Zoologische Verhandelingen Leiden 126: 1-242
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The genus Bathyporeia Lindström, 1855, in western Europe (Crustacea: Amphipoda: Pontoporeiidae) C. d’Udekem d’Acoz Udekem d’Acoz, C. d’. The genus Bathyporeia Lindström, 1855, in western Europe (Crustacea: Amphipoda: Pontoporeiidae). Zool. Verh. Leiden 348, 28.v.2004: 3-162, figs. 1-76.— ISSN 0024-1652, ISBN 90-73239-90-7. C. d’Udekem d’Acoz, Tromsø Museum (Department of Zoology), University of Tromsø, 9037 Tromsø, Norway (e-mail: [email protected]). Key words: Crustacea; Amphipoda; Bathyporeia; Atlantic, Europe; taxonomy; ecology. The amphipod genus Bathyporeia Lindström, 1855, on the European Atlantic coasts is revised and new identification keys are provided. The importance of new characters such as the morphology of the first uropod is stressed. The following species are fully redescribed and illustrated: B. elegans Watkin, 1938, B. gracilis G.O. Sars, 1891, B. guilliamsoniana (Bate, 1857), B. nana Toulmond, 1966, B. pelagica (Bate, 1857), B. pilosa Lindström, 1855, B. sarsi Watkin, 1938, and B. tenuipes Meinert, 1877. B. elegans exhibits unusual variation, of which the taxonomical significance is still unclear. It has been observed for the first time that all species of the genus exhibit an unusual sexual dimorphism on the mandibular palp: in the adult males, the ventral border of the terminal article has a comb of stiff setae which is lacking in females and immature males. Furthermore, a locking system holding together the inner plates of the maxilliped is described and illustrated. A previously overlooked vestigial dactylus on the second and the fifth pereiopod is described. Contents 1. Introduction ............................................................................................................................................... 4 2. Historical review ...................................................................................................................................... 4 3. Ecology and biology ............................................................................................................................... 9 4. Material and methods ......................................................................................................................... 12 4.1. Dissection, microscopical preparations, illustrations and descriptions ............. 12 4.2. Terminology and abbreviations ............................................................................................ 14 4.3. Practical recommendations for preservation and identification ............................ 16 5. Systematic account ............................................................................................................................... 19 5.1. Genus Bathyporeia Lindström, 1855 ...................................................................................... 19 5.2. Diagnostic importance of characters ................................................................................... 23 5.3. Keys to species ............................................................................................................................... 28 5.3.1. Simplified key ..................................................................................................................... 28 5.3.2. Diagnostic key .................................................................................................................... 29 5.4. Description of species ................................................................................................................. 32 5.4.1. Bathyporeia elegans Watkin, 1938 ................................................................................. 32 5.4.2. Bathyporeia gracilis G.O. Sars, 1891 ............................................................................ 42 5.4.3. Bathyporeia guilliamsoniana (Bate, 1857) ................................................................... 46 5.4.4. Bathyporeia nana Toulmond, 1966 .............................................................................. 52 5.4.5. Bathyporeia pelagica (Bate, 1856) .................................................................................. 56 5.4.6. Bathyporeia pilosa Lindström, 1855 ............................................................................. 61 5.4.7. Bathyporeia sarsi Watkin, 1938 ...................................................................................... 68 4 d’Udekem d’Acoz. The genus Bathyporeia Lindström in W. Europe. Zool. Verh. Leiden 348 (2004) 5.4.8. Bathyporeia tenuipes Meinert, 1877 .............................................................................. 75 6. Acknowledgements .............................................................................................................................. 79 7. References .................................................................................................................................................. 80 8. Taxonomic index .................................................................................................................................... 89 1. Introduction In his well-known monograph, Lincoln (1979) expressed the feeling that in the North East Atlantic area, there was scarcely a single amphipod species that was ade- quately described and defined at that time. Today, his statement remains largely true, although the difficulties with identification vary considerably from group to group. In this context, the genus Bathyporeia Lindström, 1855, of which a typical representative is illustrated on fig. 1, is certainly one of the most problematic taxa. This genus comprises many species, very closely related to each other, that exhibit considerable sexual dimorphism, allometric variation, and sometimes significant intraspecific varia- tion. B. nana Toulmond, 1966 is the only species, that has been fully described and illustrated. Other species are known incompletely, by short descriptions and by semi- diagrammatic illustrations of some parts only. This situation is particularly deplorable since many species are common or even abundant, and therefore often present in the samples of ecologists working on the benthos of soft bottoms. Several benthos specialists, such as Junoy & Viéitez (1990), Cattrijse et al. (1993) and Ysebaert et al. (1998) report serious difficulties in identification. Misidentifications are probably not uncommon, as is the case for the material of Dörjes & Reineck (1977), that has been re- examined during the present study. In the present paper, all species found in western Europe, from the North Cape to southern Spain are redescribed. This is the first paper in a world-scale revision of the genus Bathyporeia. Adult specimens are fully illustrated for the first time and new identification keys are provided. It is hoped that this work will facilitate their identifi- cation considerably, although Bathyporeia will undoubtedly remain a difficult genus. Unfortunately, some questions remain open concerning B. elegans Watkin, 1938, which exhibits considerable geographic variation. 2. Historical review The history of the genus Bathyporeia is rather complicated, especially with regard to the period before Watkin’s (1938) revision. For this reason, a detailed historical account is given here, the first part of it being largely a re-transcription of Watkin’s excellent review. The genus Bathyporeia was created by Lindström (1855) with the description of the type species of the genus: Bathyporeia pilosa Lindström, 1855, from specimens from the Baltic Sea. Bate (1856; 1857a) described two further species, for which he erected a new genus, Thersites Bate, 1856: Thersites pelagica Bate, 1856 and Thersites Guilliamsoniana Bate, 1857. Later, Bate (1857b) discovered he had overlooked Lindström’s (1855) paper. He sub- sequently adopted the genus proposed by the Swedish naturalist (misspelling it as Bathyporea) but erroneously considered his species Guilliamsoniana (spelled this time d’Udekem d’Acoz. The genus Bathyporeia Lindström in W. Europe. Zool. Verh. Leiden 348 (2004) 5 Fig. 1. Bathyporeia elegans Watkins, 1938 forma A, female, Grindøen, 23.viii.1884, TMU 1395. Complete animal. Composite drawing; mostly after an exuvium, except eye and third uropod: after another female mounted on the same slide. 6 d’Udekem d’Acoz. The genus Bathyporeia Lindström in W. Europe. Zool. Verh. Leiden 348 (2004) as Guilliamsonia) a junior synonym of B. pilosa. Bate (1862) gave more detailed descrip- tions of his ‘B. pilosa’ and B. pelagica, and described a further species, B. Robertsonii. His illustrations however are crude, inaccurate and very difficult to interpret. It should be noted that Thersites Bate, 1857 is a junior homonym of a mollusc genus. Bate and West- wood (1863) treated the three species B. pelagica, ‘B. pilosa’ and B. Robertsonii again in their monograph of British sessile-eyed crustaceans. Marcusen (1867: 359) described a new species from the Black Sea, B. pontica, in a minimalist diagnosis: “ähnlich der B. Robertsonii Sp. Bate, mit calceola am unteren Fühler, aber auch einer calceola am oberen.” Stebbing (1875) placed all West European species in synonymy, only recognizing the nominal species B. pilosa as a valid species. Meinert (1877) accepted Stebbing’s point of view but described one further species: B. tenuipes Meinert, 1877. Blanc (1884) too accepted Stebbing’s point of view. Later, Chevreux (1887) resurrected B. Robertsoni but suspected that B. pilosa and B. pelagica would indeed be identical. Robertson (1888) considered B. Robertsoni a junior