4 pseudocelli at base of antenna; - claw with denticle; - small anal spines, a little arched; - length - 2.5 mm. During more than one hundred years this laconic description was a basis for many faunistic records from Europe, Asia and North America (Salmon, 1964; Yosii, 1977; Christiansen & Bellinger, 1989; Jordana et al., 1997; Deharveng, 2005). Most of these records also contained a redescription of the species, but none of them was based on type or topotype material. Also Martynova’s (1976) records were based on the materials from Chukotka and Magadan. At present, there is no convincing redescription of this species in the literature. Thanks to the kindness of Dr Babenko (Moscow) we have received onychiurid material collected during ecological investigation of ecosystems of northern areas of Siberia. Some of these samples come from geographically and ecologically nearby areas (ca. 225 km eastern [69°08’N 91°49’E]) of the type locality of P. octopunctata. Within this material we have found two Protaphorura species with 4 pseudocelli at the base of antenna. Because of small size of reproductive adults (less than 1.5 mm in length) one could not be identified as P. octopunctata and represents a new species, belonging to an other Protaphorura group of species. The second one we have compared with the specimens from Chulkovo (determined by Schött) and with the type material of Protaphorura flavorufula (Martynova, 1976) and we have ascer- tained that they are conspecific. The absence of other similar Protaphorura species in the closest neighbourhood of the delta of the Yenisei river is suggesting that P. flavorufula is a junior synonym of P. octopunctata. Among the other Siberian materials (Yakutia) received from Dr Babenko we have found one species without pseudocelli at subcoxa, closely related to P. octo- punctata. Because of the presence of 4+4 pseudocelli at the antenna base Protaphorura quadriocellata (Gisin, 1947) is similar to P. octopunctata and initially the two species were confused. Protaphorura quadriocellata was first recorded from Switzerland by Gisin (1943) as “Onychiurus octopunctatus (Tullbg.) Stach (f. daviesi Bagn.)”. In the next two publications the author gave it a subspecies rank (Onychiurus armatus quadriocellatus) (Gisin 1947) and subsequently species rank (Onychiurus quadrio- cellatus), and he listed the most important characters in a table (Gisin, 1952). The presence of 3+3 pseudocelli only on abdominal tergum IV (versus — 4+4 pseudocelli) is the character which makes it possible to distinguish P. quadriocellata from P. octopunctata and was used in Gisin’s (1960) and Palissa’s (1964) keys. At present P. quadriocellata is commonly accepted as a valid species and has been recorded from many European countries, viz. Austria, Great Britain, Denmark, Germany, Norway, PROTAPHORURA OCTOPUNCTATA AND PROTAPHORURA QUADRIOCELLATA 129 South Russia, Slovakia, Spain, Switzerland and Ukraine (Deharveng, 2005). Only one attempt at a redescription of this species is Known, but it is based on specimens from Spain, far from the type locality (Jordana er al., 1997). Thanks to the kindness of Dr Lienhard (Geneva) we had a possibility to examine the holotype of P. quadriocellata. Its comparison with Ukrainian specimens from the East Carpathians (Kaprus’, 1999) reveals that they represent different species. In the light of the facts presented above we have decided to redescribe of P. octopunctata and P. quadriocellata. Besides, the present work contains descrip- tions of two new species. The material is deposited in following institutions: MC Martynova’s Collection in Moscow State Pedagogical University, Moscow; MNHG Museum of the Natural History, Genève; MNHU Museum of Natural History, Ukrainian National Academy of Sciences, L’ viv; SMNH Swedish Museum of Natural History, Stockholm; ZIW Department of Biodiversity and Evolutionary Taxonomy, Zoological Institute, Wroctaw University, Wroctaw. SPECIES DESCRIPTIONS Protaphorura octopunctata (Tullberg, 1876) Figs 1-5 Bas. Lipura octopunctata Tullberg, 1876: 40. Onychiurus (Protaphorura) flavorufulus Martynova, 1976 syn. nov. MATERIAL: 2 unreproductive males, 6 females: Russia, Putorana Plateau, Dyngengda Mt., the Alnetum forest, near Sobachje Lake (Yt Kjuul’) [69°08’N 91°49’E], litter, bush vegetation, 350 m alt., 27 VII 1997, leg. A. Babenko (MNHU, ZI). — 5 paratypes of P. flavo- rufula (2 unreproductive males, 3 females): Russia, Jakutia, harbor Ambartchik near Medvezhka river, 12 VIII 1972, leg. Shilov (MC). — 4 juvenile males, 2 juvenile females and 3 spp. fixed in alcohol: Russia, Krasnoyarskyi Krai, Chulkovo, the left bank of the Yenisei river (about 45 km north of the Sukhaya Bakhta river mouth and Bakhta settlement, 1876, leg. ? [62°45’N 88°25’E]) (originally labeled: Lipura octopunctata Tullb., Tchulkova, Jenisiejexp.76 determ. H. Schott) (SMNH). — 3 males, 5 females: Russia, Jakutia, delta of the Indigirka river, Salix, Carex and moss community on bog, 14-16 VII 1994, leg. A. Babenko. — 1 male, 1 female: Russia, Magadanskaya oblast, delta of the Kolyma river, Pokhotskaya jedoma, meadow, 18 VII 1994, leg. A. Babenko (ZIW). Diacnosıs: Within the group of Protaphorura with four and more pseudocelli at base of antenna P. octopunctata is characterized by lack of pseudocelli on sub- coxal and yellowish orange body color. It is closely related with described below P. tolae sp. n. (differences — see diagnosis of P. tolae). REDESCRIPTION: Color in alcohol from yellowish to yellowish orange and reddish yellow. Length without antennae: males 2.5-3.1 mm, females 2.8-3.8 mm. Body shape cylindrical, with relatively small anal spines set on distinct papillae (Fig. 1). Antennae approximately as long as the head. Furca reduced to shallow cuticular pocket with 2+2 setulae — 1+1 setulae located on a cuticular fold, remaining 130 R. J. POMORSKI & I. KAPRUS deo el DI al n ea FEN né % ti } ; Sue { £ à } Hi Ev. 2 a se 1 a | Nes 0.1 mm SEN OE ie NE NN We AAT, WA (aE Al NE EP P_i 1 NN) NA. PA = N N D ET Fi er Fe NA TR ER NU NX 2°: ADE wi n, x Ve € A AAC EZ SETTA > @ @) > a 7 £ 45 € € A > € 3 \ 2 a I) v pps i NN AM NDS NAN Ny aM PRR on En allo | UE À { WS 2 ALIA | DA TAMAS PM PR fae Pa ee ee N Co sas Ka NS AAA PAT Mr ZANNI TNA Ga NU A Trim No ut | TRI 34 1 ibe if NIS 0.1 an AH \ A = Seas ee pei ar) ATARI NS ; fe VAL MdA) di NT 229 LA (ARAN, Di NOT aan ZAN WAN j I Mia, TAO | ry’ N ; i 7 N 1 \ È IN 0 0 C/G) og) IN À PAL Le IO Fics 1-5 Protaphorura octopunctata (Tullberg). (1) Habitus and dorsal chaetotaxy. (2) Antennal II sense organ. (3) Postantennal organ and anterior cephalic pseudocelli. (4) Tibiotarsal chaetotaxy and claw of legs III. (5) Chaetotaxy of abdominal sternum IV. PROTAPHORURA OCTOPUNCTATA AND PROTAPHORURA QUADRIOCELLATA 131 1+1 setulae located distinctly below of the fold (Fig. 5). Granulation is more or less uniform, distinct. Antennal area well marked. Antennal segment IV with a subapical organite. Microsensillum on antennal segment IV in latero-external position, c.1/3 length from the base. Antennal segment II with microsensillum slightly below antennal IH sense organ. Thoracic terga II and III with microsensilla laterally. Antennal III sense organ built of 5 guard setae, 5 low papillae, 2 sensory rods and 2 similarly sized sensory clubs: one is morel-like distinctly granulated, the other is sponge-like (Fig. 2). Sensory rods are relatively high, reached papillae length. Postantennal sense organ consists of 30-40 simple vesicles (Fig. 3). Pseudocellar formula dorsally: 4(5-6)3(4)/022/3335(4)3(4-5); ventrally: 1/000/00000, all subcoxal without pseudocelli. Formula of parapseudocelli ventrally: 1/000/111(0)101™, all subcoxal with 1 parapseudocellus ventrally. Dorsal chaetotaxy variable with a tendency to plurichaetosis, well differen- tiated into macro- and microsetae as in Fig. 1. Sensilla invisible. Head without d,, with p, at the same level as other p setae. Abdominal tergum V without or with seta D» abdominal tergum VI usually with two, sometimes with three medial setae. Straight lines, passing through bases of short setae situated above anal spines, parallel. Between legs on pro-, meso- and metathorax 1+1, 2+2 and 2+2 setae respectively. Tubus ventralis with ca. 13-19+13-19 setae and 2-3+2-3 setae at base. Claws always with large teeth. Empodial appendage without basal lamella, appendage longer than inner edge of the claw. Tibiotarsi with 11 distal setae (Fig. 4). Male ventral organ is absent. BIOLOGY: P. octopunctatata lives in humid habitats in mountains and low- lands. It has been collected in forest soil and litter and mosses, which covered the river bank and bog. DISTRIBUTION: The species recorded from many countries, but probably most records are misidentifications. Univocal data come only from North and Middle Siberia. Protaphorura tolae sp. n. Figs 6-10 TYPE MATERIAL: Holotype female and 14 paratypes (7 adult females, 7 unreproductive males): Russia, Yakutia, Suntar Khayata Mt. Range, upper currents of Kyubyume river (63°13’N; 139°36°E, about 250 km to the east [along the only road] of Khangyga on Aldan River), 1430 m alt., southern slope, forest edge, droppings of mouse-hare or rock rabbit (= Ochotona), 7 VII 2002, leg. O. Makarova. The type material is preserved in following collections: ZIW — holotype and 5 paratypes (3 males and 2 females), MC — 2 females and 2 males, MNHG — 1 female and 1 male, MNHU - 3 males and 2 females). ETYMOLOGY: The species is dedicated to our friend Dr Anatolyi Babenko (Moscow), for his material and valuable help. DIAGNOSIS: Yellowish orange body color, relatively low papillae in antennal III sense organ, a shallow cuticular pocket as a remnant of furca and particularly the absence of pseudocelli on subcoxae are suggesting a close relationship of P. tolae sp. n. with P. octopunctata. The new species distinctly differs from P. octopunctata in pseudocellar formula and in more symmetrical chaetotaxy, without plurichaetosis. 132 R. J. POMORSKI & I. KAPRUS vous SÉ a D vovlly on en | elev LI al N EN Ln an nantes, Fe 3 n a < à È, 0.1 mm cha N ) arr, bi © >> Pas < Pb SI LSS, = fi ni ER RP Nas a ec TES CI, AD ) —— a \ 1 SIE si » Ud 100000 990 N «1152200066, 9009 nai A IR = RSM ESE ; È a = 1 n nes AU J H 2 \ ì 20 PMOAOA CH GY 09 Tal | NR N x eodaqane? LEE Uf \ N AN + EUX PRO RL ROIO d gi VE i NN PS < Hi = TI Dy Y NL eg Al 0.1 mm È \ Fics 6-10 Protaphorura tolae sp. nov. (6) Habitus and dorsal chaetotaxy. (7) Antennal III sense organ. (8) Postantennal organ and anterior cephalic pseudocelli. (9) Remnant of furca. (10) Tibiotarsal chaetotaxy and claw of legs III. PROTAPHORURA OCTOPUNCTATA AND PROTAPHORURA QUADRIOCELLATA 133 DESCRIPTION: Color in alcohol from yellowish to yellowish orange and reddish yellow. Length without antennae: males 1.8-2.0 mm, females 2.5-2.75 mm. Body shape cylindrical, with relatively small anal spines set on distinct papillae (Fig. 6). Antennae approximately as long as the head. Furca reduced to shallow cuticular pocket with 2+2 setulae — 1+1 setulae located on a cuticular fold, remaining 1+1 setulae located distinctly below of the fold (Fig. 9). Granulation more or less uniform and distinct. Antennal area well marked. Antennal segment IV with subapical organite. Microsensillum on antennal segment IV in latero-external position, c. 1/3 length from the base. Antennal segment II with microsensillum slightly below antennal III sense organ. Thoracic terga II and III with microsensilla laterally. Antennal III sense organ built of 5 guard setae, 5 low papillae, 2 sensory rods and 2 similarly sized sensory clubs: one is morel-like distinctly granulated, the other sponge-like. (Fig. 7). Sensory rods relatively high, reached papillae length. Postantennal sense organ consists of 36-40 simple vesicles (Fig. 8). Pseudocellar formula dorsally: 43/012/332-343 (on thoracic tergum IV lack of anterolateral pseudocelli); ventrally: 1/000/00000, all subcoxal without pseudocelli. Formula of parapseudocelli ventrally: 1/000/100000, all subcoxal with 1 parapseudo- cellus ventrally. Dorsal chaetotaxy rather regular, well differentiated into macro- and micro- setae as in Fig. 6. Sensilla invisible. Abdominal tergum V with seta p , abdominal tergum VI with one medial setae. Straight lines, passing through bases of short setae which are situated above anal spines are parallel. There are 1+1, 2+2 and 2+2 setae respectively. Tubus ventralis with ca. 10-11+10-11 setae and 2+2 setae between legs on pro-, meso- and metathorax at the base. Claws always with large teeth. Empodial appendage without basal lamella, appendage longer than inner edge of the claw (Fig. 10). Tibiotarsi with 11 distal setae. Male ventral organ is absent. Protaphorura quadriocellata (Gisin, 1947) Figs 11-15 Bas. Onychiurus armatus quadriocellatus Gisin, 1947: TYPE MATERIAL: Holotype — unreproductive/reproductive, moulting male, (G930) Switzerland, Langenthal (BE), Sphagnum in Fagus forest, 24 VIII 1941 (MNHG). OTHER MATERIAL EXAMINED: | male, 2 females, 7 juveniles Germany, Hessen, Hochtaunus near Frankfurt am Main, litter of cultivated, mixed forest; 21 VII 2001. leg. A. Smolis (ZIW). — 5 males, 4 females, Germany, Augsburg, near Munich, litter +soil , Fagus- Abies forest; 15 IV 2003. leg. T. Zieche (MNHU). DIAGNOSIS: P. quadriocellata belongs to the group of Protaphorura species with 4+4 pseudocelli at base of antenna, 2+2 pseudocelli on thoracic terga II and II and without anterolateral pseudocelli on abdominal tergum IV. It is closely related with described below P. saltuaria sp. n. (differences — see diagnosis of P. saltuaria). REDESCRIPTION: Color white. Length without antennae: males 1.8-2.0 mm, females 2.1-2.4 mm. Body shape is cylindrical, with relatively small anal spines set on distinct papillae (Fig. 11). Antennae approximately as long as the head. Furca 134 R. J. POMORSKI & I. KAPRUS = N \ u LU AR sercennn “ey, — B Fics 11-15 Protaphorura quadriocellata (Gisin). (11) Habitus and dorsal chaetotaxy. (12) Antennal II sense organ. (13) Postantennal organ and anterior cephalic pseudocelli. (14) Tibiotarsal chaetotaxy and claw of legs III. (15) Chaetotaxy of abdominal tergum V and VI (11 specimen from Germany; 12-15, holotype). PROTAPHORURA OCTOPUNCTATA AND PROTAPHORURA QUADRIOCELLATA 135 reduced to cuticular pocket with 2+2 setulae. Granulation more or less uniform, distinct. Antennal area well marked. Antennal segment IV with a subapical organite. Microsensillum on antennal segment IV in latero-external position, usually at 1/3 height of antennal IV, somewhat above the second row of setae. Antennal segment III with microsensillum slightly below antennal III sense organ (Fig. 12). Thoracic terga II and III with microsensilla laterally. Antennal III sense organ built of 5 guard setae, 5 low papillae, 2 sensory rods and 2 similarly sized sensory clubs: one is morel-like finely granulated, the other is sponge-like (Fig. 12). Postantennal sense organ consists of 28-36 simple vesicles (Fig. 13). Pseudocellar formula dorsally: 43(4)/022/33333; ventrally: 1/000/00000, each subcoxal with 1 pseudocellus. Formula of parapseudocelli ventrally: 1/000/111101”, allsubcoxal with | parapseudocellus ventrally. Dorsal chaetotaxy symmetrical, well differentiated into macro- and microsetae as in fig. 11. Sensilla invisible. Head without d,, with p, shifted forward in relation to other p setae. Abdominal tergum V with seta p,, abdominal tergum VI usually with two medial setae (Figs 11, 15). Straight lines, passing through bases of short setae situated above anal spines, parallel. Between legs on pro-, meso- and metathorax 1+1, 2+2 and 2+2 setae respectively. Tubus ventralis with ca. 13-19+13-19 setae and 2+2 setae at base. Claws with large teeth (claw III of holotype is without teeth). Empodial appendage without basal lamella, appendage longer than inner edge of the claw. Tibiotarsi with 11 distal setae (Fig. 14). Male ventral organ absent. Protaphorura saltuaria sp. nov. Figs 16-20 TYPE MATERIAL: Holotype — 1 male and 14 paratypes (4 males, 10 females) Ukraine, East Carpathians, Chornoghora Mt., Vorochta, soil and litter in Piceetum forest, 1450 m alt., 24 VIII 1993, leg. I. Kaprus’. The type material is preserved in following collections: MNHU —holotype and 11 paratypes (4 males and 7 females), MNHG - 3 females. OTHER MATERIAL EXAMINED: 3 males, 4 females, Ukraine, East Carpathians, Skolivs’ki Beskydy Mt., Kamianka, soil and litter in Fagaetum forest, 900 m alt., 25 X 1989, leg. I. Kaprus’ (MNHU). — 1 male, Ukraine, East Carpathians, Chyvchyny Mt., Burkut, soil and litter of Piceetum forest, 1100 m alt., 4 VIII 1991, leg. I. Kaprus’ (MNHU). — 2 males, Ukraine, East Carpathians, Chorna Mt., Vynoghradove, soil and litter in Fagaetum forest, 450 m alt., 5 IV 1989, leg. I. Kaprus’ (MNHU). — 1 female, Poland, East Carpathians, Bieszczady Mt., Muczne, soil and litter in Abieto-Fagaetum forest, 650 m alt., 1 X 1996, leg. I. Kaprus’ (MNHU). — 1 female, Poland, East Carpathians, Bieszczady Mt., Sekowiec, soil and litter in Abieto-Fagaetum forest, 550 m alt., 1 X 1996, leg. I. Kaprus’ (ZIW). ETYMOLOGY: The species name is derived from the Latin word “saltus” — mountain forest. DIAGNOSIS: Because of the presence of pseudocelli on all subcoxa and the same dorsal pseudocellar formula P. saltuaria sp. n. is closely related with P. qua- driocellata. This new species distinctly differs from P. quadriocellata in the lack of parapseudocelli on abdominal sterna II-IV, in the presence of only one medial seta in 136 R. J. POMORSKI & I. KAPRUS 9 C 02090 Ceotco gy / AUS A - ‘| a ) i SES 0.01 mm AUS NDR FLOTTE 0.1 mm Fics 16-20 Protaphorura saltuaria sp. nov. (16) Habitus and dorsal chaetotaxy. (17) Chaetotaxy of abdominal sterna I-VI. (18) Antennal III sense organ. (19) Postantennal organ and anterior cephalic pseudocelli. (20) Distal part of leg III. abdominal tergum VI and in location of p, seta on the head. P. quadriocellata has parapseudocelli on all abdominal sterna, p, seta in anterior position and usually has two medial setae on abdominal tergum VI. DESCRIPTION: Color in alcohol yellowish white. Length without antennae: males 1.8-2.0 mm, females 1.7-2.2 mm. Body shape cylindrical, with strong anal PROTAPHORURA OCTOPUNCTATA AND PROTAPHORURA QUADRIOCELLATA 137 spines set on distinct papillae (Fig. 16). Antennae approximately as long as the head. Furca reduced to cuticular pocket with 2+2 setulae. Granulation more or less uniforms and distinct. Area antennalis well marked. Antennal segment IV with a subapical organite. Microsensillum on antennal segment IV in latero-external position, c. 1/3 length from the base. Antennal segment II with microsensillum slightly below of antennal III sense organ. Thoracic terga II and III with microsensilla laterally. Antennal III sense organ consists of 5 guard setae, 2 sensory rods, 2 straight and granulated sensory clubs and 5 papillae (Fig. 18). Postantennal sense organ consists of 35-42 simple vesicles (Fig. 19). Pseudocellar formula dorsally: 4(3)3/022/33333; ventrally: 1/000/00000, all subcoxal with pseudocellus. Formula of parapseudocelli ventrally: 1/000/100001”, all subcoxal with | parapseudocellus. Position of pseudocelli and parapseudocelli is presented in Figs 16 and 17. Dorsal chaetotaxy, usually symmetrical, well differentiated into macrochaetae and microchaetae as in fig. 16. Sensilla weakly marked. Head without seta d,, with p, at the same level as other p-setae. Abdominal tergum V with seta p , abdominal tergum VI with one medial seta. Straight lines, passing through bases of short setae with are situated above anal spines, parallel. There are 1+1, 2+2 and 2+2 setae between legs on pro-, meso- and metathorax, respectively. Tubus ventralis with ca. 9+9 setae and 2+2 setae at the base. Ventral abdominal chaetotaxy as in Fig. 17. Claws are with small teeth. Empodial appendage without basal lamella, slightly longer than inner edge of the claw. Tibiotarsi with 11 distal setae (Fig. 20). Male ventral organ absent. BIoLOGY: P. saltuaria lives in humid litter and soil of beech and spruce forests of the Eastern Carpathians. A KEY TO THE PALEARCTIC PROTAPHORURA SPECIES WITH 4 AND MORE PSEUDOCELLI AT BASE OF ANTENNA 1 Subcoxael of I, II and III pair of legs without pseudocelli ............... 2 - Subcoxael of I, II, and III pair of legs with 1,0,0 pseudocelli respectively . . 3 - Subcoxael of I, II, and III pair of legs with 1, 1, 1 pseudocelli respectively . . 4 2 Pseudocellar formula dorsally: 4-63-4/022/3334-53-5 dorsomedial pseudocelli on thoracic tergum I and anterolateral pseudocelli on abdo- minal tergum IV present BN TEE APE PUSS. Ss P. octopuntata (Tullberg, 1876) North Asiatic part of Russia - Pseudocellar formula dorsally: 43/012/332-343, on thoracic tergum I lack of dorsomedial pseudocelli, on thoracic tergum IV lack of antero- lateralkpseudoce liter PR eme A ne P. tolae sp. n. Russia: Jacutia 3 Pseudocellar formula dorsally: 43/022/33342, claw with strong denticle EEE a ee PO P. mongolica (Martynova, 1970) Mongolia - Pseudocellar formula dorsally: 43/022/33343, claw without or rarely wath very smallidenticlezem 2 2.2.2 P. serbica (Loksa & Bogojevic, 1976) 138 12 R. J. POMORSKI & I. KAPRUS Central and South East Europe, Caucasus Mts., Crimea Mts., Kazakhstan, Tadzhykistan, South Siberia Thoracic terga II-II with 3+3 and 3+3 pseudocelli respectively PENIS wees dear). P. pseudocellata (Naglitsch, 1962) Great Britain, Czech Republic, Germany, Poland, Ukraine Thoracic terga II-III with 2+2 and 3+3 pseudocelli respectively............ 5 Thoracic terga II-III with 2+2 and 2+2 pseudocelli respectively ........... 6 Pseudocellar formula dorsally: 54/023/454-75-65 ee le. OR M et P. decempunctata (Kos, 1939) Slovenia Pseudocellar formula dorsally: 44/023/34353 ee A IE SARA P. caledonica (Bagnall, 1937) Great Britain Pseudocellar formula dorsally: 43/023/44453 ....P.macfadyeni (Gisin, 1953) Denmark, Finland, Germany, Iceland, Norway, Jan Mayen & Svalbard, Sweden Abdominal tergum IV without anterolateral pseudocelli ................. 7 Abdominal tergum IV with anterolateral pseudocelli .................... 9 Head with 4+4 posterior pseudocelli; pseudocellar formula dorsally: 44/022/33333 ....P. suboctopuntata (Khanislamova, 1986) Russia: Bashkiria Head with 3+3 posterior pseudocelli; pseudocellar formula dorsally: 123102233393 ee aus ione à neo taie Sean CRISI SEI IT 8 Abdominal sterna II-IV with 1+1 parapseudocelli SOUS CIN SRI EU PO NOT SONDE Re NE P. quadriocellata (Gisin, 1947) Austria, Great Britain, Denmark, Germany, Norway, Slovakia, South Russia, Spain, Switzerland Abdominal sterna II-IV without parapseudocelli EEE N PORE a 2 P. saltuaria sp. n. Ukraine, Poland Sensory rods of antennal III sense organ higher than accompanying papillae, the first one is located at external surface of base of first internal papilla, the second one located between of third and fourth papillae; pseudocellar formula dorsally: 43/022/33353(4) RR MEI Re e o P. nutak (Yosii, 1972) Japan, Far East of Russia Sensory rods of antennal III sense organ shorter than accompanying papillae and arranged typical, closely together, behind of papillae in the middle-part'of the organ .!. 17.5. eee lo RIA IT 10 Males with ventral'organ i Pte LIRE IRE 11 Males without ventral organ. 0 SRO ee ee eee 12 Males ventral organ located between abdominal sterna II and III; pseudocellar formula dorsally:43/022/3334-33 “atts iad ou abe ies SI Sa AT ee P. janstachi (Yosii, 1972) Russia: Caucasus Males ventral organ located between abdominal sterna III and IV; pseudocellar formula dorsally: 44-3/022/3335(4,6)3 or EEE ere oe P. eichhorni (Gisin, 1954) Luxemburg Pseudocellar formula dorsally: 43/022/46655 EN NE ee tee Ree NENNT Roe ande P. asensitiva (Stach, 1954) Slovenia Pseudocellar formula dorsally: 43/022/33353 DIRO Pe) RIO RITA a RO P. valsainensis (Acön, 1981) Spain PROTAPHORURA OCTOPUNCTATA AND PROTAPHORURA QUADRIOCELLATA 139 ACKNOWLEDGEMENTS We are greatly indebted to Dr Anatolyi Babenko (Severtsov Institute of Ecology and Evolution RAN, Moscow) for his material, information and valuable help in finding the places of collection of onychiurids on the map of Russia. We wish to express our sincere thanks to Dr Charles Lienhard (Museum of the Natural History, Geneve) and Dr Torbjörn Kronestedt (Swedish Museum of Natural History, Stockholm) for the loan of type and historic materials. The study was sponsored by the Wroclaw University — grant n°: 7-13-1201; 1018/S/IZ/2005. REFERENCES CHRISTIANSEN, K. & BELLINGER, P. 1989. The Collembola of North America north of the Rio Grande. Second edition. Grinnell College, Grinnel: 1518 pp. DEHARVENG, L. 2005. Fauna Europea: Collembola. Jn: DEHARVENG L. (ed.) Fauna Europea: Collembola. Fauna Europea version 1.5, http://www.faunaeur.org Gisin, H. 1943. Ökologie und Lebensgemeinschaften der Collembolen im Schweizerischen Excursionsgebiet. Revue suisse de Zoologie 50: 131-224. GISIN, H. 1947. Sur les Insectes Aptérygotes du Parc National Suisse. Ergebnisse der wissen- schaftlichen Untersuchung des Schweizerischen Nationalparks 2: 77-91. GisIn, H. 1952. Notes sur les Collemboles, avec démembrement des Onychiurus armatus, ambulans et fimetarius auctorum. Mitteilungen der schweizerischen entomologischen Gesellschaft 25: 1-22. GISIN, H. 1960. Collembolenfauna Europas. Muséum d'Histoire Naturelle, Genève, 312 pp. JORDANA, R., ARBEA, J. I. SIMON, C. & LUCIANEZ, M. J. 1997. Collembola Poduromorpha Onychiuridae Onychiurinae. Fauna Iberica. Vol. 8. Museo Nacional Ciencias Natu- rales, Madrid, 807 pp. Kaprus’ I. J. 1999. Reaction of Collembola communities to anthropogenic substitution of forests in the Upper Dniester Basin (Eastern Beskidy). Roczniki Bieszczadzkie 8: 257-290. MARTYNOVA E. F. 1976: Vidy roda Onychiurus Gervais, 1841 (Collembola, Onychiuridae) severa 1 severo-vostoka Azji [A species of the genus Onychiurus Gervais, 1841 (Collembola, Onychiuridae) of the north and north-east Asia]. 10: 5-44. In: CHEREPANOV A. I. (ed.). Novye i maloizvestnye vidy fauny Sibiri. Novosti Fauny Sibiri, Novosibirsk, Nauka Sibirskoe Otdelenie, 207 pp. [in Russian] PALISSA, A. 1964. Apterygota. Die Tierwelt Mitteleuropas, Band IV, Teil I, Leipzig: 405 pp. SALMON, J. T. 1964. An index to the Collembola. Vol. 1-2, Society Victoria University of Wellington, Wellington, 144 pp. TULLBERG, T. 1876. Collembola borealia. — Nordiska Collembola, Ofversig af Kongl Vetenskaps Akademiens Forhandlingar, Stockholm 33, 5: 23-42. Yosu, R. 1977. Critical List of the Japanese Species of Collembola. Contributions from the Biological Laboratory Kyoto University 25,2: 141-169. ‘| 7 à ay Bin PE ti f REVUE SUISSE DE ZOOLOGIE 114 (1): 141-174; mars 2007 Cestodes of the genus Biuterina Fuhrmann, 1902 (Cyclophyllidea: Paruterinidae) from passeriform and piciform birds in the Ivory Coast, with a key to the species of the genus Boyko B. GEORGIEV!:? & Jean MARIAUX? ! Central Laboratory of General Ecology, Bulgarian Academy of Sciences, 2 Gagarin Street, 1113 Sofia, Bulgaria. E-mail: bbg@ecolab.bas.bg 2 Department of Zoology, Natural History Museum, Cromwell Road, London SW7 5BD, UK. 3 Natural History Museum, P.O. Box 6434, 1211 Geneva 6, Switzerland. E-mail: jean.mariaux @ville-ge.ch Cestodes of the genus Biuterina Fuhrmann, 1902 (Cyclophyllidea: Paruterinidae) from passeriform and piciform birds in the Ivory Coast, with a key to the species of the genus.- On the basis of recently collected material from the Ivory Coast (République de Cöte d’Ivoire) and other spe- cimens from the collections of Geneva Natural History Museum, we review the species of Biuterina (Paruterinidae) known from that country. Five species are recorded, among which two are new: B. pogoniuli sp. n. from Pogoniulus scolopaceus (Piciformes: Capitonidae) and B. petroniae sp. n. from Petronia dentata (Paseriformes: Ploceidae). B. africana Joyeux & Baer, 1928, from Tchagra spp. (Passeriformes: Laniidae), is redescribed on the basis of the type material from Benin, as well as of specimens from Angola, Guinea and the Ivory Coast. The species is reported for the first time from the latter two countries. B. pentamyzos (Mettrick, 1960) from Prionops plumata in Zimbabwe is recognised as a synonym of B. africana (new synonymy). B. cordifera Murai & Sulgostowska, 1983 and B. trian- gula (Krabbe, 1869) are reported from new hosts (Acrocephalus arundi- naceus and Anthus leucophrys gouldii, respectively) and for the first time in Africa. An identification key to the 34 species of Biuterina presently recognized worldwide is given. Keywords: Parasite - helminths - Paruterinidae - Biuterina - Africa - Ivory Coast - systematics - morphology - Laniidae - Ploceidae - Capitonidae - Sylviidae - Motacillidae. INTRODUCTION Biuterina Fuhrmann, 1902 comprises cestode parasites from passeriform and coraciiform birds; the number of the included species is considered to be between 30 and 40 depending on the generic definition adopted or on the views on the species validity (Matevosyan, 1969; Bona & Maffi, 1984; Schmidt, 1986; Kornyushin, 1989; Manuscript accepted 12.12.2006 142 B. B. GEORGIEV & J. MARIAUX Mariaux & Vaucher, 1989; Georgiev & Kornyushin, 1994; Georgiev et al., 2002, 2004). The species diversity of this genus is relatively well studied in temperate lati- tudes (e.g., Matevosyan, 1969; Kornyushin, 1989; Georgiev er al., 2004). However, most of the tropical Biuterina spp. are known from single records only. Five of the six species originally described from Afrotropical birds were recently redescribed (Mariaux & Vaucher, 1989; Georgiev et al., 2002), but data on their geographical dis- tribution and host ranges remained scarce. The only extensive faunistic survey on avian cestode in Africa during the last 20 years was carried out in the Ivory Coast (for sum- marised data, see Mariaux, 1994); however, some of the paruterinid species remained identified only at the generic level and their detailed taxonomic description required additional studies. The aim of the present article is to report new data on the morphology and taxonomy of Biuterina spp. collected from birds in the Ivory Coast, West Africa. In addition, we provide an identification key to the species of the genus. MATERIAL AND METHODS In total, 1,252 birds belonging to 174 species, 104 genera and 39 families were studied in the Ivory Coast by one of the present authors (JM) during 1985-1988 (for more detailed data, see Mariaux, 1994). Birds were captured with mist nets or shot. They were dissected immediately after death. Cestodes were removed from guts and fixed in 5% hot formalin. They were stored in 70% ethanol, stained in alcoholic hydro- chloric carmine, dehydrated in ethanol series, cleared in clove oil and mounted in Canada balsam. Some scoleces were mounted in Berlese’s medium to facilitate the examination of the rostellar hooks. The specimens were deposited in the Invertebrate Collection of the Natural History Museum, Geneva (MHNG). Details on their collection data and number of specimens are given in the text for each species. The metrical and meristic data are presented as the range, with the mean in parentheses and the number of measurements or counts taken (n). The measurements are given in micrometres unless otherwise stated. The developmental stages of the proglottides are designated as previously described (Georgiev & Vaucher, 2001). The nomenclature for the birds follows Howard & Moore (1980). TAXONOMIC PART Biuterina africana Joyeux & Baer, 1928 Paruterina pentamyzos Mettrick, 1960, new synonymy Biuterina pentamyzos (Mettrick, 1960) Matevosyan, 1964 MATERIAL EXAMINED: MHNG INVE 44454-44455 (Collection Joyeux, nos CJ 28/73a and CJ 28/74a), 2 slides, labelled “Biuterina africana, Telephonus senegalus, Gendre”, one of them containing a stained scolex squashed in Canada balsam, and the other containing three stained pregravid specimens in Canada balsam; recognised here as syntypes of B. africana (see Remarks); from Tchagra senegala (synonyms Pomatorhynchus senegalus, Telephonus sene- galus) (Passeriformes: Laniidae) collected at Bohicon, Benin by E. Gendre (see Joyeux & Baer, 1928); substantial parts of strobila contracted. - MHNG INVE 44456 (Collection Joyeux, nos CJ 28/75a and CJ 28/78), 2 slides, labelled “Biuterina africana, Telephonus senegalus, Gendre, Labé”, containing longitudinal sections of strobilar fragments of mature and postmature proglot- tides; from Tchagra senegala collected at Labé, Guinea by E. Gendre; these specimens were not mentioned in the publications of Joyeux. - MHNG INVE 40301 (Collection Fuhrmann - Baer BIUTERINA SPP. FROM THE IVORY COAST 143 nos 20/60-63), 4 slides, labelled “Biuterina africana, pie grieche, Angola” (two slides) or “Biuterina africana, Pomatorhynchus, Angola” (2 slides); mentioned by Fuhrmann (1943). Exact host names are not given in the labels; according to Fuhrmann (1943), B. africana was recorded in Angola from Pomatorhynchus australis (Smith) [now Tchagra australis (Smith)], Pomatorhynchus senegalus (L.) [now Tchagra senegala (L.)] and Pomatorhynchus anchietae (Bogage) [now Tchagra minuta anchietae (Bocage)] (Passeriformes, Laniidae). - MHNG INVE 16117 (former no 988.414, see Mariaux, 1994), 4 slides (2 fragmented specimens stained in hydrochloric carmine and mounted in Canada balsam and 2 scoleces mounted in Berlese’s medium), from Tchagra senegala pallida (Neumann) (Passeriformes: Laniidae), collected at Tortyia, Ivory Coast, on 10 February 1987, by J. Mariaux; mentioned by Mariaux (1994). The host specimen has been deposited in the Ornithological Collection of the MHNG, no. 1768.047. RE-DESCRIPTION OF THE SYNTYPES: Pregravid specimens with band-like body, 11.2-14.2 mm (12.4 mm, n =3) long, consisting of 106-114 (109, n=3) proglottides; contracted, single specimen with relaxed portions of strobila; maximum width at pre- gravid proglottides, 612-734 (654, n = 3). Scolex wider than neck, laterally rounded, with maximum width at middle of suckers, 385-450 (420, n = 3) (Fig. 1). Suckers oval, 115-130 (121, n = 7) in diameter, with well-developed musculature; in single spe- cimen, inner surface of suckers with punctiform spines (not distinct in remaining specimens). Rostellum sucker-like, retracted in all available specimens, with diameter 125-145 (133, n = 3); vertical muscular fibres inside distinct. Thick layer of radial muscular fibres surround rostellum (Fig. 1); glandular cells surround radial muscu- lature. Rostellar hooks arranged in 2 regular rows; 2 of 4 available scoleces had mis- sing rostellar hooks and others had densely packed hooks, for which exact number is difficult to determine but there are at least 38. Each hook with epiphyseal thickening on both handle and guard. Anterior hooks 75-80 (78, n = 12) long; blade longer than handle (Fig. 2). Posterior hooks 55-60 (58, n = 12) long; blade shorter than handle (Fig. 3). Rostellar hooks with blades directed anteriorly when rostellum is retracted. Genital pores irregularly alternating in short series, e.g. ... 2, 1,3, 1,2,1,2,5,1,2, 2, 2; pores open about middle of lateral proglottis margin. Genital atrium represented by infundibular distal part with thick walls and tubular proximal part surrounded by aggregation of glandular cells (Fig. 5). Dorsal osmoregulatory canals 5-8 (n = 10) in diameter; ventral osmoregulatory canals with diameter 13-38 (n = 10), with transverse anastomoses along posterior proglottis margin. Genital ducts pass between osmoregu- latory canals. Testes 10-14 (11,n = 15) in number, oval, form compact group in posterior most part of median field, posterior, dorsal and lateral to vitellarium and posterior and lateral to ovary, may overlap posterior margins of ovary dorsally (Fig. 4). External vas deferens coiled, with diameter 5-8 (7, n = 10), forms dense body in anterior poral part of median field together with surrounding glandular cells. Cirrus-sac highly elongate, 193-225 x 27-41 (215 x 33,n = 10), mostly cylindrical, tapered porally and rounded antiporally, thick-walled, usually crosses poral osmoregulatory canals. Internal vas deferens coiled. Evaginated cirrus not observed; minute (>1 um long) triangular spines present in canal of withdrawn cirrus. Ovary 110-145 (122, n = 10) wide, symmetrical, consisting of 2 compact rounded wings. Vitellarium compact, oval, 42-60 (52, n = 10) in diameter, at some distance from posterior proglottis margin (Fig. 4). Mehlis’ gland indistinct. Seminal receptacle elongate, situated dorsally, mostly between wings of ovary, extends from 144 B. B. GEORGIEV & J. MARIAUX Fics 1-3 Biuterina africana Joyeux & Baer, 1928, syntypes from Tchagra senegala ın Benin. (1) Scolex. (2) Anterior rostellar hooks. (3) Posterior rostellar hooks. Scale bars: 1, 100 um; 2, 3, 50 um. poral part of median field at level anterior to ovary to level of vitellarium, 105-118 x 28-33 (112 x 30,n = 7). Vagina opens postero-laterally to male pore, with copulatory part 138-180 (169, n = 7) long and 18-23 (20, n = 10) in diameter, consisting of thick- walled vaginal canal and cellular sleeve along entire length, with diameter of lumen 4-7 (6,n = 7), wider at poral end; conductive part of vagina short, narrow. Uterus in mature proglottides sac-like, spherical or with slightly irregular shape, thick-walled (Fig. 4). Primordium of paruterine organ appears as consolidation of cen- tral part of medullary parenchyma anterior to uterus. In pregravid proglottides, uterus consists of 2 sacs connected by transverse isthmus; paruterine organ elongate, may be slightly convoluted, with tapering anterior end surrounded by granular tissue (Fig. 6). No fully-developed eggs in specimens available. DESCRIPTION OF THE MATERIAL FROM THE Ivory COAST: Strobila band-like, gradually widening in posterior direction, with maximum width 751-848 (n = 2) at level of pregravid proglottides. Scolex rounded, wider than neck, with conical anterior protrusion (Fig. 7); maximum width 372-437, at middle of suckers. Suckers oval, 130- 161 (144, n = 8) in diameter, with developed musculature; their inner surface provided punctiform spine-like structures in transverse rows (Fig. 7, 12). Rostellum sucker-like, BIUTERINA SPP. FROM THE IVORY COAST 145 Fics 4-6 Biuterina africana Joyeux & Baer, 1928, syntypes from Tchagra senegala in Benin. (4) Mature proglottis, dorsal view. (5) Genital ducts in a mature proglottis, dorsal view. (6) Gravid proglot- tis. Scale bars: 4, 6, 200 um; 5,50 um. with diameter 148 (n = 1) when retracted and 186 (n = 1) when protruded. Rostellum surrounded by thick layer of radial muscular fibres and intensely staining glandular cells (Fig. 7). Rostellum armed with 38 (n = 3) or 42 (n = 1) rostellar hooks arranged in 2 regular rows. Each hook provided with epiphyseal thickening on both handle and guard. Anterior hooks 82-85 (83, n = 6) long, with length of refractive particle 81-82; 146 B. B. GEORGIEV & J. MARIAUX blade longer than handle (Fig. 8). Posterior hooks 61-63 (62, n = 6) long, with length of refractive particle 58-60; blade shorter than handle (Fig. 9). Proglottides craspedote; mature proglottides wider than long (Fig. 10); gravid proglottides slightly longer than wide (Fig. 11). Genital pores irregularly alternating in short series, open about middle of lateral proglottis margin. Genital atrium consists of infundibular thick-walled orifice and tubular hermaphroditic canal surrounded by intensely staining cells. Dorsal osmo- regulatory canals 4-8 (n = 10) in diameter. Ventral osmoregulatory canals with diameter 22-47 (n = 10), with transverse anastomoses along posterior margin of each proglottis. Genital ducts between poral osmoregulatory canals. Testes 10-15 (12, n = 11) in number, oval, situated in compact group posterior, dorsal and lateral to vitellarium and posterior and lateral to ovary, may overlap posterior margins of ovary (Fig. 10). External vas deferens coiled, with diameter 5-12 (7, n = 10), together with surrounding intensely stained cells forming dense body in anterior poral part of median field. Cirrus-sac highly elongate, 171-232 x 30-37 (201 x 34, n = 10), mostly cylindrical, with tapering poral end and rounded antiporal end, thick-walled; in mature proglottides, usually cross poral osmoregulatory canals; in gravid proglottides, often entirely situated in lateral field or slightly crossing poral osmoregulatory canals. Internal vas deferens coiled. Evaginated cirrus not observed; minute (length < 1 um) triangular spines with pointed tips seen in canal of withdrawn cirrus. Ovary consisting of 2 compact symmetrical wings. Vitellarium compact, oval, 51-69 (62, n = 10) in diameter, at some distance from posterior proglottis margin (Fig. 10). Mehlis’ gland not observed as glandular structure. Seminal receptacle elon- gate, situated mostly dorsally between wings of ovary, extends from poral part of median field at level anterior to ovary to level of vitellarium. Vagina opens postero- laterally to male orifice, with copulatory part 174-223 (193, n = 7) long, 14-23 (18, n = 10) in diameter, consisting of thick-walled vaginal canal and thick cellular sleeve along entire length; poral part of vaginal canal wider, covered with distinct micro- triches; conductive part of vagina short and thin. Uterus in mature proglottides sac-like, spherical or with irregular shape, thick- walled (Fig. 10). Primordium of paruterine organ appears as consolidation of central part of medullary parenchyma anterior to uterus. In pregravid and gravid proglottides, uterus consists of 2 sacs connected by thin transverse isthmus; paruterine organ elon- gate, with tapering anterior end surrounded by granular tissue (Fig. 11). Eggs oval; outer envelopes thin, often not distinct. Embryophore oval, thick, with diameter 40-45 (43, n = 10). Oncosphere oval, with diameter 28-34 (31, n = 10). Embryonic hooks: central pairs 19-21 (20, n = 7) long; lateral pairs 16-18 (17, n = 10) long. REMARKS: Biuterina africana Joyeux & Baer, 1928 was described from Tchagra senegala (L.) (Passerifomes, Laniidae) at Bohicon, Benin (Joyeux & Baer, 1928). Subsequently, it was recorded from the type host in Angola (Fuhrmann, 1943) and the Ivory Coast (Mariaux, 1994), and also from two other species of Tchagra Lesson, T. minuta (Hartlaub) and T. australis, in Angola (Fuhrmann, 1943). The only information on its morphology is the brief original description (Joyeux & Baer, 1928). We found substantial differences between the material from the Ivory Coast and the original description in relation to the size of the rostellar hooks. Other characters were BIUTERINA SPP. FROM THE IVORY COAST 147 Fics 7-9 Biuterina africana Joyeux & Baer, 1928, material from the Ivory Coast. (7) Scolex. (8) Anterior rostellar hooks. (9) Posterior rostellar hooks. Scale bars: 7, 100 um; 8, 9, 50 um. difficult to compare because the original description is illustrated with drawings of the scolex and rostellar hooks only; few strobilar characters are described and none of them illustrated. This provoked the present redescription of the type material. None of the specimens of B. africana in the Collection of C. Joyeux (currently deposited in the MHNG) is indicated as type material. We examined 4 slides from the type host «Telephonus senegalus» and recognised as syntypes two slides, MHNG 148 B. B. GEORGIEV & J. MARIAUX Fiss 10-11 Biuterina africana Joyeux & Baer, 1928, material from the Ivory Coast. (10). Mature proglottis, dorsal view. (11) Gravid proglottis. Scale bars: 10, 100 wm; 11, 200 um. 44454 and MHNG 44455 (Collection Joyeux nos CJ 28/73a and CJ 28/74a), since their labels are concordant with the information included in the paper containing the original description (Joyeux & Baer, 1928). The other two slides of B. africana in Joyeux’ collection, MHNG 44456 (Collection Joyeux nos CJ 28/75a and CJ 28/78), contain BIUTERINA SPP. FROM THE IVORY COAST 149 sectioned strobilar fragments of the same species; however, according to the labels, they were collected at Labé (Guinea) and therefore do not belong to the type series. Our study revealed discrepancies in the original description in relation to the length of the posterior rostellar hooks and the number of the testes (Table 1). In addition, we found that the most developed specimens of the type series were pregravid; therefore, the original measurements of the body length and the diameter of the eggs are not reliable. The material from the Ivory Coast corresponds well with the morphology of the type series; some differences in the measurements of the rostellum and suckers are probably due to the different state of the material (rather contracted type specimens versus relaxed specimens from the Ivory Coast). We confirm the identification of Fuhrmann’s (1943) material from Angola, although we are not sure of the identity of the host species of the available specimens. Their morphology corresponds well with that of the type series, including to the respective metrical and meristic data (Table 1). The new morphological data on B. africana enables a re-evaluation of the validity of another Afrotropical species of Biuterina, B. pentamyzos (Mettrick, 1960), a parasite of Prionops plumata poliocephala Shaw (Passeriformes, Laniidae) in Zimbabwe (Mettrick, 1960). Its type material was recently redescribed (Georgiev er al., 2002). The morphology of the scolex and both mature and uterine proglottides of B. pentamyzos entirely corresponds to the present redescription of B. africana, including the metrical data (Table 1). The difference in the body dimensions is due to taking measurements from a fully-developed (gravid) and well-relaxed specimen from the type series of B. pentamyzos (see Georgiev et al., 2002) compared with the rather contracted and not fully-developed syntypes of B. africana. Therefore, we recognise B. pentamyzos as a junior synonym of B. africana. Georgiev et al. (2002) reported the presence of an armature on the inner surface of the suckers of B. pentamyzos consisting of “fine, punctiform, spine-like structures in transverse rows”. This is distinct in the material from the Ivory Coast (Fig. 12). However, it is hardly seen in one of the syntypes of B. africana and not distinct in the remaining specimens, probably because of the contracted condition of the scoleces resulting in a highly folded internal surface of suckers. Matevosyan (1969) mentioned Tunisia as the only locality for B. africana. However, we did not find any published record of the species from that country. Summarising the above data, the geographical range of B. africana includes Guinea, the Ivory Coast, Benin (type locality), Angola and Zimbabwe; its hosts are restricted to birds of the family Laniidae: Tchagra senegala (type host), T. australis, T. minuta and Prionops plumata. Biuterina cordifera Murai & Sulgostowska, 1983 MATERIAL EXAMINED: MHNG INVE 15927 (formerly no. 987.273, see Mariaux, 1994), 1 slide (fragments of 2 specimens), 14 January 1987, host collection number CI 440 and MHNG INVE 15928 (formerly no. 987.274, see Mariaux, 1994), 1 slide (fragments of 2 specimens), 15 January 1987, host collection number CI 447, from Acrocephalus arundinaceus (L.) (Passeriformes, Sylviidae), collected at Adiopodoumé. One of the host specimens (CI 447) has been deposited in the Ornithological Collection of the MHNG, no. 1769.041. DESCRIPTION: Strobila gradually widening posteriorly; maximum width at level of gravid proglottides 559-578 (n = 2). Scolex (Fig. 13) rounded, with maximum width B. B. GEORGIEV & J. MARIAUX 150 "uounoods PoXE[9I-]9X ¢ ‘Aforeredes USAIS JOU SYOOY 9IUOÂIQUIS [e.loye] pur [eNU99 JO saut} “‘Ajoyeredas USAIS JOU SYOOY 1OLISISOI pue IOHAUE JO SWFUIT ¢ ‘8999 podojoA9p-A]mJ YIM Sopiyosoid syoe] jeLspeun adA) oy} osnedsaq afqui[oI Loos JOU op s339 JO J9JaWIp Ay) UO vez "POE.NUOI WOY JO sued [enurjsqns 10 eJIqOns amuo !suswroads prARITAIg | 81-91 pII-0I 81-91 6I-LI x 17-61 pII-0I IC-61 17-07 a 1270 E vE-87 Te z 6r-Ir a St-0f ch 9e = È 6E€-SE 5 3 IT-ET : ET PI cC-El €c-81 LyT-861 3 ECTYLI 081-791 081-8€1 9€-87 97 LE-0€ Or-Te Ip-L7 LTT-08I 961 TETILI 061-€91 STT-EGI VIII 01-8 SI-OI el-Ol vi-Ol 19-09 £C8-08 €919 19-65 09-SS 58-18 £C8-08 58-08 8-08 08-SL ch ch Tr-8€ OV ge < 091 cs 981-871 Sp SpI-SCI LSI-vel cri-cel I9I-OEI SvI-cel O£I-SII SOV Lvt LEV-TLE 097 OStT-S8£ ELT I 00€ 1 8p8-ISL OFT 1-071 I 1PEL-TI9 SA ç8S È LE < ICVICII (7007) (0961) ‘]D 19 ADIB1IOIH NOLMON Apnys Juasoig Apnys Juosoig Apnys juasoig 2MqeqUuUIZ Omgequiz Jsvod AIOA] vjoduy urusg pypumjd sdouo114J soz(unjuad 'g pypund sdouot4i4 soz(upjuad ‘g DIDSaUuas DASDYI] DUDILLD *g ‘dds nuspy2y DUDILUD 'g DIDSIUAS DASDYI], DUDIIL[D * g OS 0ST-061 85 OV v8 vv 007 OI X 061 00€ X OOS OSL SI (8761) Jovg 2 xnoAor uruog pjpSauas DASYYIT DUDIIL[D 'g sured je1oye] ‚red je.nuo9 :y)3u9] ‘syooy oruoAaquiq I9PWEIP :o1sydsooug Jojoureip :oroydoArquiq JOJOUURIP :s83q UIPIA Yu] ed Aroyejndoo ‘vurse, UIPIA Yu] :IBS-SNAND JOQUINU :S9)S9], Jo119}sod JOroJue :y)3u9] ‘SYOOY IP[[9JSOY JOQUINU :SYOOY Ie][9}SOM JOJQUILIP :UUN][O)SOM JOJQUILIP !SIOYONS JOJOUPIP :X9[09S UIPIM wnwrxew :Apog (wu) yu] :Apog 991n0$ A890T 1S0H soroadg ‘(0961 PLINOW) so2Kwmjuod g wAuouAs su pue 8761 ‘eg 2 xnoAof VUDILD DULLAINIG 107 eyep INSLIEW pur [BOLO AUIOS JO uosLedwo) ‘| ATAVL, BIUTERINA SPP. FROM THE IVORY COAST 151 Fic. 12 Biuterina africana Joyeux & Baer, 1928, material from the Ivory Coast. Sucker. Note armament consisting of punctiform spine-like structures arranged in transverse rows on the internal surface of the sucker. Scale bar: 10 um. at level of middle of suckers 321 (n = 1). Suckers slightly oval, with well-developed musculature, with diameter 129-150 (142, n = 4). Rostellum of only available scolex retracted (Fig. 13), with diameter 123, sucker-like, highly muscular; no glandular cells visible within or around rostellum. Rostellar hooks 30 in number, arranged in 2 regular rows; their blades directed anteriorly when rostellum retracted; each hook provided with 2 epiphyseal thickenings, | on handle and 1 on guard. Anterior hooks 58-62 (61, n = 5) long, with blade considerably longer than handle (Fig. 14). Posterior rostellar hooks 42-45 (44, n = 4), with blade almost as long as handle (Fig. 15). Neck contracted, 207 wide; first proglottides appear at 339 from posterior margin of suckers. Proglottides craspedote (Figs 16-18); mature, postmature and pregravid proglottides wider than long; gravid proglottides as long as wide or slightly longer than wide (Fig. 18). Genital pores irregularly alternating in short series. Genital atrium infundibular, thin-walled; no surrounding intensely stained cells. Genital ducts pass between osmoregulatory canals. Dorsal osmoregulatory canals 4-7 (n = 10) in diameter. Ventral osmoregulatory canals 24-34 (n = 10) in diameter, with transverse anastomoses along posterior margin of each proglottis. Testes oval, 8-11 (9, n = 10) in number, form compact group situated laterally, dorsally and posteriorly to vitellarium, often overlap posterior margin of ovary and 152 B. B. GEORGIEV & J. MARIAUX Fics 13-16 Biuterina cordifera Murai & Sulgostowska, 1983. (13) Scolex. (14) Anterior rostellar hooks. (15) Posterior rostellar hook. (16) Mature proglottis, dorsal view. Scale bars: 13, 16, 100 um; 14, 15, 10 um. ventral osmoregulatory canals (Fig. 16). External vas deferens convoluted, surrounded by intensely stained cells but not forming dense body. Cirrus-sac oval (Fig. 16), 57-84 x 27-33 (70 x 30, n = 10), tapered porally and rounded antiporally, thin-walled, may reach or just cross poral osmoregulatory canals. Internal vas deferens forms several BIUTERINA SPP. FROM THE IVORY COAST 153 coils in antiporal part of cirrus sac. Evaginated cirrus not observed; no armament within duct of withdrawn cirrus. Ovary bi-winged, medial, symmetrical, occupies entire width of median field (Fig. 16); wings round, compact. Vitellarium compact or slightly lobed, oval or irre- gular in shape, with diameter 39-72 (48, n = 10). Seminal receptacle ovoid to elliptical, dorsal to ovary. Vagina opens dorsally to male pore; not clearly separated into copu- latory and conductive parts. Uterus in mature proglottides antero-dorsal to ovary, sac-like, thick-walled, almost spherical or with irregular shape (Fig. 16). In postmature proglottides, uterus horseshoe-shaped, with lateral ends directed posteriorly; primordium of paruterine organ as consolidation of medullary parenchyma, entirely enveloping uterus. In pregravid (Fig. 17) and gravid (Fig. 18) proglottides, uterus with elongate lateral branches; paruterine organ occupies entire median field; anterior part of paruterine organ granular. In slightly contracted gravid proglottides, uterine branches convoluted, suggesting presence of uterine capsules each containing several eggs (Fig. 18). Eggs oval; outer shell thin, often not distinct, closely envelops embryophore. Embryophore oval, 30-38 (34, n = 10) in diameter, thick-walled. Oncosphere oval, 20-25 (23, n = 10) in diameter. Embryonic hooks: central pair thin, 16-17 (n = 7) long; intra-lateral hooks thin, 14-15 (n = 7) long; extra-lateral hooks stout, 14-15 (n=7) long. REMARKS: The present material was reported as Paruterina sp. from two of nine examined specimens of the host species (Mariaux, 1994). Its morphology fits well with the previous descriptions of B. cordifera (see Murai & Sulgostowska, 1983; Georgiev et al., 2004). This species was originally described from Lusciniola melanopogon (Temmink) (type host), Acrocephalus scirpaceus Hermann and A. shoenobaenus (L.) in Hungary (Murai & Sulgostowska, 1983) and later reported from A. scirpaceus in the Czech Republic and from Erithacus megarhynchos (L.) in Bulgaria (Georgiev et al., 2004). The present study reports the first record of this species in A. arundinaceus and the first record for Africa. Biuterina triangula (Krabbe, 1869) Fuhrmann, 1908 MATERIAL EXAMINED: MHNG INVE 15961 (formerly no. 988.401, see Mariaux, 1994), 5 slides: 1 stained entire pregravid specimen, 5 stained juvenile specimens and 4 stained stro- bilar fragments without scoleces, and 3 scoleces in Berlese’s medium; 11 February 1988, host collection number CI 1051, ex Anthus leucophrys gouldii Fraser (Passeriformes, Motacillidae), collected at Tortyia. The host specimen has been deposited in the Ornithological Collection of the MHNG, no. 1765.070. DESCRIPTION: Pregravid specimen 18.5 mm long, with strobila gradually widening posteriorly and reaching maximum width of 770 (n = 1) at level of pregravid proglottides, consists of 107 proglottides: 24 juvenile, 33 premature, 21 mature, 18 postmature and 11 pregravid. Scolex with protruded apical part (Fig. 19), clearly out- lined from neck by its greater diameter, with maximum width 350-405 (381, n = 6) at level of suckers. Suckers somewhat oval, with well-developed musculature, 100-140 (121, n = 24) in diameter, with their apertures directed antero-laterally. Rostellum sucker-like (Fig. 19), 102-110 (106, n = 6) in diameter, highly muscular; numerous intensely-staining cells present in anterior half of rostellum. Rostellum separated from 154 B. B. GEORGIEV & J. MARIAUX Fics 17-18 Biuterina cordifera Murai & Sulgostowska, 1983. (17) Pregravid proglottis. (18) Gravid proglot- tis. Scale bars: 100 um. surrounding parenchyma by thick layer of radial musculature, outside of which is aggregation of intensely-staining (glandular) cells. Rostellar hooks 40-44 (n = 9) in number (40, n = 3; 42, n = 2; 44, n = 4), arranged in 2 regular rows; blades directed anteriorly when rostellum retracted. Each hook provided with slightly expressed BIUTERINA SPP. FROM THE IVORY COAST 155 19 Fics 19-21 Biuterina triangula (Krabbe, 1869). (19) Scolex. (20) Anterior rostellar hooks. (21) Posterior rostellar hooks. Scale bars: 19, 100 mm; 20, 21, 50 mm. epiphyseal thickening on both handle and guard. Anterior hooks 58-65 (62, n =24) long, with blade considerably longer than handle (Fig. 20). Posterior rostellar hooks 37-43 (41, n =23), with blade almost as long as handle (Fig. 21). Neck 160-202 (180, n = 6) wide in its narrowest part; first proglottides appear at 540-800 (620, n = 6) from 156 B. B. GEORGIEV & J. MARIAUX posterior margin of suckers. Proglottides craspedote (Figs 22, 24); mature and post- mature wider than long; pregravid as long as wide. Genital pores irregularly alternating inishort series, e.g. ... 3, 3,2, 1, 1, 1,1, 2.1, 2; 1, 3,1, 1... No genital papilla Genital atrium thick-walled, 22-27 (24, n = 10) deep, with infundibular aperture, 10-19 (14, n = 10) in diameter, and cylindrical proximal part; intensely staining cells surround genital atrium. Genital ducts pass between osmoregulatory canals. Dorsal osmoregu- latory canals 5-12 (n = 10) in diameter. Ventral osmoregulatory canals 18-52 (n = 10) in diameter, with transverse anastomoses along posterior margin of each proglottis. Testes oval, 6-9 (8, n = 14) in number, form compact group situated laterally, dorsally and posteriorly to vitellarium, overlapping osmoregulatory canals and posterior margin of ovary (Fig. 22); maximum diameter of fully-developed testes 48-58 (54, n = 10). External vas deferens convoluted, surrounded by intensely stained cells, which form dense body at anterior proglottis margin. Cirrus-sac elongate-oval (Fig. 23), with almost cylindrical poral part, thick-walled, reaches or just crosses poral osmoregulatory canals, 135-155 x 24-33 (145 x 29, n = 9) in mature proglottides, 133-155 x 30-33 (142 x 32, n = 4) in postmature and pregravid proglottides. Internal vas deferens forms several coils in antiporal part of cirrus sac. Evaginated cirrus not observed; armament within ductus cirri consisting of small triangular spines (< 1 mm long) which occupy only its distal part (Fig. 23). Ovary bi-winged, medial, symmetrical, occupies almost entire width of median field, when fully developed 147-162 (155, n = 6) wide, with round or slightly oval, compact wings (Fig. 22). Vitellarium compact, oval or with irregular shape close to oval, with diameter 45-68 (43, n = 7). Seminal receptacle elongate, almost tubular, with slightly expanded and rounded poral end 22-30 (26, n = 8) in diameter, tapers anti- porally and gradually forms straight canal passing towards Mehlis’ gland. Mehlis’ gland situated anterior to vitellarium, globular, with diameter 23-35 (31, n = 8), not distinct in some proglottides. Vagina opens dorsally to male pore; its copulatory part 113-128 (125, n = 8) long, slightly shorter than cirrus sac or of same length, consisting of thin-walled canal with diameter of lumen 4-6 (n = 8), surrounded by cellular sleeve (Fig. 23); diameter of copulatory vagina 13-18 (15, n = 8). Conductive part of vagina thin, short. Uterus in mature proglottides antero-dorsal to ovary, sac-like, thick-walled, almost spherical or with irregular shape (Fig. 22). In postmature proglottides, uterus gradually divides into 2 lateral expanded sacs connected by narrow anterior isthmus; primordium of paruterine organ forms as consolidation of medullary parenchyma anterior to developing uterus. In pregravid proglottides (Fig. 24), uterus situated in pos- terior third of median field, consisting of 2 lateral round sacs; paruterine organ cylin- drical, with rounded anterior end surrounded by granular tissue. No developed eggs in material studied. REMARKS: The original description of Biuterina triangula (= Taenia triangulus Krabbe, 1869) was based on specimens from ‘Turdus sp.’ and did not mention the type locality (Krabbe, 1869). The subsequent revisers considered that the material originat- ed from “West Europe’ (Matevosyan, 1969) or ‘Central Europe’ (Kornyushin, 1989), probably because the collector was Bremser (the naturalist J. G. Bremser of Vienna, BIUTERINA SPP. FROM THE IVORY COAST 157 Fics 22-24 Biuterina triangula (Krabbe, 1869). (22) Mature proglottis, dorsal view. (23) Genital ducts in postmature proglottis, dorsal view. (24) Pregravid proglottis, dorsal view. Scale bars: 22, 24, 200 mm; 23,50 mm. see Sattman, 2000). As demonstrated by previous studies (Kornyushin, 1989; Georgiev et al., 2004), two other species with anterior rostellar hook of a similar length but with different strobilar morphology, B. cordifera and Spasskyterina trianguloides 158 B. B. GEORGIEV & J. MARIAUX Kornyushin, 1989, were sometimes misidentified as B. triangula. Therefore, the data for the host range and the geographical distribution of B. triangula need to be re- evaluated, which is difficult for the moment because of the lack of morphological information accompanying most of previous records. The original description of B. triangula is very brief, giving information on the number and the length of rostellar hooks (Table 2) and the irregular alternation of the genital pores (Krabbe, 1869). Fuhrmann (1908) re-examined the type material and additional material from Turdus pilaris L. and reported the length of the rostellar hooks of the type as 57 and 39 um. He mentioned that the types are in bad condition and only added the length of the cirrus sac (0.1 mm) to the original description. Kornyushin (1989) published a detailed redescription on the basis of specimens from Anthus campestris (L.) from the Crimea, Ukraine, very much resembling the present material. Some differences in the metrical data (Table 2) are of minor importance and are probably due to the different methods of fixing and processing the specimens. All the previous records of B. triangula are from the Palaearctic Region (Matevosyan, 1969; Kornyushin, 1989). The present finding is the first record of this species in Africa and a new host record. Biuterina pogoniuli sp. n. MATERIAL EXAMINED: MHNG INVE 16064, 49753 (formerly no 988.174, see Mariaux, 1994), 3 slides, 1 entire specimen, 1 specimen with scolex mounted separately in Berlese’s medium and strobilar fragments of 2 further specimens, from Pogoniulus scolopaceus (Bonaparte) (Piciformes: Capitonidae), collected on 20 April 1988 at Dabou. The host specimen has been deposited in the Ornithological Collection of the MHNG, no. 1765.018. Holotype: MHNG INVE 16064, an entire specimen in Canada balsam, mounted in the same slide with another strobilar fragment. Paratypes: MHNG INVE 49753, 1 specimen with scolex mounted separately in Berlese’s medium and 2 strobilar fragments, | of them on the same slide as the holotype. ETYMOLOGY: The species name pogoniuli is a derivate of the generic name of the host. DESCRIPTION: Body band-like. Entire gravid specimen with scolex 37 mm long, consisting of 145 proglottides (44 juvenile, 24 premature, 23 mature, 19 postmature, 26 pregravid and 9 gravid). Specimen without scolex and initial part of strobila, 38 mm long, consisting of 107 proglottides (7 juvenile, 27 premature, 19 mature, 12 post- mature, 33 pregravid and 9 gravid). Maximum width 0.95-1.20 mm (1.05 mm, n = 4), at level of gravid proglottides. Scolex (Fig. 25) almost globular, with maximum diameter 420 (n = 1) at level of middle of suckers; anterior part of scolex protrusible. Suckers round, 179-192 x 179-183 (188 x 180, n = 4) in diameter, with moderately developed musculature, open via transverse slit-like aperture. Rostellum sucker-like, with concave anterior surface, 192 (n = 1) in diameter; numerous vertical muscular fibres and few glandular cells present within rostellum. Layer of radial musculature separates rostellum from surrounding parenchyma. No glandular tissue observed around rostellum. Retractor muscular bundles attached at periphery of rostellum. Rostellar armament consists of 56-62 (n = 2) rostellar hooks arranged in 2 regular rows. Each hook triangular, with epiphyseal thickening on both handle and guard; blades curved. Anterior and posterior hooks of different shape, also differ slightly in BIUTERINA SPP. FROM THE IVORY COAST 159 TABLE 2. Comparison of some metrical and meristic data for Biuterina triangula (Krabbe, 1869). Host Turdus sp. “Turdus sp.” Anthus Anthus Turdus pilaris campestris leucophrys Locality Unknown Unknown Crimea, Ukraine Ivory Coast Source Krabbe (1869) Fuhrmann (1908) Kornyushin (1989) Present study Body: length (mm) = ~30 15-20 18.52 Body: maximum width - ~1000 450-470 770 Scolex: diameter - - 320-450 350-405 Suckers: diameter - - 120-170 100-140 Rostellum: diameter - - 110-160 102-110 Rostellar hooks: number 32 - 40 40-44 Rostellar hooks, length: anterior 55 DI 60 58-65 posterior 38-41 39 40 37-43 Testes: number - - 8-10 6-9 Cirrus-sac: length - 100 120 135-155 width - - 30-40 30-33 Vagina, copulatory part: length - - - 113-128 width - - - 13-18 Embryophore: diameter - - 27-35 - Oncosphere: diameter - - 20-25 - Embryonic hooks, length: central pair - - 17! - lateral pairs - - 171 - lLengths of the central and lateral embryonic hooks not given separately. 2Pregravid specimen. size. Anterior hooks 24-25 (24.5, n = 6) long; blade considerably longer than handle (Fig. 26). Posterior hooks 22-25 (23, n = 4) long; blade slightly longer than handle (Fig. 27). Neck 224 (n=l) wide, long; first proglottides appear at 860 from posterior end of suckers. Proglottides craspedote; juvenile, premature, mature (Fig. 28) and postmature (Fig. 30) and early pregravid proglottides (Fig. 31) wider than long; pregravid proglot- tides with more advanced development and gravid proglottides (Fig. 33) almost as long as wide or slightly longer than wide. Genital pores alternating irregularly, predomi- nantly in short series but long unilateral series may also occur, e.g., ...3, 1,3, 1, 1, 2, EO 3 5222 eee lemons IE, TION ele SIENNE eles alee 1,2,2,1,2,7...; pores open at junction of anterior and second thirds of lateral proglot- tis margin in mature and postmature proglottides and slightly anterior to middle of lat- eral proglottis margin in gravid proglottides. Genital atrium 41-70 (54, n = 10) deep, thick-walled, cylindrical, slightly expanded at its base (Fig. 29), with infundibular aperture 36-66 (52, n = 10) in diameter, provided with aggregation of intensely stain- ing cells. Atrial region might be surrounded by muscular bundles (sometimes not dis- tinct), thus assuming spherical appearance; contraction of these bundles results in for- mation of short but well-expressed genital papilla. Dorsal osmoregulatory canals 5-9 (n = 10) in diameter, without transverse anastomoses. Ventral osmoregulatory canals 17-40 (n = 10) in diameter, with transverse anastomoses along posterior margin of each proglottis. Genital ducts pass between poral osmoregulatory canals. 160 B. B. GEORGIEV & J. MARIAUX Fics 25-27 Biuterina pogoniuli sp. n. (25) Scolex. (26) Anterior rostellar hooks. (27) Posterior rostellar hook. Scale bars: 25, 100 um; 26, 27,25 um. Testes 12-17 (14,n = 15) in number, round or slightly oval, form compact group occupying whole width of posterior half of median field, laterally and postero-dorsally to ovary and dorsally and laterally to vitellarium, reach posterior proglottis margin; when fully developed, testes with diameter 68-84 (76, n = 10). External vas deferens convoluted in anterior poral part of median field, surrounded by prostate cells and forming together with them compact transversely-elongate body just adjacent to anterior proglottis margin; diameter of external vas deference 4-7 (5, n = 10). Cirrus- sac oblique, elongate, thick-walled, 165-192 x 49-63 (179 x 55, n = 15), with tapered poral end and widely rounded antiporal end, overlapping or just crossing poral osmoregulatory canals (Figs 28-30). Internal vas deferens convoluted in antiporal part of cirrus-sac, with diameter 8-13 (11, n = 10); intensely staining cells surround with- drawn cirrus. Partly evaginated cirrus cylindrical, 44-66 (n = 2) long, 14-15 (n = 2) in diameter; cirral armament (distinct in both evaginated cirrus and in canal of withdrawn organ) consisting of triangular spines 2-3 um in length. Ovary arcuate, with ends directed postero-laterally (Fig. 28), compact or slightly lobate, 179-268 (230, n = 10) wide. Vitellarium with irregular shape, compact, width 98-150 (119, n = 10). Seminal receptacle oval, situated dorsally to ovary and uterus; with rounded poral end; antiporal end gradually forms wide seminal duct. BIUTERINA SPP. FROM THE IVORY COAST 161 Fics 28-30 Biuterina pogoniuli sp. n. (28) Mature proglottis, dorsal view. (29) Genital ducts in postmature proglottis, dorsal view. (30) Early postmature proglottis, dorsal view. Scale bars: 28, 100 um; 29, 50 um; 30, 250 um. Vagina opens postero-dorsally to male pore; copulatory part almost as long as cirrus- sac, covered with thick cellular sleeve; vaginal canal thick-walled, with its distal part lined with long microtriches and conductive part long, thin (Fig. 29). 162 B. B. GEORGIEV & J. MARIAUX In mature proglottides, uterus thick-walled sac, transversely elongate, situated just anterior to ovary; around it, modification of medullary parenchyma forms pri- mordium of paruterine organ (Fig. 28). With further development, uterus expands in lateral direction, becoming arcuate (Fig. 30). In late post-mature proglottides, uterus assuming shape of horseshoe; paruterine organ surrounds entire uterus, its anterior part being thicker (Fig. 31). In gravid proglottides, uterus in general appearance horseshoe- shaped, lobate, entirely embedded by paruterine organ. Anterior part of paruterine organ fills entire median field (Fig. 33). Eggs with thin, irregular outer shell, situated closely to embryophore (Fig. 32). Embryophores thick-walled, spherical or slightly oval, with diameter 41-48 (44, n = 10). Oncospheres spherical, 28-32 (30, n = 10) in diameter. Embryonic hooks of central pair 20-22 long, with collar about their middle. Embryonic hooks of lateral pairs 17-19 long, with collar at about one third of their length. REMARKS: This species was previously reported as Paruterina sp. collected from 1 of 17 specimens of the host species studied (Mariaux, 1994). The identification was based on its correspondence with the generic concept of Paruterina Fuhrmann, 1906 as presented by Matevosyan (1969) and Schmidt (1986). The current concept of Paruterina is restricted to a small group of species parasitising Strigiformes, while Biuterina includes species with two symmetrical uterine sacs connected by an anterior isthmus, no matter whether the entire uterus is embedded by the paruterine organ or only its anterior part (Bona & Maffi, 1984; Kornyushin, 1989; Georgiev & Kornyushin, 1994). Therefore, the morphology of the specimens studied corresponds to the diagnosis of Biuterina (see Georgiev & Kornyushin, 1994). According to the structure of its uterus and paruterine organ, B. pogoniuli sp. n. corresponds to a group of species having a horseshoe-shaped uterus entirely embedded in the paruterine organ. This includes B. cordifera, a parasite of European and African insectivorous birds of the families Sylviidae and Turdidae (see above), and B. chlo- rurae (Rausch & Schiller, 1949), a parasite of North-American emberizid birds (Rausch & Schiller, 1949). The new species differs from B. cordifera by its consi- derably smaller rostellar hooks, greater number of testes and larger cirrus sac (Table 3). The ovary of B. pogoniuli is arcuate, while that of B. cordifera is more or less bi- winged (Georgiev er al., 2004; present study). Other differences are related to the cirral armament (present in the new species and lacking in B. cordifera), the shape of the cirrus sac (elongate and oval, respectively) and the number of rostellar hooks (Table 3). From B. chlorurae, the new species differs in its larger rostellar hooks, greater number of testes and larger cirrus sac (Table 3). Other species possessing rostellar hooks of a similar size are B. fuhrmanni Schmelz, 1941 and B. cylindrica Fuhrmann, 1908. However, these two species possess a paruterine organ situated anterior to the uterus (Fuhrmann, 1908; Schmelz, 1941; Georgiev et al., 2004). Biuterina petroniae sp. n. MATERIAL EXAMINED: MHNG INVE 16114, 49754 (formerly no 988.186, see Mariaux, 1994), 3 entire gravid specimens and 2 strobilar fragments stained and mounted in Canada bal- sam (2 slides) plus 2 scoleces mounted in Berlese’s medium (2 slides); from Petronia dentata BIUTERINA SPP. FROM THE IVORY COAST 163 (Sundevall) (Passeriformes: Ploceidae, Passerinae), collected on 19 February 1987 at Ouango- Fitini (9°37’N, 4°03’W). The host specimen has been deposited in the Ornithological Collection of the MHNG, no. 1773.018. Holotype: MHNG INVE 16114, an entire specimen in Canada balsam, mounted on the same slide with 2 strobilar fragments. Paratypes: MHNG INVE 49754, 2 entire specimens in Canada balsam; 2 scoleces in Berlese’s medium; 2 fragments mounted on the same slide with the holotype. ETYMOLOGY: The species name petroniae is a derivate of the generic name of the host. DESCRIPTION: Body gradually expands in posterior direction to level of pre- gravid proglottides and then with almost parallel lateral sides; gravid specimens 10.6- 13.1 mm (11.7 mm, n = 3) in length, consisting of 51-56 (54, n = 3) proglottides (10- 12 juvenile, 10-16 premature, 8-9 mature, 4-5 postmature, 13-15 pregravid and 2-3 gravid). Maximum width 0.63-0.70 mm (0.67 mm, n = 3) at pregravid proglottides. Scolex (Fig. 34) clearly outlined from neck by constriction, with maximum diameter 365-405 (680, n = 3) at level of posterior margin of sucker apertures; anterior part of scolex protruded, almost dome-shaped. Suckers situated on small but well-expressed pedicles (Fig. 34), cup-shaped, oval, 120-147 (131, n = 12) in diameter, with well- developed musculature. Rostellum sucker-like, cup-shaped, with diameter 90-112 (103, n = 3); numerous vertical muscular fibres and few glandular cells present within rostellum. Thick layer of radial musculature separate rostellum from surrounding tissue. Aggregations of glandular tissue present around rostellum. Rostellar armament consisting of 42-48, i.e. 42 (n = 2) or 48 (n = 2), rostellar hooks arranged in 2 regular rows. Each hook triangular, with epiphyseal thickening on both handle and guard; blades curved. Anterior and posterior hooks of different shape and size. Anterior hooks 38-40 (39, n = 12) long; blade slightly longer than handle (Fig. 35). Posterior hooks 28-30 (29, n = 11) long; blade and handle of almost equal length (Fig. 36). Neck 175 (n = 1) wide in narrowest part (at constriction between it and scolex); first proglottides appear at 220-340 (n = 3) from posterior end of suckers. Proglottides craspedote; juvenile, premature, mature (Fig. 37), postmature and early pregravid (Fig. 39) proglot- tides wider than long; pregravid proglottides with more advanced development and gravid proglottides (Fig. 40, 41) almost as long as wide or slightly longer than wide. Genital pores alternating irregularly, predominantly in short series, e.g. ... 1, 2,3, 2,5, 1,6,1,1,1,1,1, 1,5, 1; pores open slightly posterior to middle of lateral proglottis margin in mature proglottides (at border of anterior 38-45% of length of lateral proglot- tis margin) and at middle (48-55%) in gravid proglottides. Genital atrium 22-27 (24, n = 10) deep, thick-walled, with infundibular orifice and diameter of 10-14 (12, n = 10), provided with surrounding aggregation of intensely staining cells. No genital papilla. Dorsal osmoregulatory canals 4-6 (n = 10) in diameter, without transverse anasto- moses. Ventral osmoregulatory canals 10-45 (n = 10) in diameter, with transverse anas- tomoses along posterior margin of each proglottis. Genital ducts pass between poral os- moregulatory canals. Testes 7-10 (8, n= 11) in number, round or slightly oval, form compact group situated entirely in median field, laterally and postero-dorsally to ovary and dorsally and laterally to vitellarium; diameter when fully developed 43-53 (49, n = 10). External vas deferens convoluted in anterior poral corner of median field, surrounded by pro- 164 B. B. GEORGIEV & J. MARIAUX Fics 31-33 Biuterina pogoniuli sp. n. (31) Late postmature proglottis, dorsal view. (32) Egg. (33) Gravid proglottis. Scale bars: 31, 33, 250 um; 32, 50 um. static cells and together with them forming compact body; diameter of external vas deference 5-10 (8,n = 10). Cirrus-sac usually not reaching or only slightly overlapping poral osmoregulatory canals, 82-98 x 28-33 (92 x 31,n = 10), with rounded antiporal end and gradually tapering poral end; antiporal portion thick-walled; cirrus sac con- tains glandular cells surrounding canal of withdrawn cirrus and ejaculatory duct (Fig. 38). Internal vas deferens convoluted in antiporal part of cirrus-sac. No evaginated cirrus in material studied; no cirral armament observed in canal of withdrawn cirrus. Ovary compact, bi-winged, occupies most of width of median field; wings rounded, connected by narrow isthmus (Fig. 37). Vitellarium median, oval or irregu- BIUTERINA SPP. FROM THE IVORY COAST 165 De, aS = NI SS > Al / a Pa | ii GZ | a SS Of it Sa II tiny i | 36 34 37 Fics 34-37 Biuterina petroniae sp. n. (34) Scolex. (35) Anterior rostellar hooks. (36) Posterior rostellar hooks. (37) Mature proglottis, dorsal view. Scale-bars: 34, 37, 100 Im; 35, 36, 20 um. larly suboval, slightly lobed or compact, 45-70 (57, n = 10) wide. Mehlis’ gland globular, 18-24 ( 20, n = 10). Seminal receptacle dorsal to ovary and uterus, with rounded poral end with diameter 25-35 (31, n = 8) and antiporal end gradually forms wide seminal duct. Vagina opens posteriorly to male pore; copulatory part 137-162 (148, n = 8) long, consists of thin-walled vaginal canal with luminal diameter 6-10 (8, 166 B. B. GEORGIEV & J. MARIAUX Fics 38-39 Biuterina petroniae sp. n. (38) Genital ducts, dorsal view. (39) Pregravid proglottis, dorsal view. Scale bars: 38, 50 um; 39, 200 um. n= 8) and thin, slightly expressed cellular sleeve; diameter of cellular sleeve 18-24 (21, n = 8); conductive part short, thin (Fig. 38). In mature proglottides, uterus a thick-walled sac with irregular shape situated antero-dorsal to ovary; around and anteriorly to it, modification of medullary paren- chyma forms primordium of paruterine organ (Fig. 37). With further development, uterus expands in lateral direction. In early pregravid proglottides (Fig. 39), uterus in form of single, transversely-elongate sac in most posterior part of median field; its BIUTERINA SPP. FROM THE IVORY COAST 167 Fics 40-41 Biuterina petroniae sp. n. (40, 41) Gravid proglottides showing the final stages of the develop- ment of the uterus and paruterine organ. Scale bars: 40-41, 200 um. lateral ends gradually differentiate as 2 distinct sacs divided by median constriction (Fig. 40). Paruterine organ consists of 2 parts; conical formation consisting predomi- nantly of fibrilar tissue just anterior to uterine sac and modification of medullar parenchyma situated near anterior proglottis margin (Figs 39, 40); in addition, all medullary parenchyma are compacted. In most developed gravid proglottides available, eggs pass from uterine sacs into paruterine organ (Fig. 41). Eggs with thin, often indistinct, outer shell closely covering embryophore. Embryophore thick-walled, oval, with diameter 38-45 (42,n = 10). Oncosphere spherical, with diameter 27-34 (31, n = 10). Embryonic hooks of central pair 17-18 long, of lateral pairs 12-13 long; intralateral and extralateral embryonic hooks of the same shape. REMARKS: The present material was reported as Biuterina sp. by Mariaux (1994). Judging by the size of the rostellar hooks, it is close to B. macrancistrota (Fuhrmann, 1908), B. trapezoides Fuhrmann, 1908, B. campanulata (Rudolphi, 1819) and B. landsdowni Malhotra & Capoor, 1987 (Table 3). The latter species, a parasite of Upupa epops L. in India (Malhotra & Capoor, 1987), is the only member of the genus for which the posterior hooks are reported to be longer than anterior hooks. The structure of its uterus and paruterine organ and the shape of the rostellar hooks, as illustrated by Malhotra & Capoor (1987), very much resemble those of Neyraia Joyeux B. B. GEORGIEV & J. 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B. GEORGIEV & J. MARIAUX & Timon-David, 1934 (see Georgiev & Kornyushin, 1994), a genus which includes specific parasites of hoopoes (Upupidae and Phoeniculidae); therefore, we believe that the affiliation of this species to Biuterina requires further confirmation. Nevertheless, B. petroniae differs from B. landsdowni by the considerably smaller number of testes and a substantial difference in the length of the posterior rostellar hooks (Table 3). B. campanulata and B. trapezoides are poorly known Neotropical species; the only sources concerning their morphology are their original descriptions (Fuhrmann, 1908). They are both characterised by a considerably smaller number of rostellar hooks (26 and about 30, respectively) than in the present material (42-48). Our worms are most similar to B. macrancistrota (for a redescription, see Mariaux & Vaucher, 1989). This species has a body about twice the length of our material (22 versus 10-13 mm) and consists of many more proglottides (180 versus 51-56). A substantial part of its cirrus sac is situated in the median field in mature proglottides, while in the new species this organ occurs in the lateral field. Another difference is the shape of ovary, which is clearly bi-winged in the new species and a massive, rather transversely elongate oval structure in B. macrancistrota. On the basis of these differences, we consider that the present material is a new species. DISCUSSION In addition to the material described above, Mariaux (1994) reported a species of Biuterina from Sylvietta virens Cassin (Sylviidae). This material does indeed belong to this genus and might represent a new species. However, the specimens available are fragmented and do not permit the preparation of a complete description. The concept of Biuterina adopted in the present article corresponds to those presented in previous studies (Bona & Maffi, 1984; Kornyushin, 1989; Georgiev & Kornyushin, 1994). It is defined by the presence of two symmetrical uterine sacs connected by an anterior isthmus, no matter whether the entire uterus is embedded by the paruterine organ or only its anterior part. However, whether this is a synapomorphy of a monophyletic Biuterina or not will be a subject for further studies. Furthermore, other paruterinid genera (Triaenorhina Spasskii & Shumilo, 1965, Neyraia Joyeux & Timon-David, 1934 and Spasskyterina Kornyushin, 1989) are characterised by two symmetrical uterine sacs, although of various shape. Currently, we recognise 34 species as belonging to this genus (Table 3). The position of two of them in Biuterina is doubtful. B. lansdowni Malhotra & Capoor, 1987, from Upupa epops L. in India (Malhotra & Capoor, 1987), has hooks and gravid proglottides strongly resembling Neyraia intricata (Krabbe, 1878) from the same host species. Whereas B. diversihamata Sawada, Hassan & Saleen, 1994, from Coracias garrulus L. in Iraq (Sawada et al., 1994), we suspect to be a synonym of Triaenorhina rectangula (Fuhrmann, 1908). It is our opinion that these two taxa represent species inquirendae, but they are retained in the key below, pending a re-examination of the type material. Six species have previously been considered members of Biuterina (see Matevosyan, 1969; Schmidt, 1986) and are currently placed in other genera. B. rec- tangula Fuhrmann, 1908 from Coracias garrulus L. (Europe?) was selected as the type BIUTERINA SPP. FROM THE IVORY COAST 171 species of Triaenorhina (see Spasskii & Shumilo, 1965). Three cestode species from the same host were recognised as synonyms of T. rectangula; these are B. coracii Chiriac, 1963 from Romania, B. garrulae (Matevosyan, 1950) from Smolenskaya Oblast’, Russia, and B. uzbekiensis Matevosyan, 1964 from Uzbekistan (synonymy after Spasskaya & Spasskii, 1971; Kornyushin, 1989; Georgiev & Gibson, 2006). In addition, B. dunganica Skrjabin, 1914 from Oriolus oriolus L. in Kazakhstan was transferred to Spasskyterina Kornyushin, 1989 (see Kornyushin, 1989) and B. meggitti (Johri, 1931) from bucerotid birds in India and Somalia is currently recognised as a species of Triaenorhina (see Kornyushin, 1989; Georgiev & Gibson, 2006). Recent studies have provided new morphological data for most of the Old- World species and re-evaluated their validity (Kornyushin, 1989; Mariaux & Vaucher, 1989; Georgiev et al., 2002, 2004). However, most of the New-World species are known from their original descriptions only and are in need of taxonomic revision. Nevertheless, we propose the following key based on our current knowledge of Biuterina spp. (B. motacillabrasiliensis and B. motacillacayanae are poorly described parasites of Neotropical passerines and lack reliable distinguishing features. KEY TO THE SPECIES OF BIUTERINA (See Table 3 for authorities for species names) la. Parasites of Paradisaeidae in the Australian Region .................... 2 IbaggpeRarasitessofiothenbirds asta 2 ea 3 2a. Testes -30 in number. Genital atrium without distinct dilatory muscles Eee eas en LEN EL Nears e MAS N B. clavula 2b. Testes ~15 in number. Genital atrium with distinct dilatory muscles Di BOP Bele, NE NORCO EER DREHEN RIO ERNEST AN gata Cae Wea B. mertoni Sal Anterior rostellar hooks < 17 long; if anterior rostellar hooks 17 long thenstotalenumber,ofrostellanhooksis 32 u. m RR 4 3b. Anterior rostellar hooks 017 long; if anterior rostellar hooks 17 long or slightly longer then total number of rostellar hooks O40................ 5 tae Rostellanhooks 2A: innumben ter B. globosa 4b. Rostellar hooks 30-32 in number se oh ya A ga pe igs A eS ul et B. motacillabrasiliensis and B. motacillacayanae Sa. Uterus entirely embedded in paruterine organ in pregravid proglottides ..... 6 Sb. Paruterine organ situated anterior to uterus in pregravid proglottides, not entirely embed dim sinters RE ec ee Dee 8 6a. Developing uterus horseshoe-shaped, with posteriorly directed lateral EXtTEMMIIIES". i E e voice IO REN RE ENORME 7 6b. Developing uterus consisting of two rounded lateral chambers connected anteriorly by narrow, branched isthmus ............ B. passerina Ja. Rostellar hooks 26-30 in number; anterior hooks 056 long ...... B. cordifera 7b. Rostellar hooks 56-62 in number; anterior hooks 24-25 long . . . . B. pogoniuli 7c. Rostellar hooks 40-42 in number; anterior hooks 20 long ....... B. chlorurae $a, Anterionrostellamhooks)=aslonsviaa TRE RR 9 éb AN Anterior rostellarmhooks SSN ONE ae See: 21 172 9a. Ob. 10a. 10b. lla. lb. 12a. 12b. 13a. 185% 14a. 14b. 15a. 15b} 16a. 16b. 17a. 17b. 18a. 18b. 19a. 19b: 20a. 20b. 21a. 21b. DIR. 22b. 23a. 236: 24a. B. B. GEORGIEV & J. MARIAUX Numberoferostellar hooks < 2052. Sn el eee 10 Numberof rostellar hooks 26 +40 =. EEE 11 Number of rostellar hooks 14; anterior rostellar hooks 56-60 long; cirrus SACEHMAOMONPLA RT POE. oe RR OBS a ee eee B. erithaci Number of rostellar hooks 18-20; anterior rostellar hooks 44-48 long; CITUSISAC 90 long wv: Sees Py a. PPS RCE B. fallax Amtenorrostellan hooks} 75-Solong RO B. africana Anterior rostellar/hooks'= 70 long Li RESSE 12 Anterior rostellarshooks 58-66 long. Pr oe ee eee 13 Anterionrostellar hooks 38-49 long MC eee eee 14 Rostellar hooks 34-36 in number; testes 15-18 in number; cirrus sac 86- lOSHlOn gry SAYS NIEREN EEE B. morgani Rostellar hooks 40-44 in number; testes 6-9 in number; cirrus sac > 120 ON ER AR ELE RUN Sie, MEER EA EEE A B. triangula Rostellauhooksispine=lke mes EEE B. lansdowni Rostellar hooks triangular, of typical shape for Biuterina ............... 15 Rostellarhooks = 30. in’number 225 CEE EROE NE Des SE I 16 Rostellar hooks > 35 in number... . 22... sa 17 Rostellar hooks 26 in number, anterior rostellar hooks 43-46 long; paru- terine organ wide, occupying almost entire width of median field PIPES ee NEN SLO ANS, AL EEE B. campanulata Rostellar hooks ~30 in number; anterior rostellar hooks 41 long; paru- tenmneorganun middle ofmedian field ee Sam. Sener ae B. trapezoides Anternorrostellar hooks':45 long Tit, Er SE N eee 19 Gravid strobila consisting of ~180 proglottides; cirrus sac crosses poral osmoregulatory canals and substantial part of it situated in median field BE DE ESS B. macrancistrota Gravid strobila consisting of -50-60 proglottides; cirrus sac in lateral Helden Au A RATTEN NEE ED EN B. petroniae Posterior rostellar hooks not longer than 55% of length of anterior ros- tellariho0ksı fa 2 EN ER RE ES B. planirostris Posterior rostellar hooks longer than 60% of length of anterior rostellar Hook MET DIR RSS PLOT SOG BA RI ee 20 Blade of rostellar hooks 4-5 times shorter than handle .... . B. diversihamata Blade of rostellar hooks slightly longer than handle or of same length RPMS ene BONS LUE TONI SOA IE NE B. sobolevi Rostellar hooks -20#n number 2... 222 E sea eee B. distincta Rostellarlhooks'’> 30, ininumbert. 4 RR ORE INR 22 Anteriorrostellar hooks!= 30 lons ens. 225 5 SS ae eee B. reynoldsi Anterior rostellar hooks < 30 long. Ir SS 23 Rostellar hooks >56 ininumberi ia: . 29 255 Re Sa SE 24 Rostellar hooks)|=52 infaumber SCE EIER 21 Rostellum consists of weak musculature and strongly developed glan- dulartissue:.. 2.2.2... ll Pe NT SR PR RENE B. quelea BIUTERINA SPP. FROM THE IVORY COAST 173 24b. Rostellum consists of strongly developed musculature, with no glandu- lar tissue or glandular tissue weakly developed ....................... 25 25a. Paruterine organ in pregravid proglottides occupies entire width of medianimel desi ar Bie Sek ee ne: LIA B. ugandae 25b. Paruterine organ distinctly narrower than width of median field .......... 26 26a. Parasite of Furnariidae in the Neotropics (Brazil) .......... B. trigonacantha 26b'Parasite OP Dicrurida cin india EM A B. singhi 2.0 ce Parasite of lBanndae- in therBalasarcher Wr we ee B. collurionis Aa BERostellar/nooks AU NUMBER en ee. 28 27b. Rostellar hooks 048 (possibly, as exception, 42 in B. fuhrmanni) ........ 29 2825 Anteriortostellarhooks27a10N 94 ne ara anne ee B. meropina 2°bAnterionrostellas hooks;<20Jlongrrr a ae N ee B. dicruri 20 a Miestes.numberEI2I ANY KIN 20 RI ee a B. fuhrmanni PO DIRETE stestnumDer al. ERE ORI REL) E RIMA Ne a 30 30a. Rostellar hooks 52 in number; anterior rostellar hooks 23-25 long D A I AT AS e SE o A IO LL a ba CN U8 B. cylindrica 30b. Rostellar hooks 48 in number; anterior rostellar hooks 19-22 long ET ee I I RI IR B. zambiensis ACKNOWLEDGEMENTS We are grateful to Dr D.I. Gibson (London) for reading the manuscript. The Natural History Museum, Geneva, funded research visits of the senior author, enabling Joint studies. REFERENCES BONA, F. 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The genus Biuterina Fuhrmann, 1902 (Cestoda, Paruterinidae) in the Old World: redescriptions of three species from Palaearctic Passeriformes. Systematic Parasitology 57: 67-85. HOWARD, R. & Moore, A. 1980. A complete checklist of the birds of the world. Oxford University Press, Oxford, 701 pp. JOYEUX, C. & BAER, J.-G. 1928. Cestodes. (pp. 17-54). In: JOYEUX, C., GENDRE, E., & BAER, J.- G. (eds). Recherches sur les helminthes de 1’ Afrique occidentale française. (Collection de la Société de Pathologie Exotique, Monographie II.), Masson et C'€, Paris. KORNYUSHIN, V. V. 1989. [Fauna of Ukraine. Volume 33. Monogenea and Cestoda. Part 3. Davaineoidea. Biuterinoidea. Paruterinoidea.] Naukova Dumka, Kiev, 252 pp. (In Russian). KRABBE, H. 1869. Bidrag til Kundskab om Fuglenes Baendelorme. Videnskabelige Selskab Skrifter, 5. Raekke, Naturvidenskabelig og Mathematisk Afdel 8: 251-389. MALHOTRA, S. K. & CAPOOR, V. N. 1987. Avian cestodes of the Himalayan ecosystems. II. 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Species belonging to families Davaineidae and Biuterinidae. Indian Journal of Helminthology 11: 1-24. SPASSKAYA, L. P. & SPASSKII, A. A. 1971. [Cestodes of birds in Tuva.] Shtiintsa, Kishinev, 252 pp. (in Russian). SPASSKII, A. A. & SHUMILO, R. P. 1965. [The phenomenon of the postlarval development of the rostellum and the hooks in the cestodes of the genus Triaenorhina, n. gen. (Paruterinidae).] Doklady Akademii Nauk SSSR 164: 1436-1438. (In Russian). SUDARIKOV, V. E. 1950. [New cestode species from the basin of Middle Volga.] Trudy Gel’mintologicheskoy Laboratorii Akademii Nauk SSSR 3: 142-151. (In Russian). REVUE SUISSE DE ZOOLOGIE Tome 114 — Fascicule 1 LOURENCO, Wilson R. & QI, Jian-Xin. Additions a la faune des scorpions de l’Etat du Amapa, Brésil (Chelicerata, Scorpiones) ............ KOTTELAT, Maurice, BARBIERI, Roberto & STOUMBOUDI, Maria Th. Aphanius almiriensis, a new species of toothcarp from Greece (dteleostei4Gyprinodentidas) 3.2. aus na en DI GIULIO, Andrea, KAUPP, Andreas, FATTORINI, Simone, VIGNA TAGLIANTI, Augusto & NAGEL, Peter. Pupal morphology in the subfamily Baussinaei(@oleopteray@arabidae)er „nm ae Novak, Tone, SIVEC, Ignac, JANZEKOVIC, Franc & CHRISTIAN, Erhard. Chionea austriaca in caves and artificial galleries of Slovenia WipteraslEiimonudae)ie ss ek ie a. ne. Xu, Xiang & Li, Shugiang. A new genus and species of the spider family Agelenidae from western Sichuan Province, China (Arachnida: ATANE AS) RSS IA IO TERE WITTMANN, Karl J. Continued massive invasion of Mysidae in the Rhine and Danube river systems, with first records of the order Mysidacea (Crustacea: Malacostraca: Peracarida) for Switzerland ........... AMIET, Jean-Louis. Les Phlyctimantis et Kassina du Cameroun (Amphi- biayAnura,Hiyperolüdae)ery. ar en ne n POMORSKI, Romuald J. & KAPRUS, Ighor. Redescription of Protaphorura octopunctata (Tullberg, 1876) and Protaphorura quadriocellata (Gisin, 1947) with description of two new related species from Siberia and Europe (Collembola: Onychiuridae) ................ GEORGIEV, Boyko B. & MARIAUX, Jean. Cestodes of the genus Biuterina Fuhrmann, 1902 (Cyclophyliidea: Paruterinidae) from passeriform and piciform birds in the Ivory Coast, with a key to the species of the OT Sper ar macs cist Nie tenses kau eer heeded ee end: 33-48 49-57 59-64 65-86 87-126 127-139 141-174 REVUE SUISSE DE ZOOLOGIE Volume 114 — Number 1 LOURENÇO, Wilson R. & Qi, Jian-Xin. Addition to the scorpion fauna of the Amapä State in Brazil (Chelicerata, Scorpiones) ............. KOTTELAT, Maurice, BARBIERI, Roberto & STOUMBOUDI, Maria Th. Aphanius almiriensis, a new species of toothcarp from Greece (FeleosteriGypunodontida ERE CERO OE Di GruLio, Andrea, KAUPP, Andreas, FATTORINI, Simone, VIGNA TAGLIANTI, Augusto & NAGEL, Peter. Pupal morphology in the subfamily Paussinae (Coleoptera: Carabidae) Hat. Wat. were Suse NovaK, Tone, SIVEC, Ignac, JANZEKOVIC, Franc & CHRISTIAN, Erhard. Chionea austriaca in caves and artificial galleries of Slovenia (Diptera9lEimoniidac ees Erna ER Cee Xu, Xiang & Li, Shugiang. A new genus and species of the spider family Agelenidae from western Sichuan Province, China (Arachnida: INTANECAC) Richtee class, Stations NAT SSR i BESTE ye WITTMANN, Karl J. Continued massive invasion of Mysidae in the Rhine and Danube river systems, with first records of the order Mysidacea (Crustacea: Malacostraca: Peracarida) for Switzerland ........... AMIET, Jean-Louis. The Phlyctimantis and Kassina of Cameroon (Amphi- bias AnurasHyperolidae).2 00. rm ram NC ee Ber POMORSKI, Romuald J. & KAPRUS, Ighor. Redescription of Protaphorura octopunctata (Tullberg, 1876) and Protaphorura quadriocellata (Gisin, 1947) with description of two new related species from Siberia and Europe (Collembola: Onychiuridae) ................ GEORGIEV, Boyko B. & MARIAUX, Jean. Cestodes of the genus Biuterina Fuhrmann, 1902 (Cyclophyliidea: Paruterinidae) from passeriform and piciform birds in the Ivory Coast, with a key to the species of the PETS HN Pr Math aan MI EN he tok ONE SAE ES SEE Indexed in CURRENT CONTENTS, SCIENCE CITATION INDEX 49-57 59-64 65-86 87-126 127-139 141-174 PUBLICATIONS DU MUSEUM D’HISTOIRE NATURELLE DE GENEVE CATALOGUE DES INVERTEBRES DE LA SUISSE, N95 1-17 (1908-1926) ........ serie Fr. (prix des fascicules sur demande) REVUE DE PALEOBIOLOGIE rn een Echange ou par fascicule Fr. LE RHINOLOPHE (Bulletin du centre d’étude des chauves-souris) . ...... par fascicule Fr. THE EUROPEAN PROTURA: THEIR TAXONOMY, ECOLOGY AND DISTRIBUTION, WITH KEYS FOR DETERMINATION Jo INOS SAPORE NOIA ope meron cable sole aot Fr. CLASSIFICATION OF THE DIPLOPODA RBICAHOFEMANSZITIP IDEE ET sesh RE MIS Te Fr. LES OISEAUX NICHEURS DU CANTON DE GENÈVE P. GEROUDET, C. GUEX & M. MAIRE 35iip-mombreuses cartesiet tigures, 19831. nen... ei en Fr. CATALOGUE COMMENTÉ DES TYPES D'ECHINODERMES ACTUELS CONSERVES DANS LES COLLECTIONS NATIONALES SUISSES, SUIVI D'UNE NOTICE SUR LA CONTRIBUTION DE LOUIS AGASSIZ A LA CONNAISSANCE DES ECHINODERMES ACTUELS IMETANGOUXAOM ID MISI wera cure) sity else lesen III OA II Fr. RADULAS DE GASTEROPODES LITTORAUX DE LA MANCHE (COTENTIN-BAIE DE SEINE, FRANCE) NEEINED I) WiUESISAIKeIMAREDAS 62) pr OOM ae ie nal a Fr. GASTROPODS OF THE CHANNEL AND ATLANTIC OCEAN: SHELLS AND RADULAS NA EINED SE WiUESTHOeKIMARED AGW OO Da N eile eines ee el Fr. O. SCHMIDT SPONGE CATALOGUE R. DESQUEYROUX-FAUNDEZ & S.M. STONE, 190 p., 1992 ....................... . Fr. ATLAS DE REPARTITION DES AMPHIBIENS ET REPTILES DU CANTON DE GENEVE A. KELLER, V. AELLEN & V. MAHNERT, 48 p., 1993 ............................ Fr. THE MARINE MOLLUSKS OF THE GALAPAGOS ISLANDS: A DOCUMENTED FAUNAL LIST CINICO SE O Fr. NOTICE SUR LES COLLECTIONS MALACOLOGIQUES DU MUSEUM D’HISTOIRE NATURELLE DE GENEVE DÉCACAILLIEZ SAID. III e I A I er I VA NOAA tot ECS Fr. PROCEEDINGS OF THE XIIIth INTERNATIONAL CONGRESS OF ARACHNOLOGY, Geneva 1995 (ed. V. MAHNERT), 720 p. (2 vol.), 1996 ...... Fr. CATALOGUE OF THE SCAPHIDINAE (COLEOPTERA: STAPHYLINIDAE) (Instrumenta Biodiversitatis I), I. LOBL, xii + 190 p., 1997...................... Fr. CATALOGUE SYNONYMIQUE ET GÉOGRAPHIQUE DES SYRPHIDAE (DIPTERA) DE LA REGION AFROTROPICALE (Instrumenta Biodiversitatis U), H. G. DIRICKX, x +187 p.,1998 ..... ............ Fr. A REVISION OF THE CORYLOPHIDAE (COLEOPTERA) OF THE WEST PALAEARCTIC REGION (Instrumenta Biodiversitatis II), S. BOWESTEAD, 203 p., 1999 ................... Fr. THE HERPETOFAUNA OF SOUTHERN YEMEN AND THE SOKOTRA ARCHIPELAGO (Instrumenta Biodiversitatis IV), B. SCHATTI & A. DESVOIGNES, SD OO AA INNI ROIO I NIE RE Fr. PSOCOPTERA (INSECTA): WORLD CATALOGUE AND BIBLIOGRAPHY (Instrumenta Biodiversitatis V), C. LIENHARD & C. N. SMITHERS, XL AS ND 2002 a OLA EA LR Fr. REVISION DER PALAARKTISCHEN ARTEN DER GATTUNG BRACHYGLUTA THOMSON, 1859 (COLEOPTERA, STAPHYLINIDAE) (1. Teil) (Instrumenta Biodiversitatis VI), G. SABELLA, CH. BUCKLE, V. BRACHAT SACIBESUEHET, VIA) 283,92, 2004 ROTTE TE EE Fr. PHYLOGENY, TAXONOMY, AND BIOLOGY OF TEPHRITOID FLIES (DIPTERA, TEPHRITOIDEA) Proceedings of the “3rd Tephritoid Taxonomist’s Meeting, Geneva, 19.-24. July 2004” (Instrumenta Biodiversitatis VII). B. 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Primary types must be deposited in a museum or similar institution. In taxonomic papers the species heading should be followed by synonyms, material examined, description, distribution, and comments. All material examined should be listed in similar, compact and easily intelligible format; the information should be in the same language as the text. Sex sym- bols should be used rather than “male” and “female” (text file: $ = d,£= @). Discussion. This should not be excessive and should not repeat results nor contain new information, but should emphasize the significance and relevance of the results reported. References. The author-date system (name-year system) must be used for the citation of references in the text, e.g. White & Green (1995) or (White & Green, 1995). For references with three and more authors the form Brown et al. (1995) or (Brown et al., 1995; White et al., 1996) should be used. In the text authors’ names have to be written in standard type face. However, in the list of references they should be formatted in SMALL CAPITALS (see below). The list of references must include all publications cited in the text and only these. References must be listed in alphabetical order of authors, in the case of several papers by the same author, the name has to be repeated for each reference. The title of the paper and the name of the journal must be given in full in the following style: PENARD, E. 1888. Recherches sur le Ceratium macroceros. These, Genève, 43 pp. PENARD, E. 1889. Etudes sur quelques Héliozoaires d’eau douce. Archives de Biologie 9: 1-61. MERTENS, R. & WERMUTH, H. 1960. Die Amphibien und Reptilien Europas. Kramer, Frankfurt am Main, XI + 264 pp. HANDLEY, C. O. Jr 1966. Checklist of the mammals of Panama (pp. 753-795). In: WENZEL, R. L. & TIPTON, V. J. (eds). Ectoparasites of Panama. Field Museum of Natural History, Chicago, XII + 861 pp. Tables. These should be self-explanatory, not integrated in the text-file, with the title at the top, organised to fit 122 x 180 mm, each table on a separate sheet and numbered consecutively. Figures. These may be line drawings or half tones, not integrated in the text-file, and all should be numbered consecutively. Figures should be arranged in plates which can be reduced to 122 x 160 mm. Drawings and lettering should be prepared to withstand reduction. Magnification should be indicated with scale lines. Authors should refrain from mixing drawings and half tones. Originals of figures (ink drawings, photographs, slides) should be submitted together with the revised version of the accepted manuscript. Original drawings will not be returned automatically. The Revue suisse de Zoologie declines responsibility for lost or damaged slides or other documents. If scanned figures are submitted on CD, this should be clearly indicated on the print-out. Scanned line drawings must be saved as TIF files in bitmap mode with a resolution of at least 600 dpi. Half tone illustrations and photos must have at least 300 dpi reso- lution. Legends to figures. These should be typed in numerical order on a separate sheet. Proofs. Only page proofs are supplied, and authors may be charged for alterations (other than printer’s errors) if they are numerous. Offprints. The authors receive a total of 25 offprints free of charge; more copies may be ordered at current prices when proofs are returned. Correspondence. All correspondence should be addressed to Revue suisse de Zoologie, Muséum d’histoire naturelle, CP 6434, CH-1211 Genève 6, Switzerland. Phone: +41 22 418 63 33 - Fax: +41 22 418 63 01. E-mail: danielle.decrouez@ville-ge.ch Home page RSZ: http://www. ville-ge.ch/musinfo/mhng/page/rsz.htm 3 9088 01325 4958