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Freshwater Gammarus Species from Europe, North Africa and Adjacent Regions of Asia (Crustacea-Amphipoda)

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Freshwater Gammarus Species from Europe, North Africa and Adjacent Regions of Asia (Crustacea-Amphipoda) FRESHWATER GAMMARUS SPECIES FROM EUROPE, NORTH AFRICA AND ADJACENT REGIONS OF ASIA (CRUSTACEA-AMPHIPODA) PART II. GAMMARUS ROESELI-GROUP AND RELATED SPECIES by GORDAN S. KARAMAN Biological Institute, Titograd, Yugoslavia & SJOUK PINKSTER Institute of Taxonomic Zoology, University of Amsterdam, The Netherlands FRESHWATER GAMMARUS SPECIES FROM EUROPE, NORTH AFRICA AND ADJACENT REGIONS OF ASIA (CRUSTACEA-AMPHIPODA) PART II. GAMMARUS ROESELI-GROUP AND RELATED SPECIES by GORDAN S. KARAMAN Biological Institute, Tito grad, Yugoslavia 8Z SJOUK PINKSTER Institute of Taxonomic Zoology, University of Amsterdam, The Netherlands ABSTRACT 9) For all species complete lists of all studied localities 1) This part of the revision of the freshwater Gammarus are given (except for G. roeseli, for economical reasons), species deals with the G. roeseli-group. Members of this together with two maps in which the distribution is illus- artificial group are characterized by the presence of dorsal trated. processes (carinae) on the metasome segments. RESUME 2) A historical review is given from the first description of G. roeseli and the taxonomic confusion that followed, to 1) Cette deuxieme partie de la revision traite des Gam- the start of this study when 3 species and at least 15 sub- marus d'eau douce du groupe roeseli. Les membres de ce species were known. groupe artificiel sont caracterises par la presence de carenes 3) Where possible, the type material of all these taxa is dorsales sur les segments du metasome. re-examined and compared with hundreds of newly collected 2) Un apercu historique est donne des la premiere samples and material from all major museums in Europe. description de G. roeseli, et de la confusion taxonomique sui- 4) In this study the same morphological characters are vante. Au debut de cette etude, trois especes et au moms 15 used as in the revision of the G. pulex-group (Karaman sous-especes etaient connues. Pinkster, 1977). The variability of the various characters is 3) Autant que possible, le materiel-type de tous ces taxa a discussed. ete reexamine et compare avec des centaines d'echantillons col- 5) Complete descriptions and many illustrations are given lectionnes recemment et avec du materiel provenant de tous of the males of all species, along with a key for their identifi- les grands musees de l'Europe. cation. Where necessary, figures are given of those details in 4) Dans cette etude les memes caracteres morphologiques which females differ from the males. ont ete utilisees que dans la revision anterieure du groupe 6) It is proved that G. triacanthus and its many subspecies pu/ex (Karaman 8z Pinkster, 1977). La variabilite des divers must be considered synonymous with G. roeseli. G. argaeus caracteres a ete discutee. stojicevici is raised to specific rank. Four new species (G. 5) Des descriptions completes et beaucoup d'illustrations goedmakersae, G. mladeni, G. obnixus and G. pavo) are ont ete donnees des males de toutes les especes, accompagnees described, resulting in a total number of seven species within dune cle de determination. Si necessaire, les caracteres des this group. femelles, lorsqu'ils sont differents, ont ete figures. 7) In many localities members of the G. roeseli-group have 6) Il a ete prouve que G. triacanthus et toutes ses sous- been found together with members of the G. pulex- or G. especes doivent etre consideres comme synonymes de G. balcanicus-group. Experimental work on G. roeseli showed roeseli. Quatre especes nouvelles ( G. goedmakersae, G. mla- that in such cases G. roeseli is more eurybiont than the deni, G. obnixus et G. pavo) ont ete decrites, aboutissant members of the G. pulex- or G. balcanicus-group (higher un nombre total de sept especes dans ce groupe. resistance against low oxygen concentrations and elevated 7) Dans plusieurs localites des populations mixtes des temperatures). Subterranean members of this group have not membres du groupe roeseli et des groupes pu/ex ou bal- been found so far. canicus ont ete trouvees. Des travaux experimentaux ont 8) From zoogeographical data it can be concluded that this montres que G. roeseli est plus eurybionte que les membres group originates from southeastern Europe and Asia Minor. des autres groupes (resistance plus forte aux basses con- Only in Recent times G. roeseli has penetrated inland waters centrations d'oxygene et aux temperatures elevees). Nous in western Europe. Members of this group are lacking in n'avons pas encore observe des membres hypoges de ce groupe. northern Europe, the British Isles, Italy and the Iberian 8) En se basant sur des donnees zoogeographiques, on pout Peninsula. conclure que ce groupe est originaire de l'Europe sud-est et 166 G. S. KARAMAN & S. PINKSTER - GAMMARUS ROESELI-GROUP de l'Asie Mineure. G. roeseli n'a penetre les eaux continen- was given by Rosel von Rosenhof in 1755. Al- tales de l'Europe occidentale que pendant l'Holocene. Les though Rose! gave a very detailed and well-illus- membres de ce groupe sont absents de l'Europe septentrionale, des lies Brittanniques et de la peninsule Iberique. trated description of his Squilla fluviatilis, it was 9) Pour toutes les especes (abstraction faite de G. roeseli in later years often confounded with the species pour des raisons economiques) des listes completes du now known as Gammarus pulex. Although the materiel etudie sont donnees. En plus l'aire de distribution est illustree a l'aide de deux cartes. original description of the latter by Linnaeus, 1758 (as Cancer pulex) was not very detailed, it is INTRODUCTION nevertheless very clear that it was a completely In the first part of the revision of the freshwater different species without the dorsal carinae so taxa belonging to the genus Gammarus, Karaman clearly described by Rosel for Squilla fluviatilis. & Pinkster, 1977, subdivided them into three Nevertheless later authors identified their keeled artificial groups, viz. the G. pulex-group, the G. gammarids as Gammarus pulex until Gervais, balcanicus-group, and the G. roeseli-group. This 1835, made it clear that there existed at least two subdivision was merely an artificial one, based on different species: a keeled one which he called morphological characters only. Gammarus Roeselii and a species without dorsal The members of the Gammarus roeseli-group carinae, called Gammarus pulex. It lasted until differ from those of the other two groups in the 1905 before another keeled, closely related species presence of a middorsal process (carina) on each was described. of the three metasome segments. The setation of When the present study was started only three the pereiopods 3 and 4 as well as of the uropod 3 species were known to belong to this group, viz. can be long (as in the G. pulex-group) or short (as G. roeseli Gervais, 1835, G. triacanthus (Schafer- in the G. balcanicus-group). na, 1922) and G. argaeus Vávra, 1905, but at least The first description of a member of this group 15 subspecies were attributed to them. Most of t • ° • 00 • t • Gammarus pavo • Gammarus goedmakersae • Gammarus argaeus • Gammarus stojicevici Map I. The distribution of four species of the Gammarus roeseli-group in southeastern Europe and Asia Minor. BIJDRAGEN TOT DE DIERKUNDE, 47 (2) - 1977 167 these subspecies were poorly described, often are given on the ecological range of the different defined on basis of one or two characters only, species. which we now know to be quite variable. ACKNOWLEDGEMENTS AND RESPONSABILITIES Therefore it was thought wise to re-examine The authors are indebted to Dr. Sandro Ruffo and Dr. G. the type material of all known taxa. Likewise we Osella of the Museo civico di Storia naturale, Verona, Italy; studied the material present in all major museums to Dr. G. Hartman and Dr. C. Kosswig of the Zoolog. and private collections in Europe, as well as a rich Museum und Institut of the University of Hamburg, G.F.R.; to Dr. L. B. Holthuis of the Rijksmuseum van Natuurlijke material collected by the second author. It became Historie, Leiden, the Netherlands; to Dr. J. Forest of the clear that this group of species has its main distri- Museum national d'Histoire naturelle, Paris, France, for the bution area in southeastern Europe and Asia loan of material studied in this work. Special thanks are due to Dr. Mladen Karaman of the Minor. Six out of seven species are confined to University of Prigtina, Yugoslavia, for his help in obtaining this part of the world and only one species, G. material from some museums; to Mrs. I. Pinkster-de Graaf, roeseli, has a much wider distribution reaching Drs. Annemarie Goedmakers, Drs. J. Dieleman, and Mr. J. van Veen for their assistance to the second author during from the eastern parts of Turkey to the borders of his fieldwork in many parts of the area studied. the Atlantic. In the present paper complete descriptions of Although the present paper is, we hope, a unity, all members of the G. roeseli-group and a key for each of the authors took special care for certain their identification are given. In doing so, we used sections, which were discussed afterwards with the same taxonomic characters and terminology as his co-author. Therefore, both authors share the we used previously in the revision of the G. pulex- full responsability for the total work. group (Karaman & Pinkster, 1977). The distribu- ABBREVIATIONS USED tion of the various species is discussed and Z.M.A. = Zoologisch Museum Amsterdam, the Nether- illustrated in two maps, and where possible notes lands. In Gam marus mladeni • Gammarus obnixus Map II. The distribution of two species of the Gammarus roeseli-group in Asia Minor. 168 G. S. KARAMAN & S. PINKSTER — GAMMARUS ROESELI—GROUP K.C. = Collection of S.
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