Systematic, Phylogenetic and Biological Considerations on The
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'' BULLETIN DE L' INSTITUT ROYAL DES SCIENCES NATURELLES DE BELGIQUE, BlOLOGlE, 76: 33-110,2006 BULLETIN VAN HET KONINKLIJK BELGISCH lNSTITUUT YOOR NAT UURWETENSCHAPPEN, BlOLOGIE, 76: 33-11 0, 2006 Systematic, phylogenetic and biological considerations on the genera Bathyporeia, Amphiporeia, Pontoporeia and Priscillina, with redes~ription of the West-Atlantic Bathyporeia species and description of a new Priscillina from Svalbard (Crustacea, Amphipoda) by Cedric d'UDEKEM d' ACOZ Abstract Resume The two West-Atlantic species of Bathyporeia, B. parkeri and B. Les deux especes ouest-atlantiques de Bathyporeia, B. parkeri et B. quoddyensis are re-described. They exhibit a large number of differ quoddyensis sont redecrites. Elles presentent un grand nombre de ences with East-Atlantic and Mediterranean Bathyporeia species. differences avec les especes est-atlantiques et mediterraneennes de Cladistic analyses of morphological characters of all known Bathyporeia. Des analyses cladistiques des caracteres morphologi Bathyporeia species indicate that the two geographic groups are sis ques de toutes les especes connues de Bathyporeia indiquent que les ter clades. There is strong evidence that in the genus Bathyporeia deux groupes geographiques sont des clades sceurs. II y a de serieu the character state 'short antennae in mature males' is the ses evidences qu'au sein du genre Bathyporeia, l'etat de caractere plesiomorphic condition and that ' long antennae in mature males' is 'antennes courtes chez le male mature' soil plesiomorphique. Des the apomorphic condition. Representatives of the genera representants des genres Amphiporeia, Pontoporeia et Priscillina Amphiporeia, Pontoporeia and Priscillina are characterised and il sont caracterises et illustres, en tant que point de comparaison avec lustrated, as a basis of comparison with Bathyporeia. These three Bathyporeia. Ces trois genres sont traditionnellement consideres genera are traditionally considered to be the closest relatives of comme les plus proches parents de Bathyporeia et sont aJors regrou Bathyporeia and grouped together in the family Pontoporeiidae. pes dans Ia famille des Pontoporeiidae. La position systematique de The systematic position of these four alleged ' pontoporeiid' genera ces quatre genres supposes appartenir aux Pontoporeiidae est is investigated in a cladistic analysis includi ng several other investiguee au moyen d' une analyse cladistique incluant plusieurs amphipods which are either morphologically similar or potentially autres amphipodes, lesquels sont soil morphologiquement similai plesiomorphic. In this analysis the monophyly of the genus res, soit potentiellement plesiomorphiques. Dans cette analyse, Ia Bathyporeia is supported by a very high number of monophylie du genre Bathyporeia est etayee par un nombre tres synapomorphies. Bathyporeia itself forms a strongly supported eleve de synapomorphies. Bathyporeia lui-meme forme un clade clade (the Bathyporeiidae) with the genus Amphiporeia. There is no tres robuste (les Bathyporeiidae) avec le genre Amphiporeia. II n' y a cladistic support for a sister relationship between Bathyporeia + pas Ia moindre evidence cladistique que le clade Bathyporeia + Amphiporeia and Pontoporeia + Priscillina. The Bathyporeiidae Amphiporeia soit le taxon sceur. de Pontoporeia + Priscillina. Les seem to have stronger affinities with the Gammaridae sensu stricto Bathyporeiidae semblent avoir davantage d'affinites avec les and especially with the Pontogammaridae. The Bathyporeiidae are Gammaridae sensu stricto et surtout avec les Pontogammaridae. therefore considered as belonging to the superfamily Gammaroidea. Les Bathyporeiidae sont done consideres comme appartenant a Ia Cladistic analyses of morphological characters suggest loose rela superfamille des Gammaroidea. L'analyse cladistique de caracteres tionships between the genera Haustorius, Pontoporeia and morphologiques suggere une parente possible entre les genres Priscillina. However these three genera are separated by a consider Haustorius, Pon.toporeia et Priscillina. Cependant, les evidences able morphological gap. The morphological isolation of the genus sont fa ibles et peu convaincantes, et qui pl us est, ces trois genres Priscillina justifies the creation of a monotypic fa mily Priscillinidae sont separes par des differences morphologiques profondes. L' isole fam. nov. for it. T he superfamily Pontoporeioidea is retained for the ment morphologique du genre Priscillina justifie Ia creation de Ia Pontoporeiidae and Priscill inidae fam. nov. The materiaJ of famille monotypique des Priscillinidae fam. nov. La superfamille Priscillina used in this study includes a new species from Svalbard, des Pontoporeioidea est retenue pour les Pontoporeiidae et les which is described as Priscillina herrmanni sp. nov. Priscillinidae fam. nov. Le materiel utilise pour cette etude inclut une nouvelle espece du Svalbard qui est decrite sous le nom de Key-words: Bathyporeia, Amphiporeia, Pontoporeia, Priscillina, Priscillina herrmanni sp. nov. Bathyporeiidae, Gammaridae, Melitidae, Melphidippidae, Pontogammaridae, Pontoporeiidae, Priscill inidae fam. nov. , Mots-cles: Bathyporeia, Amphiporeia, Pontoporeia, Priscillina, Amphipoda, Crustacea, taxonomy, phylogeny, cladistic, North At Bathyporeiidae, Gammaridae, Meli tidae, Melphidippidae, lantic. Pontogammaridae, Pontoporeiidae, Priscillinidae fam. nov., Amphipoda, Crustacea, taxonomie, phylogenie, cladistique, Atlan tique Nord. 11 34 CEDRIC d' UDEKEM d'ACOZ 1. Introduction ratios. The values given should be considered as indicative only, since ratios have been measured on one or two speci The taxonomy and morphology of the genus Bathyporeia mens only. After the examination of all known species of LfNDSTROM, 1855 have been the object of several recent pa Bathyporeia, some ratios used in previous papers have pers: D'UDEKEM D' ACOZ & BERGE (2003), D' UDEKEM proved to be redundant or to have low information content. D' ACOZ (2004), D'UDEKEM D' ACOZ & MENIOUI (2004), Such ratios have been abandoned in the present paper. The D' UDEKEM D' ACOZ & VADER (2004, 2005a, 2005b) and term ' pseudorostrum' denotes tne enlarged first article of the D'UDEKEM D' Acoz et al., (2006). The present paper, which peduncle of :he first antenna. The terminal crown of setae of is the last part of this study, consists of two sections. In the the third article of the maxillipedal palp is not illustrated. The first section, the two West-Atlantic Bathyporeia species are 'carpal fang' is a long and strong modified seta found on the re-described, illustrated, and compared with the Bathyporeia carpus of the third and fourth pereiopods in all Bathyporeia species from the other side of the Atlantic. Cladistic analyses species. In Bathyporeia, the medial border of the outer ramus are run on all known species of Bathyporeia. They show that of the third uropod normally has long plumose setae orien the West-Atlantic species are the sister clade of the East-At tated in the same plane as the outer ramus; each plumose seta lantic I Mediterranean species and that evolutionary direction is either single or forms a group with one (sometimes two or apparently goes towards an elongation of the second anten three) non-setulose spiniform setae pointing obliquely up nae in males. wards; these spiniform setae, which are not present in all spe The second section of this paper is focused on the systematic cies, are called 'accessory spiniform setae' . position of the genus Bathyporeia and its alleged and puta The following abbreviations of morphological features are tive relatives. In literature the genus Bathyporeia adopted in the present paper, either systematically or in some LINDSTROM , 1855 is traditionally assigned to the family parts of the text: A1: first antenna; A2: second antenna; Pontoporeiidae DANA, 1853, along with Amphiporeia SHOE Coxae 1-7: coxal plates of the first to seventh pereiopods; MAKER , 1929, with Pontoporeia KR0YER, 1842 and its satel Ep1-Ep3: first to third epimeral plates; Md: mandible; Mx1: lites Diporeia BOUSFIELD, 1989 and Monoporeia first maxilla; Mx2: second maxilla; Mxp: maxilliped; P 1: BOUSFIELD, 1989, and with Priscillina STEBBfNG, 1888. first pereiopod (first gnathopod); P2: second pereiopod (sec However, these genera are grouped together more by tradi ond gnathopod); P3-P7: third to seventh pereiopods; U1 -U3: tion than on the basis of serious phylogenetic evidence. The first to third uropods. morphology and biology of these genera are briefly outlined The following abbreviations of Scientific Institutions are and compared, and at least one representative of every group used: ARC: Atlantic Reference Centre, Huntsman Marine is illustrated in detail. A cladistic analysis is run with 4 Sciences Centre, St Andrews, New Brunswick, Canada; Bathyporeia species, one Amphiporeia, one Pontoporeia, A WI: Stiftung Alfred-Wegener-Institut fi.ir Polar- und one Priscillina and 21 other species of amphipods. From tllis Meeresforschung in der Helmoltz-Gemeinschaft, analysis, it appears that the genus Bathyporeia LINDSTROM, Bremerhaven, Germany; CMNC: Canadian Museum of Na 1855 is the sister taxon of the genus Amphiporeia SHOE ture, Ottawa, Canada; NOAA: NOAA National Benthic In MAKER, 1929. On the other hand, there is no evidence that ventory, NOS Center for Coastal Environmental Health and Pontoporeia and Priscillina are very closely related to the Biomolecular Research, Charleston, South Carolina, USA; Amphiporeia I Bathyporeia lineage