(Crustacea, Springs ( B. Group, Ingolfiella
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Bijdragen tot de Dierkunde, 51 (2): 345-374 — 1981 Amsterdam Expeditions to the West IndianIslands, Report 14. The taxonomy and zoogeography of the family Bogidiellidae (Crustacea, Amphipoda), with emphasis onthe West Indian taxa by Jan H. Stock Institute of Taxonomie Zoology, University of Amsterdam, P.O. Box 20125, 1000 HC Amsterdam, The Netherlands Abstract In the four additional are present paper, species named (two from Tortola, and one each from of of the The diagnosis a family groundwater Amphipoda, and and several unnamed Based cladistic the Saint John Margarita), Bogidiellidae, is revised. on a analysis, former is subdivided. In its con- forms recorded Puerto Rico and Cu- genus Bogidiella present are (from ception, the Bogidiellidae comprise eleven named genera, raçao ). and 50 named whereas several other seven subgenera, species, all have all Several authors, above Ruffo, taxa remain unnamed. These are distributed over major 1973, continents (except Antarctica), and some oceanic islands. This attempted in the past to find some order in the distribution pattern is presumably due to at least two major seemingly randomly distributed morphological of the Mesozoic vicariant processes: the breakup Pangaea in features of the various members of what is and the geological regression movements in the Tertiary. usually A number of West Indian taxa is described, including four considered The worldwide one genus, Bogidiella. new species. occurrence of the presumed Bogidiella ’s and their from cold mountain B. Résumé presence springs ( glacialis at 4° C from an altitude of 1900 m in Yugoslavia) La d’une famille stygobies, les diagnose d’Amphipodes in to warm mineral springs ( B. hebraea at 29° C Bogidiellidae, a été revisée. Basée sur une analyse cladistique, and to marine conditions l’ancien été divisé. Dans Israel) infralittoral, (B. genre Bogidiella a sa conception nommés, from of 6 Ischia and actuelle, les Bogidiellidae comprennent onze genres tyrrhenica a depth m on an tandis sept sous-genres et 50 espèces nommées, que plusieurs unnamed form from 42-51 m in the British taxa doivent rester inédits. Les Bogidiellidae sont distribués made in Channel), it, analogy with similar cases dans tous les continents majeurs (abstraction faite de l’Ant- arctique), et dans certains îles océaniques. Cette distri- in Gammarus Amphipoda (the group, Ingolfiella, bution deux vicariantes peut s’expliquer par au moins causes the that more than hadziids), likely one genus majeurs: la fragmentation de la Pangée dans l’ère Mesozoïque An was represented. analysis, others et des mouvements géologiques de régression pendant l’ère using among of the Tertiaire. a cladistic approach, various character Un nombre de des Indes Occidentales été taxa a décrit, that within the states occur group, brought me to nouvelles. comprenant quatre espèces a subdivision of the old good genus Bogidiella into 7 subgenera, whereas a couple of other units INTRODUCTION were separated from it on generic level. Moreover, named and three unnamed number of described Only two bogidiellid a recently genera are included thus far West in the The careful and amphipods were known from the Bogidiellidae. illuminating Indies. The named species treatment Ruffo of the named known two are Bogidiella by 19 taxa bredini in has been a most tool Shoemaker, 1959, from a cave on Barbuda, 1973, valuable for the and B. martini of the Stock, 1978, from wells on Saint start my own work. In present work, 50 Martin. Unnamed taxa have been reported from named taxa (not counting synonyms) have been marine beach used for the construction of a sandy on Curaçao (Stock, 1978), a cladogram (see from a cave on Marie-Galante (Stock, 1980b) and from a cave on Bermuda & Ilife, (Sket 1980). 13 is of *) Report published in the same issue this journal. 346 J. H. STOCK - BOGIDIELLIDAE table I). This steep augmentation in number of sufficient basis for delimiting the Bogidiellidae shows taxa clearly that our knowledge of the hypo- as a family. The discovery of several new taxa has in the outside the in the last few gean amphipods increased explosively Tethyan area years last decade, in particular that of non-European has not facilitated the diagnosis of the Bogi- taxa. diellidae. the In above conception, 50 named species or THE BOGIDIELLIDAE HERTZOG, 1936 subspecies (not counting synonyms) belong to I with Barnard & Karaman's the On the basis of our inter- Essentially, agree family. phylogenetic outspoken opinion (1980: 12) that the bogidiel- pretations (see section on cladistics) we have these 50 in of lids are "not worth superfamily distinction" and arranged species 11 genera, one which is subdivided in 7 table represent only "a weak cluster". subgenera (see I). For the moment, I have retained the bogidiellids as a family, characterized by a combination of char- TABLE I all of are not valid and in the acters, which, however, always Named, genera subgenera family Bogidiel- and number of valid in unit level. charac- lidae, the species assigned, or provi- present every on generic Such sionally assigned, to each. absence of vermiform ters are: eyes; elongated, body (exceptions: Spelaeogammarus, Artesia); 1. Genus Artesia Holsinger, 1980 (1) coxal plates wider than long and not or hardly 2. Genus Spelaeogammarus da Silva Brum, 1976 (1) 3. Genus Somagidiella nov. (1) overlapping (exceptions: again Spelaeogammarus 4. Genus Parabogidiella Holsinger, 1980 (1 + 1 unnamed, and Artesia); reduction of the number of coxal dubious species) gills to three pairs (exceptions: Dussartiella, Bolle- 5. Genus Bogidiella Hertzog, 1933 Subgenus Bogidiella s. str. ( 16) gidia, Parabogidiella, Artesia, Pseudingolfiella , Subgenus Medigidiella nov. (8) Kerguelenicola) associated with P4, P5, and P6; Subgenus Orchestigidiella nov. (1) of 2 smaller than propodus gnathopod (rarely Subgenus Stygogidiella nov. (4 + 1 unnamed species) that of of Subgenus Mexigidiella nov. (4) subequal to) gnathopod 1; carpus Subgenus Guagidiella nov. (2) 1 with gnathopod a distoposterior projection (ex- illo ella sub- Subgenus Ant gidi nov. (1 + 1 unnamed ceptions: Kerguelenicola, Dussartiella, Pseudingol- species ) 6. Actogidiella nov. (1 + 1 unnamed species) fiella); uropod 3 biramous, magniramous (excep- 7. Marigidiella nov. (3) tions: with Dussartiella, Pseudingolfiella), a 1-seg- 8. Kerguelenicola Ruffo, 1974b (1) mented tel- exopodite (exception: Dussartiella); 9. Bollegidia Ruffo, 1974a (2) 10. Dussartiella Ruffo, 1979 (1) son entire, without, or with a shallow, distal notch 11. Pseudingolfiella Noodt, 1965 (2) Artesia (exception: ); pleopodal endopodite seg- fused reduced ments to a single segment, to a Total: Eleven with 7 50 named genera, one subgenera, lodging or absent vestige, entirely (exceptions: Parabogi- species and 4 unnamed species/subspecies. diella, Artesia) ; in the more apomorphous genera, the urosomal Additional pleosomal and/or appendages have Bogidiellidae, not brought to any particular genus of lack of or subgenus because information are known from been for modified, presumably sperm transfer; Ecuador (Sket, 1979), Curaçao in the Antilles (Stock, 1978), oöstegites with limited (terminal) setation only; Marie-Galante in the Antilles (Stock, 1980b and present accessory flagellum short, 1- to 3-segmented; no paper), the English Channel (Spooner, 1959, I960), Réunion Island in the Indian Ocean (Ruffo, 1973), Japan (Matsu- calceoli on A2; mandible palp usually 3-seg- moto, 1976), and Heron Island on the Great Barrier Reef but in some in mented, genera reduction; mandible (unpublished personal observation). These records are not if palp segment 3, present, with relatively simple counted in the above table, but they are incorporated in the distribution map (fig. 2). armature; sexual differences in the gnathopods feeble, in other pereiopods absent (exception: CLADISTICS Orchestigidiella ■); no sternal gills. It remains to be seen if this combination of char- Generalities. — Twenty-three character of them not acters, many synapomorphous, forms states (see table II) have been used to construct BIJDRAGEN TOT DE DIERKUNDE, 51 (2) - 1981 347 and the evolution slowed a cladogram (fig. 1 ) for the Bogidiellidae. Several reduced, rate down, so of these character states relate to secondary sexual that at present the phreatic forms have retained which in character states. I unable differences, presumably play a role copu- many plesiomorphous am to decide which of these two theories is lation (character 23) or sperm transfer (charac- right. it should be out also the ters 11 through 16). However, pointed (see the first that of on of the that It looks disconcerting, at sight, one paragraph biogeography family) the which has coxal the to be a old genera plesiomorphous plates, bogidiellids appear very family, which had over the fresh waters of mesosomal, pleosomal and urosomal appendages, already spread the continents before the of viz. Artesia, has apomorphous mouthparts. Para- break-up Pangaea. This to that in and seems mean were bogidiella agrees mouthparts body append- they already adapted to freshwater life in the earlier with but has coxal Mesozoic, and ages Artesia, apomorphous of that — that these old freshwater taxa plates. Such cases of mosaic evolution (one set supposing evolved from marine ancestors — their ancestors characters being advanced, another set being are now extinct. The living marine taxa, rather retarded) are in reality quite common and do not in few number as compared to the freshwater undermine the utility of cladistic analysis.