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Révision Du Genre Geophagus

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Révision Du Genre Geophagus Académie royale des Sciences d'Outre-Mer :iasse des Sciences Naturelles et Médicales, N.S., XIX-3, Bruxelles, 1975 Révision du genre Geophagus (Pisces Cichlidas) PAR J.-P. GOSSE Chef de Travaux à l'Institut royal des Sciences naturelles de Belgique Mémoire publié avec le concours du Fonds Leopold III pour l'Etude et la Conservation de la Nature 1000 F .oninklijke Academie voor Overzeese Wetenschappen Klasse voor Natuur- en Geneeskundige Wetenschappen, N.R., XIX-3, Brussel, 1975 Académie royale des Sciences d'O utre-M er Classe des Sciences Naturelles et Médicales, N.S., XIX-3, Braxelles, 1975 Révision du genre Geophagus (Pisces Cichlidas) PAR J.-P. GOSSE Chef de Travaux à l'Institut royal des Sciences naturelles de Belgique Mémoire publié avec le concours du Fonds Léopold III pour l'Etude et la Conservation de la Nature Koninklijke Academie voor Overzeese Wetenschappen Klasse voor Natuur- en Geneeskundige Wetenschappen, N.R., XIX-3, Brussel, 1975 Mémoire présenté à la Séance du 26 juin 1973 Rapporteurs : MM. M. POLL, F. EVENS et P. BRIEN D/1975/0149/3 RESUME L'examen et la mensuration des spécimens types et de grandes séries d'exemplaires des espèces du genre Geophagus et de cer• tains genres voisins permet une meilleure compréhension de ce groupe de Cïchlidae néo-tropicaux. Cette étude a porté sur les espèces généralement comprises dans le genre Geophagus, c.a.d. les Cichlides sud-américains portant les branchiospines le long du bord libre d'un lobe charnu à la partie supérieure du premier arc branchial et dont la ligne latérale supérieure est écartée de la base de la nageoire dorsale par au moins deux rangées et demie d'écaillés. Ces espèces doivent être séparées en trois genres distincts, qui se caractérisent, entre autres, par des particularités ostéologiques: Geophagus HECKEL, 1840 - Un seul supraneural; Biotodoma EIGENMANN & KENNEDY, 1903 {Mesops GÜN- THER, 1872) - deux supraneuraux; Gymnogeophagus RIBEIRO, 1918 - Pas de supraneural, un processus épineux devant le premier ptérygophore dorsal. Le genre Geophagus comprend actuellement dix espèces, dont une espèce nouvelle, Geophagus harreri, découverte dans le bas• sin du Maroni. Les espèces du genre Geophagus sont réparties dans le bassin amazonien et dans les bassins guyanais, dans la région centrale du Panama, en Colombie et dans les bassins côtiers du sud-est brésilien. Geophagus steindachneri est synonyme de Geophagus hondae et Geophagus mapmtensis est synonyme de Geophagus jurupari. Outre ces nouvelles synonymies nous signalons que les spécimens du Zoologisches Museum de Berlin portant les indications « Type de Geophagus tuberosus MÜLLER & TROSCHEL » « types de Geo• phagus olfersi MULLER & TROSCHEL », n'ont jamais été décrits et doivent être rapportés à Geophagus brasiliensis. Le genre Biotodoma comprend deux espèces du bassin amazo• nien, des Guyanes et du Haut Orénoque: Biotodoma cupido (HECKEL, 1840) et Biotodoma wavrini (GOSSE, 1963). 4 RÉVISION DU GENRE GEOPtlAGUS (PISCES CICMLIDiï) Le genre Gymnogeophagus est redéfini, il compte quatre espè• ces du bassin du Parana et des bassins côtiers de l'Uruguay et du Rio Grande do Sul. La radiographie et l'examen des spécimens types permet d'établir ou de corriger les synonymies suivantes: Gymnogeophagus cyanopterus est synonyme de Geophagus bal- zanii, Geophagus rhabdotus n'est pas synonyme de G. hrasilien- sis, Geophagus camurus n'est pas synonyme de G. gymnogenys mais de G. rhabdotus; G. bucephalus, G. labiatus, G. scymno- philus et G. pygmaeus ne sont pas des synonymes de G. brasi- liensis mais bien de G. gymnogenys. Pour chacune des espèces de ces trois genres, les spécimens holotypes ou les spécimens choisis comme lectotypes ont été redécrits et figurés. Les résultats des mensurations des séries de spécimens de chaque espèce sont portés en tableaux avec leurs données statistiques. Pour les espèces à large répartition {G. surinamensis, G. jurupari) les exemplaires ont été groupés d'après leur origine géographique de manière à tenter une com• paraison de ces populations par divers tests statistiques. Les différences significatives observées par ces tests sont cependant jugées insuffisantes pour la création de sous-espèces. Les clés de détermination et les tableaux figurant en annexe seront utiles pour séparer ces trois genres et leurs diverses espèces. Etudiant la phylogénie de ces trois genres par l'examen du squelette dorsal, de la dentition et la répartition géographique on doit conclure que le genre Biotodoma est le plus primitif et que ces genres et espèces se sont différenciés à partir du bassin amazonien, le plus souvent par isolement géographique. RÉVISION DU GENRE GEOPHAGUS ( PISCES CICHLID^Îl) 5 SUMMARY Examination and mensuration of the types and of a large number of other specimens of species of the genus Geophagus and of certain related genera has resulted in a better under• standing of this group of neotropical cichlids. This study has been carried out on species generally included in the genus Geophagus, that is to say the South American cichlids bearing the gill rakers along the free border of a fleshy lobe of the upper part of the first branchial arch, and of wich the upper lateral line is separated from the base of the dorsal fin by at least two and a half rows of scales. These species can be separated into three distinct genera which are characterised by, among others, the following osteological attributes: Geophagus MECKEL, 1840 - A single supraneural; Biotodoma EIGENMANN & KENNEDY, 1903 {Mesops GÜN- THER, 1872) - Two supraneurals; Gymnogeophagus RIBEIRO, 1918 - No supraneural, a spinous process in front of the first dorsal pterygophore. The genus Geophagus includes ten species of which a new species, Geophagus harreri, has been found in the Maroni Basin. The species of Geophagus are distributed throughout the Ama• zon and Guyanan basins, in the central region of Panama, in Columbia, and in the coastal drainage areas of South East Brazil. Geophagus steindachneri is a synonym of Geophagus hondae and Geophagus mapiritensis is a synonym of Geophagus juru- pari. In addition to these new synonymies we report that speci• mens in the Berhn Zoological Museum carry the designations « Type of Geophagus tuherosus MÜLLER & TROSCHEL », « types of Geophagus olfersi MÜLLER & TROSCHEL », which have never been described and should be assigned to Geophagus brasiliensis. The genus Biotodoma includes two species of the Amazon Basin, the Guyanas and upper Orinocco: Biotodoma cupido (MECKEL, 1840) and Biotodoma wavrini (GOSSE, 1963). The genus Gymnogeophagus is redefined. It includes four species of the Parana Basin and of the coastal regions of Uru• guay and Rio Grande do Sul. 6 RÉVISION DU GENRE GEOPHAGUS (PISCES CICHLID^) Radiography and the examination of type specimens has made it possible to establish or correct the following synonymies: Gymnogeophagus cyanopterus is a synonym of Geophagus hal- zanii; Geophagus rhabdotus is not a synonym of G. hrasiliensts; Geophagus camurus is not a synonym of G. gymnogenys but of G. rhabdotus; G. bucephalus, G. lahiatus, G. scymnophilus and G. pygmaeus are not synonyms of G. brasiliensis but of G. gym• nogenys. For every species of each of these three genera the holotype or specimen chosen as lectotype is redescribed and illustrated. Measurements and statistics relating to a series of specimens of each species are given in tables. For species of wide distribution {G. surinamensis, G. jurupari) the material is grouped according to its geographical origin in such a manner as to allow a compa• rison of these populations by various statistical tests to be at• tempted. The significant differences revealed by these tests are, however, regarded as insufficient to merit the erection of sub• species. Keys for identification and the tables given in the appendix are used to separate these three genera and their various species. Consideration of the phylogeny of these three genera based on the condition of the dorsal skeleton, their dentition and their geographical distribution, leads to the conclusion that the genus Biotodoma is the most primitive and that these genera and spe• cies have been derived from Ama2onian ancestors most often in geographical isolation. L INTRODUCTION Cette révision du genre Geophagus est basée sur l'examen des spécimens types et sur l'étude biométrique de très nombreuses séries d'exemplaires appartenant aux grandes institutions zoolo• giques mondiales. Un grand nombre d'espèces ont été décrites ou rapportées à ce genre Sud-américain dont le statut est resté mal défini. Trop souvent, les descriptions d'espèces nouvelles et les révisions n'ont été basées que sur la bibliographie et non sur la comparaison des exemplaires types. Il m'est un agréable devoir de remercier le Roi Léopold qui, au cours de ses missions scientifiques en Amérique latine, aux• quelles j'ai eu l'honneur de participer, a récolté lui-même de très nombreux exemplaires de poissons faisant l'objet de cette étude. Qu'il trouve ici l'expression de ma reconnaissance et de mon attachement. J'exprime ma gratitude toute particulière au Dr. J.W. Atz de l'American Museum of Natural History qui, très absorbé par son important travail de la Dean Bibliography of fishes, a accep• té de me confier le matériel qu'il avait réuni en vue d'une étude de ce genre. Je le remercie ainsi que le Dr. D.E. Rosen pour l'accueil et les facilités qu'ils m'ont accordés. Je remercie cordialement le professeur K. Deckert et le Dr. C. Karrer qui m'ont permis d'examiner dans leur laboratoire les importantes collections du Zoologisches Museum an der Hum• boldt Universitât Berlin. Mes remerciements vont également à toutes les institutions qui m'ont aidé en prêtant les exemplaires de leurs collections: "The Academy of Natural Sciences of Philadelphia, Dr. J.E. Bölke; British Museum (Natural History), Dr. P.H. Greenwood; California Academy of Sciences, San Francisco, Dr. W.N. Esch- meyer; Estacao Experimental de Biologia e Piscicultura Pirassu- nunga, Sao Paulo, Dr.
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