The Freshwater Crabs of West Africa : Family

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The Freshwater Crabs of West Africa : Family F' CI ,u ~n 'e e' t: ·.f 1 c). r e t; r 0' IP 'i, cCii ,e ·s N il Cumberli ge The Freshwater Crabs of West Africa Family Potamonautidae For Louise Neil CUMBERLIDGE The Freshwater Crabs of West Africa Family Potamonautidae Éditions de l'IRD -\ ~ ,- Institut de Recherche pour le Développeme Collection Faune et Flore Tropicale: ~o r ' Paris, 1999 Préparation de copie Pierre Rondeau Maquette intérieure et mise en page IODE Informatique Maquette de couverture Michelle Saint-Léger © IRD éditions, 1999 ISBN 2-7099-1433-6 ISSN 1286-4994 Contents Préface 11 Summary 15 Résumé '" 23 Introduction 33 History of freshwater erab studies in West Mriea 34 The taxonomie structure of the West Mriean freshwater erabs today .44 The Gecarcinucidae .45 Madagascan freshwater crabs .. .46 Conclusions .48 List of speeies 50 Material examined 51 Colleeting loealities 52 Repositories and abbreviations 53 Aeknowledgements 54 Cladistic analysis of the West African freshwater crabs 63 Freshwater erab morphology and explanation of terminology 63 Body plan 63 Cephalothorax 64 Head 69 Thorax 71 Sternum 72 Pereiopods ............... 75 Abdomen 77 Gonopod terminology 80 The systematie position of the West Mriean freshwater erabs 99 Characters examined 101 Gonopods 101 Sternum 105 Mouthparts 105 Carapace 107 Pereiopods 110 Abdomen 1 10 Data analysis 111 Summary of findings 118 Systematic study 125 Family Potamonautidae 125 Potamonautes 129 Liberonautes 143 Sudanonautes 172 Potamonemus 208 Family Uncertain 214 Subfamily Globonautinae 214 Globonautes 216 Afrithelphusa 218 Louisea 226 Morphometric relationships 275 Biogeography and Biology 305 Distribution of the species 305 Climate and vegetation 306 Patterns of distribution, species diversity. and endemism 310 Interpretation of distribution patterns 314 Biology 316 Respiratory system 316 Reproductive system 318 Literature 341 Gazeteer 357 Index 365 List of figures, tables and plates 377 Préface Plus particulièrement spécialiste de crabes marins, j'ai grand plaisir à préfacer un ouvrage consacré aux crabes d'eau douce, qui sont intèressants à plus d'un titre, Les crabes d'eau douce (en anglais "true freshwater crabs") se sont complètement affranchis du berceau originel, la mer, et peuplent les régions tropicales et les zones chaudes-tempérées à tempérées des cinq continents, Ils n'ont pas de parents reconnaissables dans les océans. Ils passent toute leur existence en dehors du milieu marin. Ils habitent à l'intérieur des rivières et des cours d'eau, ou près des berges, dans les lacs, les étangs, les mares, parfois au cœur des montagnes jusqu'à 3 000 m d'altitude dans les tor­ rents, ou ils s'aventurent à terre dans la forêt humide pour séjourner sous les feuilles ou parmi les racines, généralement dans une atmosphère humide, ou bien encore ils pénètrent au fond des grottes, devenant de vrais troglobies profondément modifiés. Mais certaines espêces peuvent non seulement survi­ vre en dehors de l'eau mais aussi tolérer des conditions relati­ vement sèches. Les crabes d'eau douce sont donc bien différents des diverses catégories de crabes marins qui ont pu coloniser les eaux saumàtres puis continentales, et même conquérir des habitats terrestres. Contrairement à nos crabes d'eau douce, ces représentants de familles marines qui, d'abord amphibies, se sont complètement adaptés à un nouvel environnement pour devenir crabes des rochers, crabes de sable, crabes violonistes, crabes des arbres, crabes de terre, montrent des liens de paren­ té avec d'autres familles vivant dans la mer. Et ils y retournent pour leur vie larvaire, parfois au prix de longues migrations, En revanche, chez les vrais crabes d'eau douce, le développe­ ment est direct et se passe complètement en dehors de la mer. Après une longue période d'incubation, d'un trés gros œuf sort un petit crabe directement éclos qui a déjà la forme des parents: c'est une miniature de l'adulte. Pendant un certain temps les jeunes vivent sous l'abdomen de la femelle, se pro­ menant sur sa carapace ou alentour avec une protection parentale, comme on le voit chez des araignées ou des cra­ pauds, Contrairement à ce qui se passe chez les crabes marins, il n'y a pas de stade planctonique, la zoé, et donc pas de dissémination puisque les phases nageuses ont été élimi­ nées, C'est pourquoi un isolement géographique souvent très Important caractérise les formes d'eau douce, La reconnais- sance des espèces, souvent extrêmement difficile, doit faire appel à la morphologie des appendices sexuels mâles. Le mèmoire du Dr. Neil CUMBERLIDGE est une synthèse des connaissances sur les crabes d'eau douce de l'Ouest Africain et il couvre la faune de dix-huit pays de ce continent. Une seule famille, cene des Potamonautidae, est représentée, mais avec sept genres et 32 espèces, soit le tiers des espèces et sous-espè­ ces de crabes dulçaquicoles actuellement recensées sur le conti­ nent africain. Cinq nouvelles espèces sont décrites et, au passa­ ge, est distinguée une nouvelle famille, dont les reprèsentants habitent le lac Tanganyika. Le présent mémoire vient jalonner l'histoire des crabes d'eau douce africains, marquée par l'œu• vre de Mary J. RATHBUN (1904-1906), en partie réalisée au Muséum national d'histoire naturelle, Paris, et de Richard Bott dont la monographie de 1955 traitait toute la faune bra­ chyourienne de l'Afrique. Deux articles de Théodore MONOD, 1977 et 1980, avaient plus particulièrement concerné la faune sahélo-soudanaise. N. CUMBERLIDGE couronne ses recher­ ches passées sur la faune africaine, ponctuées de nombreuses notes, par ce travail qui rassemble de très nombreuses données. Il s'est appuyé sur ses récoltes personnelles ainsi que sur le matériel déposé dans les grands musées, et notamment au Muséum national d'histoire naturelle, Paris. Pour chaque espè­ ce, figurent une liste des synonymies et des références, une dia­ gnose, des renseignements biogéographiques et écologiques. En outre, est dressée une histoire phylogénétique de la famille des Potamonautidae, où la question cruciale de la monophylie est discutée. Pour cela, nous est proposée la première analyse cla­ distique des crabes d'Afrique, qui se base sur 33 caractères morphologiques. Le clade Potamonautidae est en effet reconnu, avec deux branches principales qui sont interprétées comme deux sous-familles. Une très intéressante étude morphomètrique concernant les proportions du corps fait ressortir qu'une carapace plate est associée à une existence complètement aquatique, tandis qu'u­ ne carapace plus épaisse est mise en rapport avec une existen­ ce semi-terrestre ou terrestre. Le mémoire du Dr. Neil CUMBERLIDGE sur les crabes d'eau douce de l'Ouest Africain fait suite à deux ouvrages consacrés à deux familles de crabes des eaux douces d'Amérique (tous deux par G. RODRlGUEZ, 1982 et 1992), pareillement publiés dans la série « Faune tropicale» de l'Orstom. L'Institut fran­ çais de recherches pour le développement en coopération s'honore de publier des travaux d'ordre fondamental et. plus particulièrement, de systématique. Le savoir zoologique, avec la connaissance précise de la taxonomie des animaux, est à la 12 - Cumberlidge: Freshwater Crabs base de toutes les autres sciences et peut seul permettre des recherches d'ordre biologique et appliqué. Les crabes d'eau douce ont une grande importance économique. Ils forment une part non négligeable de l'alimentation dans cer­ tains pays. On sait qu'lIs peuvent abriter le dernier stade larvai­ re (métacercaire) d'un ver trématode à l'origine de la paragoni­ mose, ou distomatose pulmonaire. une maladie qui sévit grave­ ment en Asie du Nord-Est et du Sud-Est, en Afrique et en Amérique latine. Le premier hôte est un mollusque {dont l'iden­ tité n'est pas connue en ce qui concerne l'Afrique!. le crabe est le deuxième hôte, et le dernier un mammifère carnivore ou un primate. L'homme s'infeste en mangeant des crabes insuffisam­ ment cuits: la métacercaire résiste à un quart d'heure d'ébulli­ tion. Le Bulletin de l'OMS de 1995 faisait état, pour le mondé entier, de 30 millions de patients et, par ailleurs, estimait à 190 millions d'individus la population exposée du fait de l'ingestion de crabes. En Afrique, les pays les plus touchés actuellement seraient le Cameroun et le Nigeria. Seules deux espèces du tréma-tode Paragonimus sont connues en Afrique mais leur nom­ bre est probablement plus élevé. Au moins sept espèces ouest­ africaines de crabes d'eau douce ont été identifiées comme les hôtes intermédiaires de ce parasite pathogène: Liberonautes lati­ dactylus, L. paludicolis, L. chaperi, Sudanonautes africanus, S. aubryi, S. orthostylis, S. granulatus, mais 11 est probable que d'autres espèces, récoltées dans les régions humides ou dans la savane de l'ouest africain et utilisées comme nourriture, sont dangereuses. Comment reconnaître les espèces en question, comment en dis­ tinguer les formes les plus proches? Dans quelles régions sont­ elles présentes et dans quel habitat? Quelles sont les autres for­ mes potentiellement dangereuses? Nous trouverons toutes ces informations dans le mémoire qui est publié ici. Diverses clefs dichotomiques pour permettre la reconnaissance à tous les niveaux, une excellente 1llustration, de nombreuses cartes de répartition, une bibliographie complète, ainsi qu'un index facili­ teront l'utilisation de cet ouvrage. Nous savons gré à l'auteur de nous offrir une œuvre de cette qualité, qui sera utile non seulement au taxonomiste, au bio­ géographe, à l'écologiste, mais aussi à tous ceux qui sont en charge de la gestion et de la protection de la diversité du vivant, à ceux qui étudient les problèmes de l'alimentation dans le monde, au médecin. au parasitologiste. Danièle GUINOT Professeur au Muséum national d'Histoire naturelle Préface - 13 Summary The freshwater crabs of West Africa The area of West Africa covered by this book includes eighteen countries: Mauritania, Senegal, The Gambia, Guinea Bissau, Guinea, Sierra Leone, Liberia, Côte d'Ivoire.
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