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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 I. GAMMARUS PULEX-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 (CRIJSTACEA-AMPHIPODA) PART I. GAMMARUS PULEX-GROUP AND RELATED SPECIES by GORDAN S. KARAMAN Biological Institute, Titograd, Yugoslavia SJOUK PINKSTER Institute of Taxonomic Zoology, University of Amsterdam, The Netherlands ABSTRACT 10) Within some taxa (e.g. G. fossarum and G. p. pulex) morphologically aberrant populations can be found. Since these 1) Within the genus Gammarus s.str., which is widely distrib- populations do successfully cross-breed and are sympatric they uted throughout the northern hemisphere, more than 100 must be considered mere variations. freshwater species can be distinguished. 11) We had serious problems to determine the identity 2) These freshwater species can be classified in three of several Gammarus species, especially from Asia Minor, artificial groups: (a) the G. pulex-group (species without because of the impossibility to obtain literature and type dorsal carina and with dense setation on pereiopods 3 and 4 material of some Russian authors. (So we cannot exclude the and uropod 3), (b) the G. balcanicus-group (species without possibility that our species described from Asia might be dorsal carina and poorly setose pereiopods 3 and 4 and identical with a species formerly described by a Russian uropod 3) and (c) the G. roeseli-group (species with dorsal author). carina). These groups are merely artificial ones since tran- 12) For all species, except the most common ones, com- sitive (intermediate) species do exist. Moreover, the origin plete lists of all localities studied are given. Moreover, in 3 of the species is not known, so that polyphyly is not excluded. maps the distribution of the various species and subspecies 3) In the present work the Gammarus pulex-group is is illustrated. revised, based on rich material from Europe, North Africa, 13) It was not possible to illustrate all morphological Asia minor and adjacent areas. details of every taxon mentioned in the present work. Only 4) The taxonomic differences between the various species G. pulex, the type species of the genus Gammarus and the are usually small but distinct and stable. This is especially nominal form of the entire group, is figured completely. evident in mixed populations of two or more species. In those For the other taxa, only those parts are illustrated that are populations no intermediates between the taxa have been fundamentally different from those of G. p. pulex. observed. 5) In some cases, morphological differences between two RSUME species are hardly discernible although reproductive isolation is present (e.g. G. fossarum and G. wautieri). 1) Au sein du genre Gammarus s. str. qui est largement 6) Hybridization experiments can solve taxonomic prob- repandue dans l'hemisphere nord, l'on peut distinguer plus lems and test the taxonomic value of morphological dif- que cent especes d'eau douce. ferences between populaticrns. Such experiments between many 2) On peut subdiviser ces especes d'eau douce en trois Asiatic and European populations might clarify their groupes artificiels: a) le groupe G. pulex (especes sans taxonomic status. carenes dorsales avec des pereiopodes 3 et 4 et uropode 3 7) The taxonomic characters may largely be variable within tres setoses), b) le groupe G. balcanicus (especes sans carenes one population as well as in different populations of the same dorsales et avec les pereiopodes 3 et 4 et uropodes 3 peu species. setoses) et c) le groupe G. roeseli (especes avec carenes 8) Characters that are very stable in one species can be dorsales). Ce ne sont que des groupes artificiels, puisque largely variable in other species (e.g. presence of calceoli, des especes transitives (intermediaires) sont connues, comme length of rami of uropod 3). Nevertheless we can distinguish nest pas connu l'origine des especes. certain stable characters (e.g. the structure of the mandibular 3) Dans l'etude presente une revision est donnee du palp), but also instable ones (e.g. the number of dorsal and groupe pulex basee sur un materiel riche originaire de lateral spines on the urosomites) in all species. l'Europe, de l'Afrique du Nord, l'Asie Mineure et quelques 9) Gammarus pulex has given rise to several isolated regions adjacentes. populations, adapted to subterranean life (being blind or 4) Les differences taxonomiques entre les differentes having enlarged eyes). These populations are considered especes sont legeres, habituellement, mais distinctes et stables. distinct subspecies. Ceci est surtout evident dans des populations mixtes corn- 2 G. S. KARAMAN & S. PINKSTER - GAMMARUS PULEX-GROUP portant deux ou plusieurs especes. Dans ces populations I on period, numerous different species, many of them n'a jamais observe des specimens intermediaires entre deux now considered representatives of other genera especes. 5) Dans certains cas, les differences morphologiques entre were included in the genus Gammarus. deux especes sont difficiles a reconnaitre, quoiqu'il existe In the second half of the 19th and in the 20th une barriere reproductrice entre ces especes (par exemple century, many species at first attributed to the G. fossarum et G. wautieri). 6) Les experiences d'hybridation sont une bonne methode genus Gammarus were transferred to new genera. pour resoudre des problemes taxonomiques, et pour verifier Nevertheless, up to this moment, the genus Gam- la valeur taxonomique de differences morphologiques qui marus is not yet satisfactorily delimited, while the existent entre des populations. II serait recommandable d'ef- fectuer des experiences d'hybridation entre un grand nombre taxonomy of the various species within this genus, de populations asiatiques et des populations europeennes af in especially of the freshwater forms, is not yet de verifier leur valeur taxonomique. settled. 7) Les caracteres taxonomiques peuvent varier de facon considerable au sein dune meme population, ainsi que dune Because of the extreme variability at the one population a l'autre. hand and the occurrence of convergent characters 8) Des caracteres qui sont tres stables dans une espece at the other hand, during a long time the opinion peuvent etre tres variables dans une autre (par ex. la pre- sence de calceoles, la longueur des rames de l'uropode 3). prevailed that only a limited number of very Neanmoins, nous pouvons distinguer des caracteres stables variable Gammarus species occurred in Europe and (par ex. les palpes mandibulaires) et des caracteres instables the adjacent parts of Africa and Asia. (par ex. le nombre d'epines de l'armature dorsale). 9) Gammarus pulex a cree plusieurs populations isolees, Hybridization experiments as well as detailed adaptees a la vie sous-terraine (aveugles ou a yeux agrandis). studies on variability, distribution and ecology of Lan a considere ces populations comme des sous-especes the various morphologically different populations distinctes. 10) Au sein de certaines unites taxonomiques (par ex. showed, that a very considerable number of inter- G. fossarum et G. p. index) I on peut trouver des popu- sterile Gammarus species inhabits the inland waters lations morphologiquement distinctes. Puisque ces popula- in the area studied. tions sinter-fecondent avec succes, et un mecanisme d'isolation geographique n'existe pas, i on doit considerer ces popu- During the second colloquium on the genera lations comme des simples variations. Niphargus and Gammarus in Lyon, 1973 it ap- 11) Pendant le present travail nous nous sommes heurtes peared that the present authors both tried to solve a de serieux problemes de determination de beaucoup d'especes du genre Gammarus, en particulier ceux de l'Asie the complicated taxonomic position of the various Mineure, a cause de l'impossibilite d'obtenir la litterature et freshwater species within the genus Gammarus. le materiel typique de certains auteurs russes. (Par consequent In order to make the situation not more complex nous ne pouvons pas exclure la possibilite que certaines de nos especes asiatiques sont identiques avec des especes decrites than it was already, both authors decided that it precedemment par des auteurs russes). would be wise to join forces. In this light they 12) Pour toutes les especes, sauf pour quelques especes decided to study the genus Gammarus in Europe, tres communes (par ex. G. pulex, G. fossarum et G. lacus- iris) pour des raisons de limitation d'espace, des listes North Africa and the adjacent countries of Asia completes du materiel etudie sont donnees. En plus l'aire de using identical taxonomic criteria for all taxa distribution connue est decrit et illustre a l'aide de trois involved. The study was based on the material cartes. 13) II n'etait pas possible de faire des figures de tous les from all the major museums and private collections details de chaque unite taxonomique mentionne dans ce from Europe, and a rich material collected by the travail. Seulement G. pulex, l'espece typique du genre Gam- second author. marus et la forme nominale de tout le groupe presente ici a ete dessine entierement. Pour tous les autres (sous)especes, Because of the large number of limnic Gam- seulement les parties qui different principiellement de celles marus species living in the area studied, the genus illustrees pour G. p. pulex sont representees. was subdivided into three artificial species-groups. This paper is limited to members of the Gammarus INTRODUCTION pulex-group only. In following papers the mem- Among the various amphipod genera, Gammarus bers of the other groups will be discussed. is probably the genus with the highest number of During this study the authors had to cope with epigean freshwater taxa. The first members of many difficulties, the most important being the this genus already were described during the Lin- often very incomplete existing descriptions and nean period, about 200 years ago.
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