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Phylogeny and Biogeography of of Species Bijdragen tot de Dierkunde, 63 (3) 149-161 (1993) SPB Academie Publishing bv, The Hague Phylogeny and biogeography of the Niphargus transitivus group of species (Crustacea, Amphipoda) Boris Sket & Jos Notenboom of Biology, Biotechnical Faculty, University of Ljubljana, Askerceva 12, 61000 Ljubljana, 2 Slovenia; National Institute of Public Health and EnvironmentalProtection, P.O. Box 1, 3720 BA Bilthoven, The Netherlands Keywords: Amphipoda, Niphargus, phylogeny, zoogeography Abstract 1. Introduction The transitivus includes 12 The transitivus introduced Niphargus group groundwater- Niphargus groupwas by dwellingspecies. It is defined by an uprightbody position during Sket (1971) to delimitate four species of Niphargus locomotion,a small, stout body with short appendages, except Schiödte, 1849, which on the one hand were all very V-VI for long coxae and dactyli, coxae equilobateor posterolo- aberrant, but on the other hand intermediate in bate, accessory flagellum subject to reduction, and the body en- some characters between N. kochianus Bate, 1859, dowed with adaptations for volvation ( = ability to enroll into and Karamaniella 1962. The latter a ball). Sket, genus was The of this southeastern European of biogeography group erected for K. pupetta Sket, 1962, a species which species is discussed and a cladistic analysis of the group is pre- did not fit withinthe Niphargus diversity spectrum sented. Some cases of parallel character evolution in ground- at the time. At present Karamaniella is considered water amphipodsare mentioned. The inference of biogeograph- with Niphargus, and K. has ic and phylogenetic information in elucidating the evolutionary synonymous pupetta of the is ofthe data. been into the N. transitivus history group hampered by inconsistency incorporated group(cf. One of the hypotheses about the evolutionary history of the Sket, 1972: 106, 111; Karaman, 1975). For the group is thought to be the most parsimonious. related but morphologically very distinct K. para- doxa(Sket, 1964) the genus CarinurellaSket, 1971, Résumé has been established. Since establishment of the transitivus N. group, Le transitivus du 12 groupe genre Niphargus comprend espèces several additional taxa have been described by souterrains. Il est défini surtout par la positionverticale du corps Dancàu (1971), Karaman (1972, 1984a, 1989), and pendant le déplacement,par un corps petit et trapu aux appen- Karaman& Sket after dices des les (1991). Moreover, reanalysis courts (à l’exception longues coxae et dactyli), par of the known but de- coxaeV-VI equilobés ouposterolobés, par le flagelleaccessoire previously insufficiently à réduction le doué sujet une accusée, et par corps d’adaptions scribed N. kochianus longidactylus Ruffo, 1937, it la volvation. permettant transitivus appeared to belong to the N. group as La biogéographiede cegroupe d’espèces d’Europe du S.E. est well (cf. Sket, 1971; Karaman, 1981). discutée et une analyse cladistique est présentée. Sont men- tionnés certains cas d’évolution parallèle des caractères chez des souterraines. L’utilisation de l’information bio- Amphipodes Altogether 12 nominal species are now consid- géographiqueet phylogénétique dans le but de l’élucidation de ered as the N. transitivus group, and we decided to l’histoire évolutive du groupe est rendue difficile par le caractère treat the systematics, phylogeny, and biogeography contradictoire des données. Une des hypothèses sur l’histoire of this of évolutive group species more properly as du groupe est considérée comme étant la plus par- Niphargus cimonieuse. a whole. 150 B. Sket & J. Notenboom - Phytogeny and biogeography of the Niphargus transitivus group 2. Definition and peculiarities of the Niphargus west Bosnia (Ruffo, 1937; D'Ancona, 1942a& b; Sket & transitivus group Benedetti, 1942; Karaman, 1981; Velkovrh, 1981). 2.1. List of species and distribution • N. transitivus transitivus Sket, 1971; found directly in interstitialwaters and in wells along Pi- the N. and Torrein northeast also The following taxa compose transitivus ave Italy, occasionally group: in caves at the karst margins in the same area • N. carcerarius G Karaman, 1989; two Karaman, . only speci- (Sket, 1971; 1975, 1984a, 1985). • dissonus found mens (!) found in a small cave lake near Pljevlja N. transitivus G . Karaman, 1984; (cave of Sljivanski potok near Djurdjevica Tara), in wells in the Bergamo region, northern Italy northern Montenegro (Karaman, 1989). (Karaman, 1984a, 1985). Sket (1971) opened the • N. asper G. Karaman, 1972; only one specimen question of a possible conspecifity of N. transiti- in well (!) found a near Podgorica (= Titograd), vus with N. pupetta, according to the great varia- Montenegro (Karaman, 1972). bility of some N. transitivus populations. Kara- • N. factor G. Karaman &Sket, 1991; inhabitantof man (1975) denied this possibility since he found small cave lakes in lower parts of Popovo polje no intermediatespecimens in mixed populations, (Vjetrenica at Zavala; Baba pecina at Strujici), but we think that more material should be ex- southeast Hercegovina (Karaman & Sket, 1991). amined before the last decision is conclusive. This is similar to both • N. species very N. asper, may pupetta pupetta (Sket, 1962) (syn.: Karama- be varieties of a single species. niella pupetta Sket, 1962); found in interstitial • N. brevirostris Sket, 1971; only two specimens (!) waters and wells in Bergamo and probably in found in interstitialwaters of the Gacko polje (at Belluno regions in north and northeast Italy, as Licko in southwest Croatia well in northwest Lesee) Lika, (Sket, as (Bovec, new data), central, 1971). and eastern Slovenia (Sket, 1962,1972; Karaman, • N. numerus G. Karaman & Sket, 1991; only one 1975, 1984a). specimen (!) found in a small residual lake of an • N. pupettaparapupetta G. Karaman, 1984 (syn.: River and active cave near Zrmanja (Cavlinska peci- N. parapupetta in Karaman, 1984a, Kara- northern in found in in- na), near Obrovac, Dalmacija (Kara- maniellapupetta Dancâu, 1972); man & Sket, 1991). N. brevirostris and N. numer- terstitial waters and wells near Zagreb, western us also show many similarities pointing to their Croatia, and near the river Dunarea-Dunav possible conspecifity. (Danube) in southwest Rumania. This taxon was • Since it N. pectinicauda Sket, 1971; inhabitantof intersti- described as a separate species. seems to tial of the Sava River between and be and the waters Bohinj completely allopatric to N. p. pupetta, Ljubljana, northwest Slovenia (Sket, 1971; Sket differences are extremely small, we now suggest & Velkovrh, 1981). that both taxa are conspecific. • N. rostratus Sket, 1971; found in small lakes of a • Carinurella paradoxa (Sket, 1964) (syn.: Kara- cave near Cetina River (Dragica pecina at Malj- maniella paradoxa ■ Sket, 1964); found in intersti- kovo), central Dalmacija, Croatia (Sket, 1971). tial waters of the lower reaches of Soca-Isonzo • N. alutensis Dancâu, 1971; probably an inhabi- and its tributaries: Vipava, Torre-Ter, and of tant interstitial waters of the Olt River valley Nadiza-Natisone, all on both sides of the Slo- (found in wells, village Jilbea), south-central veno-Italian boundary (Sket, 1964, 1971, 1972). Rumania attributed this (Dancâu, 1971). Although to a separate genus, spe- • N. longidactylus Ruffo, 1937; inhabitant of inter- cies has drawn much concern. The acceptability stitial able build waters, to large populations in of making the genus(Niphargus) paraphyletic has wells near rivers outside open karst areas, in been discussed elsewhere (Sket, 1990). northeast Italy, and in central and northeast Slovenia, northwest Croatia, central and north- In conclusion, therepresentatives of the N. transiti- - Bijdragen tot de Dierkunde, 63 (3) 1993 151 transitivus vus group are not all sufficiently characterized, 2.2. Characterization of the N. group primarily because of lack of material. Information about morphological variation within and between 2.2.1. Analysis of characters. - The most constant adjacent populations is lacking and the possibility characters of the Niphargus transitivus group nominal in- size that some of the species are merely (some apparently synapomorphies) are: body traspecific varieties must be considered. small to very small, stout and upright-walking; cox- The group includes taxa that are evidently mem- ae of pereiopods II—III very deep, at least twice as of in of bers the benthos larger water bodies the deep as wide; coxa of pereiopod V equilobate or karst; a quite remarkable feature, since in the posterolobate, that of VI posterolobate; bases of Dinaric karst such an ecology is mostly exhibited by posterior pereiopods broad, with widely rounded larger niphargids of a centimetre or larger. Other distoposterior lobes; dactyli of pereiopods very of of the members the group inhabit interstitial waters in slender and long; gnathopods basically N. river also in lon- kochianus of wider valleys, some hilly country (N. type (synapomorphy a group?). gidactylus), but all species may build denser popu- However, there exists a number of characters, lations in open wells - eitherin response to greater present in some of the taxa, that are functionally richer food that above-mentioned characters and space or to supplies are usually connected by the present there. Therefore, the group as a whole, as absent in other niphargids. well as some of the species, appears as quite ubiqui- We will try to evaluateall specific character states animals. tous stygobiontic Some species (N. bre- appearing in
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