ACTA BIOLOGICA SLOVENICA LJUBLJANA 2018 Vol. 61, Št. 1: 93-102 ABS Family Gammaridae (Crustacea: Amphipoda), mainly its Echinogammarus clade in SW Europe. Further elucidation of its phylogeny and taxonomy Družina Gammaridae (Crustacea: Amphipoda), posebej njena veja Echinogammarus v JZ Evropi. Nadaljnja razjasnitev filogenetskih in taksonomskih odnosov Boris Sketa*, Zhonge Houb a Biology department, Biotechnical faculty, University of Ljubljana, p.p. 2995, 1001 Ljubljana, Slovenia b Institute of Zoology, Chinese Academy of Sciences, Beijing 100101, China ^correspondence: boris.sket@bf.uni-lj.si Abstract: Most parts of the Echinogammarus clade of Gammaridae have been appropriately classified with the help of a molecular analysis, ultimately freed of the Echinogammarus-Chaetogammarus malediction. Among defining morphological characters, the gnathopod interrelations are comparatively well congruent with molecular markers. Genus Homoeogammarus distribution area extended from Mediterranean to Atlantic islands. Chaetogammarus and Trichogammarus are firm members of the morphologically very diversified Ponto-Caspian group genera, not closely related to the morphologically similar Echinogammarus, Marinogammarus or Homoeogammarus. Genus Pectenogammarus (along with Neogammarus and Laurogammarus) synonymized with Homoeogammarus. Parhomoeogammarus diagnose corrected, based on topotype samples of the type species. Freshwater species of the 'European Eulimnogammarus' in SW Europe defined as Iberogammarus gen. nov. Continental Homoeogammarus-like, but molecularly distinct group, defined as Dinarogammarus gen. nov. Some here accepted genera are molecularly well supported, some are morphologically difficult to distinguish, but each one is morphologically homogeneous; they are also geographically well defined. In both respects, the very speciose and widely spread Homoeogammarus is different. Keywords: Amphipoda, Gammaridae, systematics, molecular phylogeny, bioge-ography, new genera Izvleček: Večino taksonov klada Echinogammarus družine Gammaridae smo uspeli z molekulsko analizo primerno razvrstiti in končno rešiti prekletstva imen Echinogammarus-Chaetogammarus. Med morfološkimi znaki omogoča velikost gnatopodov I in II klasifikacijo, ki je dokaj skladna s klasifikacijo po molekulskih znakih. Izkazalo se je, da razširjenost rodu Homoeogammarus seže od sredozemskih otokov daleč v Atlantik. Chaetogammarus in Trichogammarus sta člana morfološko zelo razčlenjene ponto-kaspijske skupine rodov in nista blizu morfološko podobnim rodovom Echinogammarus, Marinogammarus ali Homoeogammarus. Rodovi Pectenogammarus, Neogammarus in Laurogammarus so sinonimni s Homoeogammarus. 94 Acta Biologica Slovenica, 61 (2), 2018 Na osnovi topotipskih osebkov tipske vrste smo popravili diagnozo rodu Parhomoe-ogammarus. Sladkovodne vrste 'evropskega Eulimnogammarus' iz JZ Evrope smo definirali kot Iberogammarus gen. nov.. Kontinentalne populacije, podobne rodu Homoeogammarus, vendar molekulsko drugačne, smo definirali kot Dinarogammarus gen. nov.. Nekateri tukaj definirani rodovi so molekulsko podprti, nekatere je težko določiti morfološko, vsak zase pa je morfološko enoten; so tudi geografsko definirani. Vrstno razčlenjen in geografsko zelo razširjen rod Homoeogammarus je težko definirati le morfološko. Ključne besede: Amphipoda, Gammaridae, sistematika, molekulska filogenija, biogeografija, novi rodovi. Introduction The previous attempt to classify the family Gammaridae on numerous samples from the entire distribution area, of the essentially entire ecological range and with most of its supposedly subordinate groups, has shown that a purely morphological characterisation can not give us a phylogenetically plausible system. The most surprising result of molecularly grounded studies is the nesting of endemic, highly aberrant (disparate) endemic 'families' from the lake Bajkal (e.g. Baikalogam-maridae, Macrohectopidae, Pallaseidae; Lowry and Myers 2013, Kamaltynov 1999, Tahteev 2000), within the genus Gammarus; this had been pointed out by Macdonald et al. (2005) and verified by Hou and Sket (2016). Similar is the situation with Pontic families (Pontogammaridae beside Gammaridae; Grabowski 2014, Lowry and Myers 2013) which however still retained a phylogenetic status of genera, but within the family Gammaridae. Gammaridae with shortened endopodite of uropod III make the family particularly difficult to classify. Even among the molecularly well supported genus Gammarus outside the strange Bajkal, there are species, which have erroneously been classified as separate genera, like Fontogam-marus (S. Karaman 1931). The majority of gammarids with shortened uropod III endopodite have been classified as species of Echinogammarus or Chaetogam-marus, some of them changing their position between these two genera, occasionally completed by Marinogammarus and some others. The authors either did not make much effort with reasoning their decisions or the reasoning was not very convincing. But even the very serious attempt of Stock (1968) gave no happy result. After the ancient Schellenberg's (1937a), among the most comprehensive revisions were made by Stock (1968) and by Karaman (1977a). Stock synonymized the taxa Homoeogammarus, Parhomoeogammarus, Ostiogammarus with Echinogammarus, while in Chaetogammarus he united the Pontocaspian member with some Mediterranean and Atlantic species. Karaman included Chaetogammarus, Marinogammarus, Pectenogammarus, the European Eulimnogam-marus into Echinogammarus and rejected the majority of previously used taxonomical characters as not being genus specific. Later, Karaman (1977b) rejected also the genus Neogammarus. Nevertheless, Lowry and Myers (2013) revived all these genera (except Eulimnogammarus). Only recently (Hou and Sket 2016) a molecularly based analysis separated most of above mentioned genera. However, some parts of the family still remained unresolved. The most confusing is the Stock's (1969) 'west European Eulimnogammarus', homonymous with a Bajkalian genus, but evidently being molecularly close to European genera. In this paper we are trying to solve this and some additional problems. Material and methods Samples Animals were caught by a hand net and preserved in 96% ethanol, exchanged after the first fixation. At least taxonomically important parts Sket and Hou: Gammaridae (Amphipoda): phylogeny and taxonomy 95 of the biochemically treated specimens have been preserved for a later morphological examination and/or study. This paper is aimed to answer some phyloge-netic and taxonomical questions left open in Hou and Sket (2016). We could direct our sampling mainly to areas of highest diversity at the genus level, which appeared to be the SW Europe and the shallow seas washing it. This includes the Macaronesian islands, Canaries and Madeira. The main subject of the analysis are groups of the clade Echinogammarus, while the clades Gammarus and Sarothrogammarus have only been used in some comparisons, evaluations. As genera have been designated the sub-clades characterised by a high molecular or morphological singularity (e. g. Iberogammarus) or separated by other branches of such a character (e. g. Chae-togammarus within the Ponto-Caspian group of genera), if at least 80% support. Therefore, our attention was focused on obtaining topotype samples of species, particularly type species of established or potential genera. We only attributed the species identity to samples from type localities and being morphologically adequate. The species identity can only be assured in topotype samples, since even the morphology of a population may change through the year (Pinkster 1988). For some details on sampling see Hou and Sket (2016). Molecular methods Total genomic DNA was extracted from specimens using the Tiangen Genomic DNA kit. Four different gene regions were amplified with primers in Hou et al. (2011), including nuclear fragments of 28S rRNA, 18S rRNA, elongation factor 1a (EF-1a), and a portion of mitochondrial cytochrome oxidase subunit I (COI). Sequence chromatograms were proofed and edited using Sequencher 4.2 DEMO (Gene Codes Corporation, Inc). Sequences were aligned using Clustal X (Thompson et al. 1997) and adjusted by eye using MacClade 4.06 (Maddison and Maddison 2000). The COI and EF-1a fragments were translated using Drosophila mitochondrial DNA or universal genetic code on MacClade to check for the pseudogenes. The best-fitting partitioning schemes and nu-cleotide substitution models were selected using PartitionFinder 1.1.1 with the Bayesian information criterion (Lanfear et al. 2012). The four-partition scheme defined by gene region was selected as best-fitting scheme, 28S with SYM+I+G substitution model, COI with TIM+I+G model, 18S with GTR+I+G model and EF-1a with TrNef+I+G model. The phylogeny was reconstructed under maximum parsimony (MP) and maximum likelihood (ML). MP analyses were performed using PAUP* 4.0b10 (Swofford 2002). All phylogeneti-cally uninformative characters were excluded from the analysis, and gaps were treated as missing data. Heuristic searches were conducted using tree bisection reconnection branch swapping, with a limit of one million rearrangements for each replicate. Bootstrap support indices were generated based on 1000 bootstrap replicates with ten random-addition sequences. ML analysis was performed using RAxML 8.2.9 (Stamatakis 2014), staring with 1000 rapid bootstrap replications followed by a thorough tree search. The GTR-GAMMA model of rate heterogeneity was used for the four-gene partition scheme. To compare the tree topology, ML analysis was implemented using GARLI 2.01 (Genetic Algorithm for Rapid Likelihood Inference; Zwickl, 2006), with four-gene partition model. Results The alignment of the combined data set contained 266 taxa (Table S1) with 5149 base pairs (bp), including 1507 bp for 28S, 656 bp for COI, 2385 bp for 18S, and 601 bp for EF-1a. All new sequences were deposited in GenBank (accession numbers MK159866-MK159939, MK176331, MK176332). The MP and ML analyses produced congruent phylogenetic trees, except for a few disagreements with lower support values. The main discrepancy was tip nodes within the Ponto-Caspian group of genera, with short branches. A number of purposely collected new samplestaxa caused no substantial change in the previously (Hou and Sket 2016) constructed parts of the phylogram, but clarified some previously unclear situations (Fig. 1 and Suppl. Fig. 1). 96 Acta Biologica Slovenica, 61 (2), 2018 Figure 1: Maximum likelihood phylogenetic tree of Echinogammarus based on the combined analysis of mitochondrial (COI) and nuclear (28S, 18S and EF-1a) markers. Numbers represent statistical bootstrap supports of nodes. Taxonomical information is presented in S2. Slika 1: Filogenetsko drevo veje Echinogammarus po analizi maximum likelihood (ML) na osnovi kombiniranih mitohondrijskih (COI) in jedrnih (28S, 18S and EF-1a) podatkov. Številke nad vejami so vrednosti ML bootstrap. Vrstna imena so provizorična, taksonomske rešitve so prikazane v S2. Sket and Hou: Gammaridae (Amphipoda): phylogeny and taxonomy 97 After basally branching off of the Relicto-gammarus and the Sarothrogammarus clades, the gammarids divide into the Gammarus clade with generally plesiomorphic uropod III (with well developed endopodite) and the Echinogammarus clade with generally apomorphic uropod III (with more or less reduced endopodite). The plesiomor-phy of the mentioned character (discussed by Hou and Sket 2016) is not in accord with the sequence of the tree branching. The Echinogammarus clade is further clearly split into a number of well supported monophyla, while the sequence of branching off events is unclear, poorly supported. Well supported groups within the Echinogammarus large clade are: • Parhomoeogammarus clade, • Echinogammarus + Typhlogammarus group genera as well as each of both separately, • 'W European Eulimnogammarus' (sensu Stock 1969) clade, • Marinogammarus clade, • Homoeogammarus + Dinaric + Pontocaspian genera group as well as each of the last mentioned branches separately; Chaetogammarus and Trichogammarus are morphologically similar to Echinogammarus or Homoeogam-marus, but molecularly clearly nested within the morphologically very different and very diverse Pontocaspian group. Beside the strong molecular similarity, there are rarely strong, but mostly weak morphological differences between these branches. Morphologically well distinguished are Echinogammarus s. str., Typhlogammarus group, 'Eulimnogammarus'; less distinct are Marinogammarus, Parhomoe-ogammarus, while morphologically indistinct are geographically and molecularly well defined, mainly coastal Homoeogammarus and the continental Dinaric branch. Discussion Within the morphologically very diverse Ponto-Caspian group, there are again morphologically and molecularly distinct and well supported branchlets: Chaetogammarus and Trichogam-marus (sensu Hou and Sket 2016); they are the only Ponto-Caspians morphologically similar to Homoeogammarus. We will be able to classify the morphologically and molecularly very diverse rest of Ponto-Caspian genera (and species) only when we obtain more complete series of samples from Caspian area. All authors agree that they have to be divided between a number of genera, but their definitions and species contents have been conceived highly diversely (Birstejn and Romanova 1968, Barnard and Barnard 1983, Karaman and Barnard 1979, Stock 1974). It could be shown that Gammarus anisocheirus Ruffo which was (in the lack of the type sample of E. lusitanus) by Hou and Sket 2016 presented as a 'questionable Parhomoeogammarus member', is in fact not related to the type species Gammarus (Parhomoeogammarus) lusitanus Schellenberg 1943. Two populations of P. lusitanus form a well supported clade which includes also Echinogam-marus pacaudi Hubault and Ruffo 1956, thus forming the genus Parhomoeogammarus Schellenberg 1943. On the other hand, G. anisocheirus could now be enthroned a type species of Ibero-gammarus gen. nov., comprising species of the Stock's (1969) 'European Eulimnogammarus', which used to pose a particular phylogenetic and biogeographical enigma. G. anisocheirus has already been attributed to that group by Pinkster and Stock (1970) for its morphological similarity. In our new tree, Iberogammarus is a well supported monophyletic branch with G. anisocherirus and some populations resembling E. grandimanus. (1) Genus Homoeogammarus Schellenberg 1937 (= Echinogammarus p. p.) remains the most speciose and the widely spread group of the Echinogammarus clade, its majority is relatively morphologically unified, with an extensive distribution area in Mediterranean and Atlantic. Beside the previously (Hou and Sket 2016) listed species, according to further molecular analysis it includes also the following members: Marinogammarus atlanticus Dahl, 1958, Neogam-marus festae Ruffo, 1937, Pectenogammarus planicrurus Reid, 1940. So, we can confirm the previous suppositions that the genus Pectenogam-marus Reid, 1940 is superfluous, and so are 98 Acta Biologica Slovenica, 61 (2), 2018 Neogammarus Ruffo, 1937 and Laurogammarus G. Karaman, 1984 (Karaman 1977, Hou and Sket 2016). But inclusion of the molecularly relevant species into Homoeogammarus makes it even more difficult to morphologically diagnose this genus. Opposite to the hint by Hou and Sket 2016, Echinogammarus spinulicornis Pinkster and Stock 1971 is not a member of Parhomoeogammarus, it is now a molecularly proven Homoeogammarus sp., inhabiting freshwater affluents of the Atlantic. Biogeographically important is the fact that some Homoeogammarus spp. are present far within the Atlantic area. H. planicrurus reaches from French Mediterranean coast to the Great Britain. H. cf. atlanticus is present in the Atlantic Madeira, most probably (Dahl 1958; not molecularly proven) it is conspecific with H. atlanticus from Azores; its closest counterpart is the Adriatic H. cf. stocki, while we were not able to obtain the Canarian and topotypic samples of the Mediterranean 'Chaetogammarus' olivii (H. Milne Edwards 1830) with which H. atlanticus might also be conspecific (Stock 1995). Our efforts to sample probably related Gammarus nox Stock 1995 (Madeira) and Chaetogammarus chaetocerus Beyer and Stock 1994 (La Gomera) were not successful. The Italian Peninsula and adjacent big islands are occupied by species of Homoeogammarus also inland (Ruffo and Stoch 2006), while in other parts of the Mediterranean, they are limited to narrow coastal belts (estuaries and coastal springs). (2) The Stock's (1969) taxon 'western European (species of) Eulimnogammarus'' Stock (1969) classified his species Eulimnogammarus macrocarpus Stock 1969, along with Gammarus anisocheirus Ruffo 1959 and with the marine Gammarus obtusatus Dahl 1938 into the noted Bajkalian genus Eulimnogammarus Bazikalova 1945. Molecular analysis has shown that (1) the genuine Bajkalian Eulimnogammarus is phylogenetically a member of the genus Gammarus (Hou and Sket 2016) and endemic to lake Bajkal, well distinguished from Echinogammarus and its relatives, (2) G. obtusatus appears to be a member of Marinogammarus (Hou and Sket 2016), (3) G. anisocheirus is related to E. macrocarpus, but not closely related to Gammarus (or Eulimnogammarus). Our present tree clearly shows that G. anisocheirus and E. macrocarpus can form a separate genus, weakly related to Marinogammarus and Homoeogammarus, but not to Gammarus. This is the rationale of the here established Iberogammarus gen. nov. Iberogammarus gen. nov. syn. western European (species of) Eulimnogammarus Stock 1969 p. p. (Pinkster and Stock 1972). Type species Gammarus anisocheirus Ruffo 1959; type locality is a hygropetric confluent of Neste d'Aure, at St. Lary-Soulan, Hautes-Pyrénées, France (825 m a.s.l.). Additional species: molecularly proven Eulimnogammarus cf. macrocarpus. Not molecularly proven Eulimnogammarus macrocarpus Stock 1969, Eulimnogammarus toletanus Pinkster and Stock 1970. Genus diagnosis. Gammariform amphipods from SW Europe (Iberian Peninsula), similar to Homoeogammarus, but with markedly different gnathopods; gnathopod propodite II only less than 80% length and up to 60% width of propodite I. Uropod III endopodite 20-50% exopodite length, linear, with a terminal and one or more marginal spines; exopodite with marginal groups of spines and variable setation. Distribution. Fresh waters of Central and NE part of Iberian Peninsula, including central Pyrenees. Remark. The most reliable marker of Ibero-gammarus is the smaller and particularly the narrower gnathopod II propodite, which also differs remarkably in shape with its very short palmar margin. Besides, the uropod III endopodite bears also a marginal spine which is normally absent in Homoeogammarus; the setation and the endopodite length are very variable. Sket and Hou: Gammaridae (Amphipoda): phylogeny and taxonomy 99 (3) Correction of the Parhomoeogammarus definition Before obtaining a sample of the Parhomoeogammarus type species, we (Hou and Sket 2016) overestimated the similarity of G. anisocheirus with it. In both, gnathopod I is somehow larger than gnathopod II, which is a comparatively rare character in gammarids. Additional sampling allowed a correction of that mistake. Here, we have samples of two distant populations of the type species while G. anisocheirus was moved to a new genus (Iberogammarus). Corrected data are given here. Parhomoeogammarus Schellenberg 1943 (mended diagnosis) syn. Gammarus (Parhomoeogammarus) Schellenberg 1943. Type species Gammarus (Parhomoeogammarus) lusitanus Schellenberg 1943, type locality Lugar de Mantelâes, Paredes de Coura, Portugal. Additional molecularly proven species Gammarus (Echinogammarus) pacaudi Hubault and Ruffo 1956. Genus diagnosis. Gammariform amphipods from Iberian peninsula and SW France, similar to Homoeogammarus, but gnathopod propodite II may be equal to or slightly shorter (only ca 80% length of) than propodite I, while they are approximately equally wide. Eyes elongated, more than twice as long as wide. Pereopod VII basis proximally convex, distally tapering, without a marked distoposterior lobe. Uropod III endopodite less than 25% exopodite long, scale-shaped to linear, with terminal and marginal spines; exo-podite with marginal groups of spines and long setae. Distribution. The genus seems to be limited to fresh waters in northern Portugal and adjacent NW Spain (all between Porto and A Coruna), SW France and NE Spain (Pinkster 1993). Remarks. For P pacaudi, the dentate posterior margin of pleonites ('l'armature caractéristique du métasome' Hubault and Ruffo 1956) was denoted the primary specific character. In fact, within the type population there are also individuals with unarmed pleonites. Both cohabiting (syntopic) morphs are molecularly indistinguishable. Another putative candidate for this genus, Echinogammarus spinulicornis Pinkster and Stock 1971 (Hou and Sket 2016), appeared to be a Homoeogammarus (see above), although a very aberrant one. (4) Continental Dinaric gammarids Inland parts of the western Balkans host a group of gammarids, morphologically similar and provisionally attached to Homoeogammarus, making that genus paraphyletic. Some additional samples confirmed the geographical distinctness of this group and forced us to establish for it a separate genus. Dinarogammarus gen. nov. Echinogammarus Stebbing, 1899 p.p., Ostiogammarus S. Karaman, 1931 p.p. (e.g. O. acarinatus). Type species Ostiogammarus acarinatus S. Karaman 1931a, Karaman 1970 (syn. Gammarus pungens forma acarinata Schaferna, 1922 p.p.), type locality spring Vrelo Bune, Blagaj, Mostar, Bosnia and Herzegovina. Additional species, molecularly confirmed: Gammarus cari S. Kara-man, 1931b, Ostiogammarus cari bosnensis S. Karaman, 1934. Genus diagnosis. Gammariform amphipods, morphologically indistinguishable from Homoeogammarus. Pereon and pleon dorsally smooth, each telson lobe less than twice as long as broad. Antennae I and II normal, antenna II shorter than antenna I; antena II with short or long, dense or sparse, straight setae. Mouth parts as in Gam-marus. Gnathopods I and II subchelate, propodite II slightly longer than propodite I. Coxal plate IV distoposteriorly lobate. Pereopod VII basis without a distoposterior lobe. Uropods I and II usually normal, with distal and lateral spines. Uropod III exopodite with marginal groups of spines, usually accompanied by long setae that are always straight; endopodite diminished and scale-like, with terminal spine(s) only (without facial or marginal spines) Distribution. Fresh waters (springs and rivers) in Dinaric karst from Ogulin in NW to Mostar in SE (within the rhomboid: Ogulin - Knin - Mostar - Travnik - Ogulin), away from the Adriatic coast. A local contact or shuffling with the coastal Homoeogammarus spp. is not impossible, but at the 100 Acta Biologica Slovenica, 61 (2), 2018 moment, no case of shared locality of both genera was molecularly signalled. Some Dinarogammarus localities are in the Danube drainage (confluents of Sava river), the others in the confluents of the Adriatic. All Homoeogammarus localities in Dinaric area are in the Adriatic drainage, close to coast or even intertidal. (5) The Echinogammarus-Typhlogammarus genera complex appears now to consist of two biogeographi-cally distinct (groups of) genera. The subgroup-genus Echinogammarus (s. str.) inhabits originally epigean fresh waters of SW Europe. The Typhlogammarus group seems to be limited to fresh subterranean waters of Dinaric karst; all species are highly troglomorph. Acknowledgement We are obliged to the collectors of many samples, their names are listed in Suppl. table 1. Thanks also to Pedro Oromi (La Laguna) for his help in the search for Canarian species. The study was financially supported by National Natural Sciences Foundation of China (NSFC-31372156/31422048) to ZH. Thanks to our colleagues Fabio Stoch, Rudi Verovnik and Christophe Piscart for some useful remarks. Thanks to Lučka Sket for her patience and manysided support during field work. We must ask for excuse all colleagues who were or are not able to find some necessary data in Hou and Sket 2016. Due to some technical problems, some files disappeared from 'supporting files', prepared for that paper. References Barnard, J.L., Barnard, C.M., 1983. 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Genetic algorithm approaches for the phylogenetic analysis of large biological sequence datasets under the maximum likelihood criterion. PhD thesis, University of Texas at Austin, Austin, TX. GARLI, version 1.0. Published by the author. Available at: http://www. nescent.org/wg_garli SUPPLEMENT / PRILOGA Family Gammaridae (Crustacea: Amphipoda), mainly its Echinogammarus clade in SW Europe. Further elucidation of its phylogeny and taxonomy Družina Gammaridae (Crustacea: Amphipoda), posebej njena veja Echinogammarus v JZ Evropi. Nadaljnja razjasnitev filogenetskih in taksonomskih odnosov Boris Sketa*, Zhonge Houb a Biology department, Biotechnical faculty, University of Ljubljana, p.p. 2995, 1001 Ljubljana, Slovenia b Institute of Zoology, Chinese Academy of Sciences, Beijing 100101, China ^correspondence: boris.sket@bf.uni-lj.si Supplement 1: Priloga 1: Maximum likelihood phylogenetic tree of Echinogammarus based on the combined analysis of mitochondrial (COI) and nuclear (28S, 18S and EF-1a) markers. Numbers represent statistical bootstrap supports of nodes. Taxonomical information is presented in S2. Filogenetsko drevo veje Echinogammarus po analizi maximum likelihood (ML) na osnovi kombiniranih mitohondrijskih (COI) in jedrnih (28S, 18S and EF-1a) podatkov. Številke nad vejami so vrednosti ML bootstrap. Vrstna imena so provizorična, taksonomske rešitve so prikazane v S2. Sket and Hou: Family Gammaridae (Crustacea: Amphipoda), mainly its Echinogammarus clade in SW Europe. Further elucidation of its phylogeny and taxonomy Supplement 2: Taxa information and GenBank numbers. OTUs are listed in the same order as in the S1. *Taxa names in parentheses are taken from literature unchanged. Species marked with TG are type species of genera. Species names followed with "Typ" or "typ" represent the topotype populations or populations close to "Typ". The "cf." and "ag." are abbreviations of "confer" and "aggregate or group". fF = Freshwater; S = Saline (brackish to marine water). Priloga 2: Informacija o taksonih in številke v GenBank. OTU so v istem zaporedju, kot v drevesu S1. *Imena taksonov v oklepajih so nespremenjena povzeta po literaturi. S 'TG' označene vrste so tipi rodov. Če imenu vrste sledi 'Typ' ali 'typ' je osebek iz topotipske populacije ali blizu nje. 'cf.' ali 'ag.' sta okrajšavi za "confer" ali "aggregate ali skupina (group)". fF = sladkovodno; S = slana (somornica ali morska) voda. Working names; emphasized genus name Designation of own samples in the Collector(s) or Locality Habitat Country Voucher number Ecological 28S CÜI 18S EF-1a valid for all taxa below it (not so within tree Reference (r. = stream; c. = cave; N, S, statef Ponto-Caspian or Typhlogammarus group). (with year). W, E = north, south, west, east) Taxon* Ponto-Caspian group genera Black Sea - Caspian -surroundings Dikerogammarus villosus 7 (Sowinsky, 1894) 052-D.vil-RÜM B. Sket, C. Fiser Babadag, Jurilovca, Limanu Razim coast of liman Romania SLOCHN052 S KF478442 KF478533 KF478613 - River Waal Dikerogammarus villosus 4 894DvillosusNLD D.Platvoet, S. Li (Rhine Branch) near Nijmegen river Netherlands IZCASIA0894 F KF478495 KF478581 -- Dikerogammarus villosus 5 901DvillosusNLD D.Platvoet, S. Li Gouwzee, dam to Marken, near Amsterdam river Netherlands IZCASIA0901 F KF478496 KF478582 -- (Dikerogammarus villosus) 1 Cristescu & Hebert 2005 - ? AY529068 AY529048 -- Dikerogammarus villosus 8 059-D.vil-UKR-Kry B.Sket, M.Zagmajster Kyiv, r. Dnipru river Ukraine SLOCHN059 F KF478449 KF478540 -- Dikerogammarus villosus cf. bispinosus Martynov 1925 309Dkg (309-Dkg-bis-UKR) B.Sket, C.Fiser Dnistrovski liman, liman Ukraine SLOCHN309 ? MK159866 MK159941 MK160009 Dikerogam v. villosus 385Dkg (385-Dkg-v-Azov-RUS) S.Utevsky, M.Kolesnykova Morskoj Culek, Taganrog brook Russia SLOCHN385 ? MK159867 MK159942 MK160010 (Dikerogammarus villosus) 2 Macdonald et al. 2005 Black Sea - S - - AY926766+ AY926828 - (Dikerogammarus villosus) 3 Hou et al. 2007 Havel Brandenburg Germany IZCASIA0676 F EF582941 EF570297 EF582898 - (Dikerogammarus caspius) Macdonald et al. 2005 Caspian Sea - S - - AY926765+ AY926827 - Dikerog cf. haemobaphes fluviatilis Martynov, 1919 300Dkg (300-Dkg-fluv-UKR) B.Sket, C.Fiser r. Turuncuk, Biliaivka, Odesa stream Ukraine SLOCHN300 F MK159868 MK159943 MK160011 Dikerogam haem cf. haemobaphes (Eichwald, 1841) TG 301Dkghaemob (301-Dkg-haemob-UKR) B.Sket, C.Fiser Dnipro, chan. Cajka, Hola prystan, Kherson stream Ukraine SLOCHN301 F MK159869 MK159987 MK160012 Dikerogam v. villosus 412Dkg (412-Dkg-h-Odesa-UKR) B.Sket, C.Fiser Biliaivka, Odesa brook Ukraine SLOCHN412 F MK159870 MK159988 MK160013 Dikerogammarus cf. villosus 6 186-Dkg-UKR S.Utevsky, M.Kolesnykova Migija, r. Juznyj Bug river Ukraine-S SLOCHN186 F KF478483 KF478569 -- (Dikerogammarus haemobaphes) Cristescu & Hebert 2005 - ? AY529069 AY529049 -- Obesogammarus crassus (Sars 1894) 055-P-cra-RÜM B. Sket, C. Fiser Babadag, Jurilovca, Limanu Razim coast of limanu Romania SLOCHN055 ? KF478445 KF478536 KF478615 - (Obesogammarus crassus) 1 Cristescu & Hebert 2005 Bandar-e Anzali Iran - S AY529061 AY189482 -- 5 (Obesogammarus crassus) 3 Cristescu et al. 2003 Lake Kuhal Ukraine - S - AY189481 -- Obesogammarus crassus 4 056-P-cra-UKR P. Trontelj Harkiv, Biskin, r. Severski Donec river Ukraine SLOCHN056 F KF478446 KF478537 -- Obesogammarus crassus 5 058-P-cra-UKR-Kry B.Sket, M. Zagmajster Kyiv, r. Dnipru river Ukraine SLOCHN058 F KF478448 KF478539 -- (Obesogammarus crassus) 2 Cristescu et al. 2003 Lake Razim Romania - S - AY189478 -- Pontogammarus obesus 4 (G.O. Sars, 1894) TG 053-P-obs-ROM B. Sket, C. Fiser Tulcea, Nufaru, r. Dunarea/ Danube river bottom Romania SLOCHN053 F KF478443 KF478534 -- cf . Stenogammarus sp. 303Gmelnps (303-Gmelnps-UKR) B. Sket, C. Fiser Dnipro, chan. Cajka, Hola prystan, Kherson stream Ukraine SLOCHN303 F MK159871 MK159944 MK160014 cf. Shablogammarus sp. 2 307Shablg (307-Shablg-UKR) B. Sket, C. Fiser Dnipro, chan. Cajka, Hola prystan, Kherson Ukraine SLOCHN307 F MK159872 - MK160015 Turcogammarus spandli1 (S.Karaman 1931) 026-D.span-Vol 026 C.Fiser, R.Verovnik Thessaloniki, Mikri Volvi lake Greece SLOCHN026 F KF478437 KF478528 KF478611 - Turcogammarus cf. spandli 438Dikg (438-Dikg-Thessal-HEL) Apollonias Stream, Thessaloniki stream Greece F MK159873 MK159945 MK160016 Turcogammarus spandli 2 028-D.span-Nes 028 C.Fiser, R.Verovnik Xanthi, Stathmos, r. Nestos brook Greece SLOCHN028 F KF478438 KF478529 -- Pontogammarus cf. robustoides 7 057-P-rob-UKR P. Trontelj Kharkiv, Biskin, r. Severski Donec river Ukraine SLOCHN057 F KF478447 KF478538 KF478616 - Dikerogam h. haemobaphes 386Dkg (386-Dkg-h-Azov-RUS) S.Utevsky, M.Kolesnyko Taganrogskij zaliv, Morskoj Culek, Taganrog Russia SLOCHN386 ? MK159874 MK159946 MK160017 (Pontogammarus robustoides) 4 Cristescu & Hebert 2005 Ukraine Baltic Sea ? - S - AY529045 -- Pontog robustoides 4 419Pgrobust (419-Pg-robust-UKR) S.Sidorovsky r. Pivdennyi Bug, Mygiya, Pervomaysk Distr., Mykolayiv Reg. Ukraine SLOCHN419 F MK159875 MK159947 MK160018 Obesogammarus cf. acuminatus Stock et al., 1998 202-Obsg-ac-T-KAS A.Mirzajani, B.Sket Gilan Prov., Bandar-e Anzali, Anzali Thalab laguna Iran-Caspian SLOCHN202 S KF478487 KF478574 KF478634 - Pontogammarus robustoides 8 (G.O. Sars, 1894) TG 255-Ptg-robudes-RUS M.Schletterer delta of r. Volga river Russia-Caspian SLOCHN255 ? JF965822 JF965990 JF966184 - (Pontogammarus robustoides) 5 Cristescu & Hebert 2005 Astrakhan Russia - ? - AY529046 -- (Pontogammarus robustoides) 6 Cristescu & Hebert 2005 Volgograd Reservoir reservoir Russia - F AY529067 AY529047 -- Pontogammarus cf. aralensis (Uljanin, 1875) 182-Ptg.aral-UKR S.Utevsky, M.Kolesnykova Odesskaja Oblast, Katlabuh lake Ukraine SLOCHN182 ? KF478482 KF478568 KF478631 - (Pontogammarus robustoides) 1 Cristescu & Hebert 2005 Lake Ramiz liman Romania - S - AY529042 -- (Pontogammarus robustoides) 2 Cristescu & Hebert 2005 Lake Kahul liman Ukraine - S - AY529043 -- (Pontogammarus robustoides) 3 Cristescu & Hebert 2005 Dnieper River river - F AY529066 AY529044 -- Euxinia maeotica 8 (Sovinsky 1893) TG 054-P-mae-ROM B. Sket, C. Fiser Mangalia, Black sea sea littoral Romania SLOCHN054 S KF478444 KF478535 KF478614 - (Euxinia maeoticus) 4 Cristescu et al. 2003 Black Sea mar. littoral Ukraine - S - AY189490 -- Euxinia maeotica 311Euxinia (311-Euxinia-UKR) B. Sket, C. Fiser Black Sea coast, Pokrovka, Kherson mar. littoral Ukraine SLOCHN311 S MK159876 MK159948 MK160019 6 (Euxinia maeoticus) 7 Cristescu & Hebert 2005 Chernomorka Ukraine - S - AY529038 -- (Euxinia maeoticus) 3 Cristescu et al. 2003 Black Sea Ukraine - S - AY189483 -- (Euxinia maeoticus) 1 Cristescu & Hebert 2005 Black Sea Romania - S AY529062 AY189494 -- Euxinia maeotica 9 204-Pont-maeo-KAS A.Mirzajani, B.Sket Gilan Prov., NW Bandar-e Anzali, Caspian sandy beach Iran-Caspian SLOCHN204 S KF478488 KF478575 KF478635 - (Euxinia maeoticus) 5 Cristescu et al. 2003 Bandar-e Anzali Iran - S - AY189503 -- (Euxinia maeoticus) 6 Cristescu et al. 2003 Lankaran Azerbaijan - S - AY189504 -- (Euxinia maeoticus) 2 Cristescu & Hebert 2005 Baku Azerbaijan - S AY529063 AY189500 -- Euxinia maeotica 387Euxinia (387-Euxinia-Azov-RUS) S.Utevsky, M.Kolesnykova Morskoj Culek, Taganrog sandy beach Russia SLOCHN387 ? MK159877 MK159949 MK160020 Stenogammarus cf. similis (Sars 1894) 187-Steng-?sim-KAS A.Mirzajani, B.Sket Gilan Prov., Lisar, Caspian lake, - 15 m Iran-Caspian SLOCHN187 S KF478484 KF478570 KF478632 - (Pontogammarus obesus) 1 Cristescu & Hebert 2005 Lake Sinoie liman Romania - S AY529064 AY529039 -- (Pontogammarus obesus) 2 Cristescu & Hebert 2005 Volgograd Reservoir reservoir Russia - F AY529065 AY529040 -- (Pontogammarus abbreviatus) Macdonald et al. 2005 Caspian Sea lake ? - S - AY926691 AY926810+ AY926871 - (Pontogammarus obesus) 3 Cristescu & Hebert 2005 ? - S - AY529041 -- Niphargogammarus aequimanus (Sars, 1895) 206-Niphgg-aeq-KAS A.Mirzajani, B.Sket Gilan Prov., Lisar, Caspian lake, - 15 m Iran-Caspian SLOCHN206 ? KF478489 KF478576 KF478636 - Paraniphargoides cf. motasi (Carauçu 1943) cfTG 188-Niphggd-?mot-KAS A.Mirzajani, B.Sket Gilan Prov., Jireh bagh, Caspian lake, - 10 m Iran-Caspian SLOCHN188 ? KF478485 KF478571 KF478633 - cf. Akerogammarus-Chaetogammarus 298Stg (298-?Stg-DniprL-UKR) B.Sket, C.Fiser Dniprovski liman, Herois'ke, Kherson liman Ukraine SLOCHN298 S MK159878 MK159950 MK160021 Trichogammarus trichiatus (Martynov, 1932) TG 299Chtgtrich (299-Chtg-trich-UKR) B.Sket, C.Fiser Hola prystan, Kherson Ukraine SLOCHN299 F MK159879 MK159951 MK160022 Trichogammarus trichiatus 310Chtg (310-Chtg-trich-UKR) B.Sket, C.Fiser Dnistrovski liman, Ovidiopol', Odesa liman Ukraine SLOCHN310 S MK159880 MK159989 MK160023 (Echinogammarus trichiatus) 1 Cristescu & Hebert 2005 Krasnaya Kosa, Dniestr Liman Ukraine - S - AY529050 -- (Echinogammarus trichiatus) Cristescu & Hebert 2005 Romania - S AY529072 - -- (Echinogammarus trichiatus) 2 Cristescu & Hebert 2005 Istria, Sinoie Lake Romania - S - AY529051 -- Chaetogammarus cf. ischnus (Stebbing, 1899) 051-Ch-ich-ROM B. Sket, C. Fiser Babadag, Jurilovca, Limanu Razim coast of limanu Romania SLOCHN051 S KF478441 KF478532 KF478612 - Chaetogammarus cf. placidus (Sars, 1896) 126-C.placid-ROM C.Fiser, B.Sket Babadag, Jurilovca, Lacul Razim limanu Romania SLOCHN126 S KF478466 - -- Chaetogammarus cf. warpachowskii (Sars, 1894) 130-f?C.warpa-ROM B. Sket, C. Fiser Babadag, Jurilovca, Lacul Razim limanu Romania SLOCHN130 S - KF478556 -- (Echinogammarus ischnus) 1 Cristescu & Hebert 2005 Romania - ? AY529070 AY326120 -- 7 (Echinogammarus ischnus) 2 Cristescu & Hebert 2005 Volga River Delta Russia - ? AY529071 AY326125 -- (Echinogammarus ischnus) 5 Cristescu & Hebert 2005 Volga River Delta Russia - ? - AY326126 -- Cristescu & Hebert 2005 Novodniestrovsk, Middle (Echinogammarus ischnus) 4 Dniestr Ukraine - F - AY326124 -- River (Echinogammarus ischnus) 3 Cristescu & Hebert 2005 r. Irpen' Ukraine - F - AY326122 -- Chaetogammarus cf tenellus (G.O. Sars 1914) cf TG 302Chtg (302-Chtg-Khers-UKR) B. Sket, C. Fišer r. Dnipro, chan. Hola pristan, Kherson Ukraine SLOCHN302 F MK159881 MK159952 MK160024 Dikerogammarus caspius (Pallas, 1771) 493Dikg (493-Dikg-casp-RUS) M.Schletterer delta Volgi Russia SLOCHN493 ? MK159940 - V Gmelina costatal Sars, 1894 TG 050-Gm.cos-ROM B. Sket, C. Fišer Babadag, Jurilovca, Limanu Razim coast of limanu Romania SLOCHN050 S KF478467 - KF478624 - Gmelina costata 2 128-Gmelin-ROM B. Sket, C. Fišer L Razim, Dobrogea, plitvina Romania SLOCHN128 S KF478468 - KF478625 - Gammaridae 519Gdae T.Delic Krušničko vrelo, Krušnica, spring Bosnia- F (cf. Gmelina, ovig., tiny) (519-Gdae-Krupa-BiH) Bos. Krupa Herzegovina (Pontogammarus crassus) Macdonald et al. 2005 Black Sea - S - - AY926811+ AY926872 - Jugogammarus kusceri 1 (S.Karaman, 1931) (typ) TG 073-Jugog-SLO B.Sket Ivančna Gorica, Krka, springs of r. Krka spring Slovenia SLOCHN073 F KF478462 KF478552 KF478622 - Jugogammarus kusceri 2 (typ) 103-Jugog-SLO B.Sket Ivančna Gorica, Krka, springs of r. Krka spring Slovenia SLOCHN103 F KF478463 KF478553 -- Amathillinapusilla G.O. Sars, 1896 304Amathln (304-Amathln-UKR) B.Sket, C.Fišer Dnipro, chan. Čajka, Hola prystan, Kherson Ukraine SLOCHN304 F MK159882 - MK160025 (Amathillina pusilla) Macdonald et al. 2005 - ? - - AY926756+ AY926818 - Gmelinopsis cf. tuberculata G.O. Sars, 1896 cf TG 305Gmlnps (305-Gmlnps-UKR) B.Sket, C.Fišer Dnipro, chan. Čajka, Hola prystan, Kherson Ukraine SLOCHN305 F MK159883 MK159953 MK160026 Croatia - Bosnia- genus Dinarogammarus Herzegovina (Dinaric karst) Echinogammarus cari (S.Karaman, 1931) (Typ) 010-E.cari-Bis 010 B.Sket Tounj, Mikašinovici, r. Bistrac brook Croatia SLOCHN010 F JF965821 - JF966183 - (Echinogammarus) Dinarogammarus acarinatus 082-E.acar-BiH B.Sket Mostar, Blagaj, vrelo Bune spring Bosnia-Herzegovina SLOCHN082 F KF478458 KF478548 KF478620 - (S.Karaman 1931) (Typ) TG Echinogammarus sp6 131-E.Pliva-BiH B.Sket Jajce, Mile, outflow of Plivsko jezero brook Bosnia-Herzegovina SLOCHN131 F KF478469 KF478557 -- Echinogammarus acarinatus bosnensis (S.Karaman 1934) (Typ) (Synonym E. acarinatus) 133-E.bosn-T-BiH B.Sket Travnik, Šumece spring Bosnia-Herzegovina SLOCHN133 F KF478471 KF478559 KF478626 - Dinarogammarus sp. 518Eg (518-Eg-Krupa-BiH) T.Delic Krušničko vrelo, Krušnica, Bosanska Krupa spring Bosnia-Herzegovina SLOCHN518 F MK159884 MK159954 MK160027 Homoeogammarus sp. 535Hmg (535-Hmg-Krka-HRV) B.Sket Krka below spring, Krško vrelo, Knin stream Croatia SLOCHN535 F MK159885 MK159955 MK160028 Echinogammarus sp 9 158-E.Knin-HRV B.Sket Krško vrelo, Knin spring-river Croatia SLOCHN158 F KF478479 KF478565 - - Echinogammarus sp 7 132-E.Una-BiH B.Sket above Bihac, motel Sunce, r. Una river Bosnia-Herzegovina SLOCHN132 F KF478470 KF478558 -- 8 circum-Mediterranean - genus Homoeogamamrus /\LI ill I LI C Echinogammarus monomerus (Stock, 1977) 107-E.mon-ESP D.Jaume isl. Mallorca, Soller, Font Fomalutx spring Spain SLOCHN107 F KF478464 KF478554 KF478623 - Sardegna, prov. Nuoro, Echinogammarus cf. tibaldii Pinkster and Stock, 1970 243-E.Sarde-ITA F.Stoch Supramonte, Complesso Carsico di Su Cologone (Oliena) spring Italy SLOCHN243 F KF478492 KF478579 -- Neogammarus nudus Stock 1971 156-Neog.nu-FRA B.Sket Nice, Cagnes-sur-Mer beach-brook France SLOCHN156 S KF478477 KF478564 KF478630 - Neogammarus nudus 1 C156 (156-C-Neog-nud-FRA) B.Sket Nice, Cagnes-sur-Mer beach-brook France SLOCHN156C S KF478478 - -- Neogammarus festai (Ruffo 1937) 435Neog (435-Neog-fest-FRA) B.Sket Calanques-Salees, Marseilles mar. littoral France SLOCHN435 S MK159886 MK159990 MK160029 Marinogammarus atlanticus Dahl 1958 C288Matlan (C288-M-atlan-POR-M) B.Sket estuar NE del plaže, Machico aestuar PortugalMadeira SLOCHN288c S MK159887 MK159956 - Marinogammarus atlanticus 486Mg (486-Marg-atl-POR-Mad) B.Sket Lugar do Baixo, Punta do Sol aestuar PortugalMadeira SLOCHN486 S MK159888 MK159957 MK160030 Marinogammarus atlanticus 290Mfunch (290-M-Funch-POR-M) B.Sket Praia Formosa, Funchal mar. littoral PortugalMadeira SLOCHN290 S MK159889 MK159991 MK160031 Marinogammarus atlanticus C289Mrbra (?:289-M-RBrava-POR-M) B.Sket Ribeira Brava aestuar PortugalMadeira SLOCHN289c S MK159890 MK159958 MK160032 Marinogammarus atlanticus 288Matlan (288-M-atlan-POR-M) B.Sket Machico aestuar PortugalMadeira SLOCHN288 S MK159891 - MK160033 Marinogammarus atlanticus C290Mfunch (?: 290-M-Funch-POR-M) B.Sket Praia Formosa, Funchal mar. littoral PortugalMadeira SLOCHN290c S MK159892 MK159959 MK160034 Echinogammarus sp. 337Elopar (337-E-Lopar-HRV) B.Sket Zidine, Lopar, Rab isl. intertidal Croatia SLOCHN337 S MK159893 MK159960 MK160035 Echinogammarus cf. stocki G. Karaman, 1970 283Elopar (283-E-Lopar-HRV) B.Sket uvala Sice, Lopar, Rab isl. coastal spring Croatia SLOCHN283 S MK159894 MK159961 MK160036 Echinogammarus foxi 1 (Schellenberg 1928) 142-E.Estram-FRA B.Sket Leucate, Salses-de-Chateau, Font d'Estramar spring France-SE SLOCHN142 S KF478472 KF478560 KF478627 - Echinogammarus sp8 145-E.Fees-FRA B.Sket Perpignan, Leucate, Grotte de Fees cave France-SE SLOCHN145 S KF478475 KF478562 -- Echinogammarus cf. pungens 1 (Milne-Edwards, 1840) 078-Echg-FRA B.Sket Perpignan, Leucate, Grotte des Fees cave France SLOCHN078 S KF478457 KF478547 -- Echinogammarus. cf. foxi 450Egfox (450-Eg-cf-foxi-Nice-FRA) G.Bračko, M.Hrovat. Bouche-de-Loup, Villeneuve de Loubet, Nice aestuar France SLOCHN450 S MK159895 MK159992 MK160037 Homoeogammarus cf. monomerus-platvoeti-tabu 531Hg (531-Hmg-Antibes-FRA) B.Sket mouth La Brague, Biot, Antibes intersticial France SLOCHN531 S MK159896 MK159993 MK160073 Echinogammarus cf. olivii (Milne Edwards, 1830) 397Estrunj (397-E-Strunj-SLO) B.Sket hotel Salinera, Strunjan brack. spring Slovenia SLOCHN397 S MK159897 MK159962 MK160038 Echinogammarus cf. foxi 2 175-E.Ankar-SLO B.Sket Ankaran gravel beach Slovenia SLOCHN175 S KF478481 KF478567 -- Pectenogammarus planicrurus 451Pecteng (451-Pecteng-plan-GB = 446) J.Ironside Aberystwyth, W Wales littoral UK SLOCHN451 S MK159898 MK159994 MK160039 Pectenogammarus planicrurus 446Pectg (446-Pectg-planicr-UK) J.Ironside Aberystwyth, W Wales littoral UK SLOCHN446 S MK159899 MK159963 MK160040 Echinogammarus cf pungens 318Ercm (318-E-RCrn-CrG) B.Sket Rijeka Crnojevica, Cetinje stream Montenegro SLOCHN318 F MK159900 MK159995 MK160041 E cf. thoni (weak bulges) 457Erisan (457-E-Risan-CrG) B.Sket D. Morinj, Risan Montenegro SLOCHN457 F MK159901 MK159996 MK160042 9 Echinogammarus cf. pungens 277Ercmoj (277-E-RCmoj-CrG) B.Sket Rijeka Crnojevica, Cetinje stream Montenegro SLOCHN277 F MK159902 - MK160043 Laurogammarus thoni 3 064-E.tho-MO-CrG B.Sket Virpazar, Crničko polje, Malo oko spring Montenegro SL0CHN064 F KF478451 - -- Echinogammarus sp. 1 012-E.xxx-Cav 012 B.Sket Cavtat, hotel Epidaurus brook Croatia SL0CHN012 F KF478435 KF478526 -- Echinogammarus thoni 4 069-Etho-Trs-HRV C.Fišer Dubrovnik, Trsteno spring Croatia SL0CHN069 F KF478434 KF478525 KF478610 - Echinogammarus thoni 1 (Schaferna, 1922) 011-E.tho-Kom 011 B.Sket Metkovic, Komin, s. Modro oko spring Croatia SL0CHN011 F KF478433 KF478524 KF478609 - (Typ; TG Laurogammarus) Echinogammarus thoni 5 118-E.Nere-HRV G.Bračko Metkovic, r. Neretva river Croatia SL0CHN118 F KF478465 KF478555 -- Echinogammarus ruffoi 2 Typ 445Egruffoi (445-Eg-ruffoi-ITA-T) F.Stoch Roggia, tra Cascina Prada e Ponte Rosso, Italy SLOCHN445 F MK159903 MK159997 MK160044 Echinogammarus pungens 2 088-E.pun-HRV G.Bračko Metkovic, r. Neretva river Croatia SL0CHN088 F KF478459 KF478549 KF478621 - Echinogammarus cf antalyae 341Epakle (341-E-Paklen-HRV) B.Jalžic Markova špilja, Seline, Starigrad, Paklenica cave Croatia SL0CHN341 F MK159904 MK159964 MK160045 Echinogammarus sp. 506Erisan (506-E-Risan-CrG) T.Delic Risan aestuary Montenegro SL0CHN506 ? MK159905 MK159965 MK160046 Echinogammarus sp 2 024-E.xxx-Kot 024 B.Sket Kotor spring Montenegro SL0CHN024 F KF478436 KF478527 -- Echinogammarus sp 3 035-E.xxx-Krk 035 B.Sket island Krk, Vrbnik spring Croatia SL0CHN035 F KF478439 KF478530 -- Echinogammarus sp 4 072-Ech.sp-Zadar-HRV R.Verovnik Zadar, Pirovac spring Croatia SL0CHN072 F KF478452 KF478542 -- Echinogammarus cf. veneris 1 076-E.ven?-ITA C.Fišer Gargano, Carpino, spring at restaurant 'da Carlo' spring Italy SL0CHN076 F KF478455 KF478545 -- Echinogammarus sp 447Plovniy ? SL0CHN447 ? MK176331 MK176332 - Echinogammarus cf. veneris 2 077-E.ven?-ITA P.Trontelj, B.Sket S Verona, Isola, stream at road brook Italy SL0CHN077 F KF478456 KF478546 KF478619 - Echinogammarus cf. veneris 3 (Heller 1865) 238-E.ex-ven-CYP R.Verovnik Polis, Akamas, s. Loutra tis Afroditis spring Cyprus SL0CHN238 F KF478490 KF478577 - KF478659 Echinogammarus cf. stammeri 1 (S. Karaman, 1931) (Typ) 074-E.stam-ITA B.Sket Trst/Trieste, Monfalcone/ Tržič, s. Polosco spring Italy SL0CHN074 F KF478454 KF478544 KF478618 - Echinogammarus sp 10 163-E.Bolju-ITA B.Sket Trieste/Trst, Bagnolli/ Boljunec, tributary of Glinščica brook Italy SL0CHN163 F KF478480 KF478566 -- Echinogammarus cf. pungens 3 197-E-Clitu-ITA C.Douady, F.Malard, F.Stoch Perugia, Campello sul Clitunno, Fonti di Clitunno spring Italy SL0CHN197 F KF478486 KF478572 -- Echinogammarus cf. stammeri 2 242-E.Verona-ITA B.Sambugar S Verona, r. Menago -Buttapietra resurgence Italy SL0CHN242 F KF478491 KF478578 -- Laurogammarus scutarensis 1 (Schaferna, 1922) (Typ) 033-L.scu-Rib 033 B. Sket Podgorica, springs of Ribnica spring Montenegro SL0CHN033 F KF478453 KF478543 KF478617 - Echinogammarus cf. thoni2 063-E.tho-GZt-CrG B.Sket Glava Zete, Dobro Polje, at the mil spring Montenegro SL0CHN063 F KF478450 KF478541 -- Laurogammarus scutarensis2 036-L.scu-RCr 036 K.Jazbec Rijeka Crnojevica spring Montenegro SL0CHN036 F KF478440 KF478531 -- Echinogammarus cf. platvoeti Pinkster, 1993 258-E.Estram-FRA B.Sket Leucate, Salses-de-Chateau, Font d'Estramar spring France-SE SL0CHN258 F KF478493 - KF478637 - Echinogammarus pungens Typ 455Epung (455-E-pungens-T-FRA) B.Sket Salses-Chateau, Leucate spring France.SE SL0CHN455 S MK159906 MK159966 MK160047 10 Echinogammarus spinulicornis (Pinkster & Stock, 1971) Typ 523Eg (523-Eg-spinlcrn-FRA) B.Sket Touvre I spring, Angouleme, NE Bordeaux spring France.SE SLOCHN523 F MK159907 MK159967 MK160048 Echinogammarus cyrtusl Pinkster and Platvoet 1986 (Typ) 144-E.cyrtu-T-FRA B.Sket Montpellier, Sources-du-Lez river France-SE SLOCHN144 F KF478473 - KF478628 - Echinogammarus cyrtus 2 144b-E.cyrtu-T-FRA B.Sket Montpellier, Sources-du-Lez river France-SE SLOCHN144b F KF478474 KF478561 -- Echinogammarus cf. simoni (sp 5) 093-E.Cheb-TUN B.Sket Tozeur, oasis Chebika brook Tunisia SLOCHN093 F KF478460 KF478550 -- Echinogammarus cf. simoni (Chevreux, 1894) cf TG 094-E.?sim-TUN B.Sket Tozeur, oasis Tamerza brook Tunisia SLOCHN094 F KF478461 KF478551 -- Echinogammarus cf. simoni 482GaeBeni J.Notenboom, F.Malard, C.Douady Font des Admiraors, Spain SLOCHN482 F MK159908 MK159968 (setae scarce) (?: 482Gae-BeniMaur-ESP) Benimaurel, Alicante Echinogammarus cf. simoni (few setae, uropod III nearly without them) 484Gae (?:484-Gae-Deifont-ESP) J.Notenboom, F.Malard, C.Douady Source de Deifontes, ?Granada Spain SLOCHN484 F MK159909 MK159998 - Echinogammarus cf. simoni (nearly no setae, uropod III styliform) 485Gae (?:485-Gae-Sorbas-ESP) J.Notenboom, F.Malard, C.Douady Fuente del Peral, Sorbas, ?Almeria Spain SLOCHN485 F MK159910 - - genus Marinogammarus E & NW Atlantic Echinogammarus marinus (Leach, 1815) G.Yang Thurso Bay, on the north coast of Scotland UK IZCASIA0700 S KF478494 KF478580 KF478638 - (Chaetogammarus marinus) Macdonald et al. 2005 Bergin Norway - S - AY926655 AY926760+ AY926822 - (Chaetogammarus obtusatus) 2 Englisch et al. 2003 Baltic Sea ? - S - - AF419224 - Marinogammarus cf. marinus 431MgMarinus (431-Mg-marinus-NOR) A. Črne mar. littoral, Riisa, Trondheim Norway SLOCHN431 S MK159911 MK159969 - Marinogammarus marinus (Leach, 1815) TG 378Mmarinus (378-M-marinus-NOR) A. Črne mar. littoral, Korsvika, Trondheim Norway SLOCHN378 S MK159912 MK159970 MK160074 (Chaetogammarus obtusatus) l Macdonald et al. 2005 Novia Scotia Canada - S - AY926656 AY926761+ AY926823 - (Chaetogammarus pirloti) Englisch et al. 2003 Scotland UK - S - - AF419228 - genus Iberogammarus Iberian Peninsula Iberogammarus cf. macrocarpus 2 324Ealbent (324-E-Albent-ESP) C.Douady, F.Malard, C.Morvan Rio de Albentosa, Albentosa, Teruel Spain SLOCHN324 F MK159913 MK159971 MK160049 Iberogammarus macrocarpus 530Eulg (530-Eulg-mac-Tur-ESP) B.Sket R. Turia, Vilamarxant, Valencia river Spain SLOCHN530 F MK159914 MK159972 MK160050 Iberogammarus cf. macrocarpus 520Eulg (520-Eulg-mac-Ara-ESP) B.Sket S of Aranjuez Spain SLOCHN520 F MK159915 MK159973 MK160051 Iberogammarus sp. 325Egestal (325-E-Gestal-ESP) C.Douady, F.Malard, C.Morvan Fuente el Morenillo, Gestalgar, Valencia Spain SLOCHN325 F MK159916 - MK160075 Echinogammarus anisocheirus (Ruffo 1959) (Typ) TG 437Eanisoch (437-E.anisoc-T-FRA) B.Sket above St.Lary-de-Soulan hygropetric France-SW SLOCHN437 F MK159917 MK159999 MK160052 Echinogammarus anisocheirus (Ruffo 1959) (Typ) 149-E.anisoc-C62T-FRA B.Sket above St.Lary-de-Soulan hygropetric France-SW SLOCHN149 F KF478476 KF478563 KF478629 KF478658 11 genus Echinogammarus s. str. SW Europe Echinogammarus longisetosus 2 196-E-longs-ESP C.Douady, F.Malard, C.Morvan Tarragona, Creus, Santes, Rio Gaya-Santes-Creus interstitial Spain SLOCHN196 F KF478513 KF478598 - - Echinogammarus longisetosus 1 Pinkster, 1973 147-E.Nive-FRA B.Sket Uhart Cize, St-Jean-Pied-de-Port., r. Nive brook France-SW SLOCHN147 F KF478507 KF478593 - KF478662 Echinogammarus cf. berilloni few setae, pleon no setae 481Gae (481-Gae-Marfil-ESP) C.Douady, F.Malard, C.Morvan Fuente de Marfil-Valdenoceda, Burgos Spain SLOCHN481 F MK159918 MK160000 - 326Ebascons (326-E-Bascons-ESP) C.Douady, source Rio Rudron-Molino Echinogammarus cf longisetosus 10 F.Malard, C.Morvan Rasgabragas,Basconcillo, Castilla y Leon Spain SLOCHN326 F MK159919 MK160001 MK160053 Echinogammarus cf. calvus (Margalef, 1956) 106-E.cal-ESP G.Bračko Burgos, Quintana del Puente, r. Arlanza river Spain SLOCHN106 F KF478502 KF478588 KF478643 KF478660 Echinogammarus cf. tarragonensis Pinkster, 1973 195-E-targ-ESP C.Douady, F.Malard, Navarra, Nacedero, Larraun, c. Cueva Nacedero cave Spain SLOCHN195 F KF478512 KF478597 KF478650 C.Morvan de Larraun-Baraibar Echinogammarus cf. feminatus 533Eg (533-Eg-Toledo-ESP) B.Sket r. ?Tagus, 12 km E Toledo river Spain SLOCHN533 F MK159920 MK159974 MK160054 Echinogammarus cf. berilloni (less setose) 534Eg (534-Eg-Valenc-ESP) B.Sket Siete Aguas, Valencia spring Spain SLOCHN534 F MK159921 MK160002 MK160055 Echinogammarus feminatus Pinkster 1973 193-E-fem-ESP C.Douady, F.Malard, C.Morvan Oviedo, Fuesesce, Nava, s. Rio Fuente Santa Fuesenta interstitial Spain SLOCHN193 F KF478511 KF478596 KF478649 - Echinogammarus berilloni 3 (Catta, 1878) (Typ) TG 143-E.berill-T-FRA B.Sket Bidarray, Cambo-les-Bains, r. Nive river France SLOCHN143 F KF478497 KF478583 KF478639 - Echinogammarus berilloni 1 023-E.ber-Tour 023 B.Sket Tours, Chateau-Renault brook France SLOCHN023 F KF478501 KF478587 KF478642 - Echinogammarus berilloni 2 109-E.ber-FRA B.Sket Tours, Chateau-Renault brook France SLOCHN109 F KF478503 KF478589 KF478644 - Echinogammarus cf. berilloni 532EgBer (532-Eg-ber-Tou-FRA) B.Sket springs Touvre I, Angouleme, NE Bordeaux limnocrene springs France SLOCHN532 F MK159922 MK160003 MK160056 Echinogammarus cf. tarragonensis oo 525Eg (525-Eg-tarrag-FRA) B.Sket springs Touvre I, Angouleme, NE Bordeaux limnocrene springs France SLOCHN525 F MK159923 MK160004 MK160057 Echinogammarus sp. (def.) 536Eg (536-Eg-Touvre-FRA) B.Sket springs Touvre I, Angouleme, NE Bordeaux limnocrene springs France SLOCHN536 F MK159924 MK160005 MK160058 Echinogammarus cf. aquilifer Pinkster, 1969 192-E-aquil-ESP C.Douady, F.Malard, C.Morvan Santander, Tijeras, Ruiloba, c. Cueva de Tijeras-Concha cave Spain SLOCHN192 F KF478510 KF478595 - KF478663 Echinogammarus zebrinus Pinkster and Stock 1971 Typ 148-E.zebrin-TypeFRA B.Sket Uchacq-et-Parentis, Mont-de-Marsan, r. Estrigon brook France-SW SLOCHN148 F KF478508 KF478594 KF478648 - Echinogammarus zebrinus 1 Typ C148 (148-C-E-zebrinus-Type-FRA) B.Sket Uchacq-et-Parentis, Mont-de-Marsan, r. Estrigon brook France-SW SLOCHN148C F KF478509 - - - Typhlogammarus group genera Dinaric karst Typhlogammarus cf. mrazeki 252-Tyg-Krupa-HRV B.Jalžic Zrmanja, c. Krupa cave Croatia SLOCHN252 F KF478505 KF478591 KF478646 - Typhlogammarus heteropalpus. 502Tyg (502-Tyg-Hrnjak-HRV) T.Delic, M.Zagmajster Hrnjakova pecina, Bunic. cave Croatia SLOCHN502 F MK159925 MK160006 MK160059 Typhlogammarus heteropalpus G. Karaman, 1972(Typ) 436Typg (436-Typg-heterop-HRV-T) B.Jalžic Rudnica VI, Kamenica, Ogulin cave Croatia SLOCHN436 F MK159926 MK160007 MK160060 12 Typhlogammarus heteropalpus 282Trudn (282-T-Rudn-HRV) V.Jalžič Rudnica VI, Kamenica, Ogulin cave Croatia SLOCHN282 F MK159927 MK160008 MK160061 Typhlogammarus mrazeki 2 (Typ) TG 113-Typg-Typ-CrG P.Trontelj, S.Polak Cetinje, c. Lipska pečina cave Montenegro SLOCHN113 F KF478504 KF478590 KF478645 - Typhlogammarus mrazeki 1 Schaferna, 1922 020-T.mra-Vjet 020 V.Zakšek Popovo polje, Zavala, c. Vjetrenica cave Bosnia-Herzegovina SLOCHN020 F KF478500 KF478586 -- Accubogammarus sp. 114-?Accg-CrG P.Trontelj, S.Polak Grahovo, c. Vojvode Dakoviča pečina cave Montenegro SLOCHN114 F KF478506 KF478592 KF478647 KF478661 Metohia carinatal (Absolon, 1927) TG 019-M.car-übod 019 P.Trontelj, S.Polak Rijeka Crnojeviča, c. Obodska pečina cave stream Montenegro SLOCHN019 F KF478498 KF478584 KF478640 - Metohia carinata 2 025-M.car-Čič 025 B.Sket Čičevo, Velja gora, c. Šumet cave Bosnia-Herzegovina SLOCHN025 F KF478499 KF478585 KF478641 - genus Parhomoeogammarus W Iberian Peninsula Parhomoeogammarus pacaudi (Hubault & Ruffo, 1956) Typ 522Eg (522-Eg-pacaudi-FRA) B.Sket Bastarisse-Bazas, SE Bordeaux forest brook France SLOCHN522 F MK159928 MK159975 MK160062 Echinogammarus pacaudi (no setae, no denticles) 521Eg (521-Eg-Bazas-FRA) B.Sket Bastarisse-Bazas, SE Bordeaux forest brook France SLOCHN521 F MK159929 MK159976 MK160063 Parhomoeogammarus lusitanus (Typ) TG 526Prhmg (526-Prhmg-lus-T-PüR) B.Sket Formariz PdC, Paredes de Coura spring Portugal SLOCHN526 F MK159930 MK159977 MK160064 Echinogammarus 477Phg (477-Phg-lusit-PüR) (Parhomoeog) lusitanus l) (no calceoli) E.Martinez Ansemil A Coruna prov., Galicia Spain SLOCHN477 F MK159931 MK159978 MK160065 genus Relictogammarus E Atlantic Marinogammarus stoerensis (Reid, 1938) TG 250-Mg-Cork-ATL A.Myers County Cork, Fountainstown sea, beach Ireland-Atlantic SLOCHN250 S KF478515 KF478601 KF478651 - Chaetogammarus stoerensis (Reid, 1938) Macdonald et al. 2005 Maine USA - S - AY926657 AY926762+ AY926824 - Gammarus group genera (including Bajkalian genera; 22 OTUs) Gammarus + Zenkevitchia 14 + 3 OTUs Holarctic Bajkal genera 5 OTUs Bajkal Sarothrogammarus group genera (incl. ?Tadzocrangonyx; 16 OTUs) 6 OTUs Central Asia high mount. 10 OTUs Mediterranean & E Atlantic coastal genus Relictogammarus E Atlantic Marinogammarus stoerensis (Reid, 1938) TG 250-Mg-Cork-ATL A.Myers County Cork, Fountainstown sea, beach Ireland-Atlantic SLOCHN250 S KF478515 KF478601 KF478651 - (Chaetogammarus stoerensis (Reid, 1938)) Macdonald et al. 2005 Maine USA - S - AY926657 AY926762+ AY926824 - Outgroup 16 species, 8 genera, 5 families. fam. Gammaracanthidae 5 OTUs fam. Crangonyctidae 2 OTUs fam. Niphargidae 2 OTUs fam. Anisogammaridae 5 OTUs fam. Talitridae 2 OTUs 13