The Mysidae (Crustacea: Peracarida: Mysida) in fresh and oligohaline waters of the Mediterranean. Taxonomy, biogeography, and bioinvasion

Karl J. Wittmann; Mikhail Daneliya

Among a total of nine species of Mysidae so far found in fresh waters of the Mediterranean, four are stenoendemics, namely of a single lake (Diamysis lacustris), of a single stream and its springs (Diamysis hebraica), waters of a single river basin (Paramysis kosswigi) or of karstic cave waters (Troglomysis vjetrenicensis), respectively. Not less than four species, T. vjetrenicensis, D. lacustris, D. fluviatilis, and P. adriatica nov. sp., are confined to freshwater tributaries of the Adriatic Sea (NE-Mediterranean). This strengthens previous findings about the outstanding role of the Adriatic basin for the endemic diversity of freshwater Mysidae within the Mediterranean; possibly related to alternating marine and freshwater-terrestrial phases during Pliocene-Pleistocene in this semi-enclosed basin. As far as known, freshwater populations of D. mesohalobia heterandra are also confined to the Adriatic basin; however this taxon shows many more populations in brackish waters of the E-Mediterranean and Marmora basins. The remaining two freshwater species (Limnomysis benedeni, Hemimysis anomala) are wide-range invaders of Ponto-Caspian origin, with recent expansion to fresh and brackish waters of the NW-Mediterranean. Two additional species, Neomysis integer and Mesopodopsis slabberi, are known from a few fresh waters and many more brackish and marine waters of the NE-Atlantic or Black Sea basins, respectively; in the Mediterranean, however, so far only from saline waters down to almost fresh water (S >= 0.9). A well figured key to the species is given, including certain potential future invaders.

Zootaxa 4142 (1): 001–070 http://www.mapress.com/j/zt/ Copyright © 2016 Magnolia Press ISSN 1175-5326 (print edition) Monograph ZOOTAXA ISSN 1175-5334 (online edition) http://doi.org/10.11646/zootaxa.4142.1.1 http://zoobank.org/urn:lsid:zoobank.org:pub:FA423164-276C-44B0-A417-8E97AC3DF0AA ZOOTAXA 4142 The Mysidae (Crustacea: Peracarida: Mysida) in fresh and oligohaline waters of the Mediterranean. Taxonomy, biogeography, and bioinvasion KARL J. WITTMANN1, ANTONIO P. ARIANI 2 & MIKHAIL DANELIYA3 1 Institut für Umwelthygiene, Zentrum für Public Health, Medizinische Universität Wien, Kinderspitalgasse 15, A-1090 Vienna, Austria. E-mail: karl.wittmann@meduniwien.ac.at 2 Dipartimento delle Scienze Biologiche, Sezione di Zoologia, Università di Napoli Federico II, Via Mezzocannone 8, I-80134 Naples, Italy. E-mail: antonio.ariani@gmail.com 3 Department of Biosciences, POB 65, 00014, University of Helsinki, Finland; Taxonomicum, Sydäntie 8 E 16, FI-01400, Vantaa, Finland. E-mail: mikhail.daneliya@taxonomicum.com Magnolia Press Auckland, New Zealand Accepted by K. Meland: 10 Jun. 2016; published: 25 Jul. 2016 KARL J. WITTMANN, ANTONIO P. ARIANI & MIKHAIL DANELIYA The Mysidae (Crustacea: Peracarida: Mysida) in fresh and oligohaline waters of the Mediterranean. Taxonomy, biogeography, and bioinvasion (Zootaxa 4142) 70 pp.; 30 cm. 25 Jul. 2016 ISBN 978-1-77557-989-2 (paperback) ISBN 978-1-77557-990-8 (Online edition) FIRST PUBLISHED IN 2016 BY Magnolia Press P.O. Box 41-383 Auckland 1346 New Zealand e-mail: magnolia@mapress.com http://www.mapress.com/j/zt © 2016 Magnolia Press All rights reserved. No part of this publication may be reproduced, stored, transmitted or disseminated, in any form, or by any means, without prior written permission from the publisher, to whom all requests to reproduce copyright material should be directed in writing. This authorization does not extend to any other kind of copying, by any means, in any form, and for any purpose other than private research use. ISSN 1175-5326 (Print edition) ISSN 1175-5334 (Online edition) 2 · Zootaxa 4142 (1) © 2016 Magnolia Press WITTMANN ET AL. Table of contents Abstract . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 Material and methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 Systematics . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 Family Mysidae Haworth, 1825 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 Subfamily Mysinae Haworth, 1825 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 Tribe Mysini Haworth, 1825 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 Genus Mesopodopsis Czerniavsky, 1882 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 Mesopodopsis slabberi (Van Beneden, 1861) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 Hemimysini Czerniavsky, 1882, revalidated at tribe rank . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10 Genus Hemimysis G. O. Sars, 1869 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11 Hemimysis anomala G. O. Sars, 1907 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11 Paramysini Wittmann, Ariani & Daneliya, new tribe . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13 Genus Paramysis Czerniavsky, 1882 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14 Subgenus Serrapalpisis Daneliya, 2004 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14 Paramysis (Serrapalpisis) kosswigi Băcescu, 1948 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15 Subgenus Longidentia Daneliya, 2004 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18 Paramysis (Longidentia) adriatica sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19 Tribe Diamysini Wittmann, Ariani & Lagardère, 2014 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27 Genus Diamysis Czerniavsky, 1882 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27 Diamysis fluviatilis Wittmann & Ariani, 2012 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28 Diamysis lacustris Băcescu, 1940. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28 Diamysis mesohalobia Ariani & Wittmann, 2000 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29 Diamysis mesohalobia mesohalobia Ariani & Wittmann, 2000 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32 Diamysis mesohalobia gracilipes Ariani & Wittmann, 2000 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32 Diamysis mesohalobia heterandra Ariani & Wittmann, 2000 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33 Diamysis lagunaris Ariani & Wittmann, 2000 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36 Diamysis hebraica Almeida Prado-Por, 1981 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38 Genus Troglomysis Stammer, 1933 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39 Troglomysis vjetrenicensis Stammer, 1933 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39 Genus Limnomysis Czerniavsky, 1882 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41 Limnomysis benedeni Czerniavsky, 1882 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41 Tribe Neomysini Wittmann, Ariani & Lagardère, 2014 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 44 Genus Neomysis Czerniavsky, 1882 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 44 Neomysis integer (Leach, 1814) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 44 Discussion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 47 Key to the tribes of the subfamily Mysinae Haworth, 1825 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 54 Key to the species of Mysidae in fresh and oligohaline waters of the Mediterranean . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 54 Acknowledgements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 57 References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 57 Abstract A census of Mysidae yielded a total of twelve species plus two non-nominotypical subspecies found so far in fresh and oligohaline waters of the Mediterranean, all belonging to the subfamily Mysinae. Among the nine species in fresh-waters, three are stenoendemics, namely of a single lake (Diamysis lacustris), of two neighbouring river systems (Paramysis kosswigi) or of karstic cave waters (Troglomysis vjetrenicensis). Four species—T. vjetrenicensis, D. lacustris, D. fluviatilis, and Paramysis adriatica sp. nov. described in this paper—are confined to freshwater tributaries of the Adriatic Sea (NEMediterranean). This strengthens previous findings about the outstanding role of the Adriatic basin for the endemic diversity of freshwater Mysidae within the Mediterranean. This is possibly related to alternating marine and freshwater-terrestrial phases during the Pliocene-Pleistocene in this semi-enclosed basin. Based on current knowledge, freshwater populations of D. mesohalobia heterandra are also confined to the Adriatic basin; this taxon, however, shows many more populations in brackish waters of the E-Mediterranean and Marmora basins. Two freshwater species (Limnomysis benedeni, Hemimysis anomala) are wide-range invaders of Ponto-Caspian origin, with recent expansion into fresh and brackish waters of the NW-Mediterranean. A further immigrant to this part of the Mediterranean, Neomysis integer, is of NE-Atlantic origin and occurs only marginally in fresh-water. MYSIDAE IN MEDITERRANEAN FRESH-WATERS Zootaxa 4142 (1) © 2016 Magnolia Press · 3 Five among the nine species found at least once in fresh-water were also reported in oligohaline conditions, mostly also at even higher salinities. A total of eight species plus two subspecies were recorded in oligohaline waters (S = 0.5– 5). Among these, only one oligohalobious stenoendemic, Diamysis hebraica, inhabits streams at the Levantine coast. In the historical biogeographical context, the current distribution of only one out of nine indigenous species in an- to oligohaline waters of the Mediterranean, namely the mainly meso- to polyhalobious Mesopodopsis slabberi, may date back to the early Pliocene flooding of the Mediterranean by Atlantic waters and to later events. For most species, the biogeographical pattern points to a primary origin in the brackish (Miocene) Paratethys; solely the cave-dwelling T. vjetrenicensis has more ancient roots in the Tethyan (Mesogeic) Sea. Both these hypotheses are supported by chorological data and mainly the mineral composition of statoliths. The statoliths are composed of CaCO3 as the metastable crystal phase vaterite in nine species plus two subspecies considered versus the otherwise more common CaF2 (fluorite) in only three species (or in 7 + 2 versus two Mediterranean indigenes). All 12 + 2 Mediterranean taxa are figured and described in detail, particularly regarding P. adriatica sp. nov. and the substantially redescribed T. vjetrenicensis Stammer, 1933. Supplementary descriptions are given for P. kosswigi Băcescu, 1948, D. hebraica Almeida Prado-Por, 1981, and N. integer (Leach, 1814). A key to the 14 taxa is given including additional three species of potential future invaders. The tribe Mysini Haworth, 1825, is revised by detachment of the newly defined tribe Paramysini and of the revalidated Hemimysini Czerniavsky, 1882. A key to the resulting six tribes of the subfamily Mysinae is given. Key words: faunistics, fresh-water, indigenous species, neozoa, historical biogeography, morphology, revised description, new species, new tribe, Adriatic basin Introduction In a large biogeographical assessment of the order Mysida (including the Stygiomysida), Porter et al. (2008) estimated that about 7% of mysid species diversity occurs in fresh-water. The families Lepidomysidae, Stygiomysidae, and Mysidae are represented by freshwater populations in Palaearctic and Neotropical regions. Porter et al. (2008) distinguished four main groups according to different history and routes of freshwater immigration: (1) Subterranean Tethyan relicts; (2) Autochthonous Ponto-Caspian endemics; (3) Mysis spp. (subarctic immigrants as 'Glacial Relicts'); and (4) Euryhaline estuarine species. Among the species of Mysidae, only Troglomysis vjetrenicensis Stammer, 1933, and Paramysis kosswigi Băcescu, 1948, were known at that time as indigenous freshwater inhabitants of the Mediterranean. Meanwhile, Diamysis lacustris Băcescu, 1940, and D. fluviatilis Wittmann & Ariani, 2012, were added to this group by Wittmann & Ariani (2012b), and Paramysis adriatica sp. nov. is appended below. The increasing species numbers indicate that it is promising to investigate and discuss below a potential autochthonous freshwater immigration by mysids in the Mediterranean. From a different biogeographical point of view, Fukuoka (2006) distinguished ten regional groups of Mysini genera (Mysinae in current taxonomy—Meland et al. 2015) with main distribution in tropical to boreal latitudes on a world-wide scale; these groups are largely associated with sea basins, continents or subcontinents. For high latitudes, Fukuoka distinguished the circumantarctic Antarctomysis Coutière, 1906, from the circumarctic genera Mysis Latreille, 1802, and Stilomysis Norman, 1892. This distinction is less clear for Neomysis Czerniavsky, 1882, with mainly boreal, (sub)arctic, and notoboreal distribution (in our terminology). Like all indigenous Mysini (i.e. Mysinae) from the Mediterranean, the an- to oligohalobious taxa treated below belong either to Fukuoka's group 1 (European) or to group 10 (African). This omits Neomysis integer (Leach, 1814), which is most likely a recent immigrant from the NE-Atlantic, as discussed below. More recent biogeographical treatments of North Eurasian continental waters and the Ponto-Caspian region (Daneliya & Petryashev 2011, Daneliya et al. 2012) consider the entire freshwater mysid fauna of the Asia Minor peninsula to be derivates of the Ponto-Caspian fauna. This includes the Ponto-Caspian Limnomysis benedeni Czerniavsky, 1882, the autochthonous Paramysis lacustris turcica (Băcescu, 1948), the latter representing a subspecies of the Ponto-Caspian Paramysis lacustris (Czerniavsky, 1882), the freshwater Pontian Diamysis pengoi Czerniavsky, 1882, and the autochthonous Anatolian P. kosswigi, which is morphologically closely related to P. lacustris. The endemic P. kosswigi and P. l. turcica are certainly indicative of some period of isolated history and thus were assigned by Daneliya & Petryashev (2011) to a separate Anatolian Province of the Ponto-Caspian Region. Not counting populations in recently invaded waters, L. benedeni, D. pengoi and P. l. turcica inhabit waters draining into the Marmora and/or Black Sea; only P. kosswigi inhabits tributary systems of the 4 · Zootaxa 4142 (1) © 2016 Magnolia Press WITTMANN ET AL. Mediterranean. It remains obscure at which time scale and by which means the ancestors of P. kosswigi entered the Mediterranean drainage. Another centre of local endemism is reported (Ruffo 1957, Ariani 1981c, Pesce 1985, Inguscio et al. 1999, Wittmann et al. 2014) from brackish and fresh subterranean waters of Apulia, i.e. in the south-east of the Apennine Peninsula (central Mediterranean). Among these subterranean endemics, Stygiomysis hydruntina Caroli, 1937, and Spelaeomysis bottazzii, Caroli, 1924, belong to the order Stygiomysida Tchindonova, 1981. Since the Mysidacea Boas, 1883, were split into three orders by Meland & Willassen (2007), the Stygiomysida are no longer pooled together with the order Mysida Boas, 1883, and are therefore not treated in the present contribution. Besides N. integer, additional species are considered to represent immigrants or, more accurately, invaders of Mediterranean coastal waters: Hemimysis anomala G. O. Sars, 1907, and L. benedeni are found on the Mediterranean coast of France (Wittmann & Ariani 2009, Wittmann et al. 2014). Both species are of PontoCaspian origin and have already invaded vast areas of central, western and northern Europe, H. anomala in addition the Great Lakes of North America. Both may have negative impacts by grazing on indigenous zooplankton communities (Ketelaars et al. 1999, Fink et al. 2012). Nonetheless, Aßmann et al. (2009) expect partly positive effects from L. benedeni invasions due to its role in leaf litter decomposition. A number of unpublished records is given below for both species from the Rhône system and additional tributaries of the Mediterranean. In a review of the Mysida and additional orders, Wittmann et al. (2014) complained about the residual heterogeneity in the tribe Mysini Haworth, 1825. This problem is reduced below by splitting off two tribes based on gross-morphological differences. Nonetheless, one major difficulty already discussed by W. M. Tattersall (1927), i.e. the questionable allocation of the genus Kainommatomysis W. M. Tattersall, 1927, remains a challenge for future investigation. Material and methods The terminology and methods used here are given in Wittmann et al. (1993, 2014), Ariani & Wittmann (2000), and Wittmann & Ariani (2012a, 2012b). In material lists, sexes are abbreviated as 'F' or 'M'. Stages in the marsupium are eggs or embryos until egg hatching, nauplioids until first larval moult and postnauplioids until second larval moult; the latter occurs shortly before up to shortly after release from the brood pouch. Four postmarsupial stages are distinguished: juveniles (juv.) characterized by the absence of secondary sexual characteristics; immatures (imm.) by rudimentary secondary sexual characteristics; subadults (subad.) by advanced but incomplete ones, i.e. a small marsupium in females, a near-adult formation of antennae and pleopods in both sexes, absence of mature sperm in males; fully developed adults (ad.) as in Fig. 7. For reasons of brevity, 'tarsus' is often used for the ensemble of carpus, propodus, and dactylus. For most taxa we distinguish a carpopropodus from the small, clawbearing dactylus (Figs 1E, 3E). By contrast, for the genus Paramysis we follow Băcescu (1940, 1954), who distinguished a stout, one-segmented carpus with specialized setation pattern, from a more uniformly developed and subsegmented propodus (Figs 4H, 10A, D, F, H), and of course also a dactylus. 'Fenestra paracornealis' is a pale, pigment-free spot (Figs 13E, 17B) dorsally on the eyestalks near the cornea (Ariani & Wittmann 2000). After bleaching a ganglion mass becomes visible below the fenestra paracornealis; the latter resembles the organ of Bellonci in size and structure. 'Paradactylar setae' are often somewhat modified setae (Figs 4J, K, 10B, C, E, G, H) at the terminal end of the propodus, flanking the dactylus in thoracopods 3-8. 'Scutellum paracaudale' is used for the laterodorsal protrusions (Figs 1H, J, 15F-H, R, Q) of the sixth pleonite, flanking the telson (Ariani & Wittmann 2000). Body size was measured from the tip of the rostrum to the posterior end of the telson, without spines. The relation of length to maximum width of the merus in thoracic endopod 6 is indicated as R6 (Ariani & Wittmann 2000, 2002). The mineral composition of statoliths was determined with a combination of optical and chemical examinations according to Wittmann et al. (1993). Most materials were collected in 1974–2012 during own excursions to fresh, brackish, and marine waters of the Mediterranean and adjacent seas. Geographical coordinates were derived from touristic and navigation maps in 1974–1996, and in part recently corrected using satellite images provided by Google Earth®; or directly measured with satellite telemetry (GPS) in 1997–2012. Distance of sampling positions from the sea was estimated as distance along waterways, including meanders. Abiotic parameters were measured in 0.5 m depth. In 1982–1996, salinity MYSIDAE IN MEDITERRANEAN FRESH-WATERS Zootaxa 4142 (1) © 2016 Magnolia Press · 5 was measured with a hand refractometer ‘Atago S/Mill’, zero calibrated before each measurement. In 1997–2012, a ‘Multiparameter Water Quality Checker Horiba U-10’ was used to measure turbidity (in NTU = nephelometric turbidity units), O2, pH, temperature, and salinity. Salinity (S) is expressed as a dimensionless equivalent of electric conductivity (the latter in µS/cm). The Venice System was used for salinity classification of water bodies (Por 1972). Test stripes 'Aquadur' were used to measure mineral content as German Degrees (°d, also known as °dH) of water hardness. Surface water flow (v) was measured with drift objects. Types and reference material were deposited and/or taken on loan, respectively, from the Naturhistorisches Museum Wien (NHMW), the Museo Zoologico, Università di Napoli Federico II (MZUN), the Museo Regionale di Scienze Naturali, Torino (MZUT), Muséum d'Histoire Naturelle de la Ville de Genève (MHNG), Finnish Museum of Natural History, Helsinki (MZH), Zoological Collection, Hebrew University of Jerusalem (HUJ), NCB Naturalis, Leiden (RNMH), Swedish Museum of Natural History, Stockholm (SMNH), British Museum of Natural History, London (BMNH), and the Grigore Antipa National Museum of Natural History, Bucharest (MGAB). Additional materials were kindly provided by Boris Sket (Biologija Univerza Ljubljana—BUL) and Cem Aygen (Ege University, Izmir, Turkey—EGE). Materials collected by Fabio Stoch (Trieste) and his colleagues were deposited in the Museo Friulano di Storia Naturale, Udine (MFSNU). Systematics Family Mysidae Haworth, 1825 Subfamily Mysinae Haworth, 1825 Tribe Mysini Haworth, 1825 Fig. 1 Diagnosis (see also ‘Discussion’). Mysinae with antennal scale setose all around, scale in most species with small apical segment; carpopropodus of the thoracic endopod 6 with 3–26 segments; well-developed oostegites on thoracopods 7, 8, and rudimentary oostegite on thoracopod 6; all pleopods of females and pleopods 1, 2 of males rudimentary; male pleopod 3 mostly biramous, but uniramous, vestigial in Kainommatomysis, if biramous then with 1- to 15-segmented endopod and 2- to 16-segmented exopod; male pleopod 4 biramous, its endopod 1- to 14segmented, its exopod 3- to 29-segmented, terminally with modified seta(e); male pleopod 5 shorter, biramous or uniramous; statoliths (as far as known) composed of fluorite; telson variable, mostly with apical cleft, this cleft (if present) lined by spine-like laminae, telson of several genera with a pair of medio-apical plumose setae emerging from the cleft. Type genus. Mysis Latreille, 1802. Taxa included. Ten genera provisionally listed (see ‘Discussion’): Antarctomysis Coutière, 1906 (3 species), Arthromysis Colosi, 1924 (1), Hyperstilomysis Fukuoka, Bravo & Murano, 2005 (1), Kainommatomysis W. M. Tattersall, 1927 (3), Mesopodopsis Czerniavsky, 1882 (8), Mysis Latreille, 1802 (15), Nanomysis W. M. Tattersall, 1921 (3), Parastilomysis Ii, 1936 (4), Stilomysis Norman, 1892 (4; this genus defined in key by Norman, 1892: 148), Tasmanomysis Fenton, 1985 (1). Genus Mesopodopsis Czerniavsky, 1882 Mesopodopsis slabberi (Van Beneden, 1861) Fig. 1 Short selection from 39 synonymy statements with a total of 425 references: Podopsis Slabberi Van Beneden, 1861: Marcusen 1867; Marion 1894; Masi 1906; Hess 1910. Mysis Slabberi: Goës 1864. Podopsis pontica Czerniavsky, 1870: nomen nudum (in synonymy lists often cited as 1869). Macropsis Slabberi: Sars 1877; Carus 1885; Gourret 1894; Sudry 1910; Percival 1929 (partim: 102). 6 · Zootaxa 4142 (1) © 2016 Magnolia Press WITTMANN ET AL. Podopsis (Mesopodopsis) Slabberi: Czerniavsky 1882a, 1887. Podopsis (Parapodopsis) Goësi Czerniavsky, 1882a. Podopsis (Parapodopsis) cornuta Czerniavsky, 1882a, 1887. Podopsis (Parapodopsis) Goesi: Czerniavsky 1887. Parapodopsis cornuta: Butchinsky 1885, 1890; Sowinsky 1894; Retzius 1910; Knipowitsch 1925. Parapodopsis cornutum: Kowalevsky 1889; Gaskell 1908. Macropsis slabberi: Scott 1888; Graeffe 1902; Zimmer 1915b; Mazoué 1931; Nekrasova & Rakitina 1968. Leptocaris Slabberi: Aurivillius 1898a, 1898b. Mesopodopsis Slabberi: Norman & Scott 1906; Norman 1907; Colosi 1929; Fage 1933; Nouvel 1943. Podopsis slabberi: Van der Sleen 1920; Gauthier 1928; Stammer 1932 (partim: 562). Mesopodopsis slabberi: W. M. Tattersall 1922, 1927 (partim: Port Said); Colosi 1929; Băcescu 1941; Ariani 1967; Daneliya 2002; Kocataş et al. 2003; San Vicente 2010; Wittmann et al. 2014. Material examined (southern France, hand net, leg. K. J. Wittmann, if not stated otherwise; among 85 Mediterranean samples inspected, only those from salinity S < 5 are listed). 27 F ad. 7.5–9.5 mm, 97 M ad. 6.5–9.6 mm, 106 F subad., 30 M subad., 58 imm., 16 juv. (additional ~30,000 specimens sorted only by taxon), accompanied by 2 Limnomysis benedeni and 3 Diamysis lagunaris, estuary of the Petit Rhône at Tiki, near Rhônekm 337.5; southern bank, 43.4514N 004.3976E, sea distance 180 m, 0.2–1.2 m depth, taken from filiform green algae on boulders, v = 0 m/s, S = 2.2, 4080 µS/cm, 23.7°C, pH 7.71, 7.50 mg O2/l, 14 NTU, 19 June 2009, NHMW reg. no. 25699; 1 F ad. 11.1 mm, Canal d'Arles à Fos, at corner with the canal Liaison Rhône - Fos, 43.4663N 004.8338E, sea distance 7 km, altitude 0 m, 0.3–1.5 m depth, from boulders and concrete walls with filiform algae, and from mud, v = 0 m/s, S = 3.4, 6490 µS/cm, 23.1°C, pH 7.42, 5.75 mg O2/l, 43 NTU, 17 June 2009; 37 F ad. 7.3–9.2 mm, 1 M ad. 6.8 mm, 28 F subad., 13 M subad., 335 imm., 1312 juv., Départment Gard, Canal du Rhône à Sète, at canal-km PK 48 (= K.21), 43.5880N 004.2168E, sea distance 9 km, altitude 0 m, 0.3–1.5 m depth, from bank vegetation, filiform algae, and stones, v = 0 m/s, S = 3.4, 6370 µS/cm, 25.9°C, pH 7.18, 3.29 mg O2/l, 9°d, 66 NTU, 15 June 2007, NHMW 25701; 76 F ad. 11.4–14.5 mm, 3 M ad. 8.6–11.6 mm, 1 F subad., 2 M subad., North Adriatic drainage, Italy, delta of Po River, mouth branch Po di Goro, 44.7960N 012.3915E, sea distance 800 m, 0.5–1.2 m depth, from shoots and roots of Phragmites, and from wood, v = 0.05–0.2 m/s, S = 1.4, 2880 µS/cm, 14.2°C, pH 7.67, 9.21 mg O2/l, 28 NTU, 14 Apr. 2012, leg. Ariani & Wittmann, NHMW 25700. Description (Fig. 1; Mediterranean materials only, adult females 5–15 mm, males 4–12 mm). Mysini with eyes well developed, cornea globular; eyestalks smooth, cylindrical, 3–4 times as long as cornea (Fig. 1A). Peduncle of antennula much more stout in males (Fig. 1A) than in females, seta at antero-lateral corner of basal segment in males much shorter than in females; male antennula with accessory flagellum (Fig. 1A) having 3–4 small setae curled with their slender distal portions around the basis of the large, straight, smooth, terminal seta; this flagellum and the appendix masculina extend to about same length when stretched anteriorly, not taking account of setae. Antennal segment posteriorly elongate giving the cephalic region its remarkably slender appearance. Antennal scale length subequal to antennular peduncle; scale slender, terminally narrowing, bluntly ending; inner and outer margins densely setose, without spines; apical segment is 13–19% scale length, this segment with 5 plumose setae (Fig. 1A). Rostrum short, subtriangular with wide angle, anteriorly well rounded (Fig. 1A). Lacinia mobilis of right mandible with 2 large and 4–9 small teeth; median segment of mandibular palp with setae over the distal 62–93% of its length. Carapace smooth (Fig. 1A, B), antero-laterally with a pair of spines; carapace with clear cervical constriction, nonetheless cervical sulcus weak, visible only in situ (Fig. 1A), but not so in detached material (Fig. 1B); total of 20–30 cardial pores in a double wing-like arrangement (Fig. 1B, C) as typical for the genus, absence of cervical pores. Thoracic sternite 1 medially with dense fields of minute hairs, sternite 2 much less hairy, sternites 3–8 medially smooth (Fig. 1D); in accordance with the slender body shape of the animals, the sternites are narrow, laterally delimited by holdfasts for the coxae of the thoracopods; in females the distance between these holdfasts increases from sternite 4 to sternite 8 (Fig. 1D), thus providing additional space for the marsupium and its content. First thoracic endopod with a large, setose endite from the basis plus a small, often indistinct, endite from the coxa; its epipod elongate, tongue-like, terminally rounded, without seta (Fig. 1D); dactylus of endopods 1, 2 setose; claw missing in endopods 1–8; tarsus of endopods 3–7 with 5–8 segments, tarsus of endopod 8 with 4–7 segments; basal to penultimate segments of tarsus 3 each with modified seta at outer distal border, this seta basally barbed and distally armed with minute spines (Fig. 1E)—such a seta only on penultimate segment of tarsus 8, yet with the spines larger and more recurved, no such setae on tarsi 4–7; these last tarsi with basally barbed setae (without MYSIDAE IN MEDITERRANEAN FRESH-WATERS Zootaxa 4142 (1) © 2016 Magnolia Press · 7 FIGURE 1. Mesopodopsis slabberi (Van Beneden, 1861) from the oligohaline reach of the Po di Goro, a mouth branch of the Po River, males with 8.6 mm (A) or 10.7 mm (F–H, L), and female with 13.5 mm (B–E, J, K) body length. A, anterior body region of adult male, dorsal view; B, carapace expanded on slide, dorsal; C, detail of (B) showing cardial pore group, dorsal; D, thoracic sternites (ventral) with caudal faces of left thoracopods 1 and 6; E, 'tarsus' of third thoracopod, rostral face; F, male pleopod 3, rostral; G, male pleopod 4, caudal; H, J, posterior margin of sixth pleonite in male (H) and female (J), lateral; K, uropods, ventral; L, telson, dorsal. 8 · Zootaxa 4142 (1) © 2016 Magnolia Press WITTMANN ET AL. spines) on their basal segments but not on their terminal 2–4 segments. Basis of all thoracic exopods with well rounded outer corner; flagellum 8-segmented in exopods 1 and 8, whereas 9-segmented in exopods 2–7, not counting the large intersegmental joint between basis and flagellum which may be mistaken as a segment. Large marsupial plates on thoracopods 7, 8; the smaller first oostegites with posterior lobe bearing a number of long, backwards directed setae that are spinose on their distal half; a number of such setae but no such lobe on the large second oostegites; a posterior lobe with only 2 (1–3) such setae present (Fig. 1D) also on the sympod of thoracopod 6 in females only. Penes normal, large, each with semicircle of only 4–5 smooth, weakly curved setae anteriorly close to the ejaculatory opening; brush of 3–7 large, barbed setae on the outer face at 31–45% penis length from tip; inner margin with area of acute (in part hair-like) scales. All pleopods of females and pleopods 1, 2, 5 of males reduced to undivided endopods with indistinct outer apophysis; male pleopods 3, 4 (Fig. 1F, G) each with large, 2-segmented sympod, terminal segment of each sympod with area of scales similar in arrangement and relative position to that on the penis; endopod of male pleopods 3, 4 with distinct, setose apophysis directed outwards in subbasal position; male pleopod 3 (Fig. 1F) with comparatively large, unsegmented endopod, and with clearly shorter, somewhat reduced, 2- to 3-segmented exopod; pleopod 4 (Fig. 1G) with minute, 2- to 3-segmented endopod and with long, 3-segmented exopod, the latter showing a large modified seta plus a much shorter one on its short terminal segment (Fig. 1G). On each side of the pleon, the scutellum paracaudale represents a roughly triangular plate with acute tip and undulate upper and lower margins (Fig. 1H, J); the scutellum is forward displaced from the posterior margin of the sixth pleonite by slightly less than its own length. Endopod of uropods with blunt projection above statocyst and with one spine below statocyst (Fig. 1K). Statoliths composed of fluorite. Telson shorter than last abdominal somite, hirsute shortly behind basal corners, terminally ending with two lateral and one median lobe (Fig. 1L), the latter 3–5 times the length of the lateral lobes; only distal 28–60% of lateral margins armed with 3–8 spines, not counting the larger spine at the tip of each lateral lobe; margin of medio-terminal lobe with 20–36 densely set spines (spine-like laminae). Distribution (Fig. 2). This mysid is known from coastal marine and brackish waters of the Ponto-Azov, Mediterranean and Baltic Seas, and of the NE-Atlantic from Norway to Morocco (59°N-33°N). It occurs in anhaline to hyperhaline conditions, with main occurrence in meso- to polyhaline waters, very rarely found in freshwater. Nonetheless it may be abundant also in coastal marine environments (Mees et al. 1993). According to Vilas et al. (2006) it shows strong osmoregulatory capability in the range of S = 7–29, with elevated oxygen consumption already at S = 6. Brood pouch young tolerate only more narrow ranges of salinity compared to adults (Greenwood et al. 1989). Brun (1967) reported Mesopodopsis slabberi from the chlorinity range of 0.5–21‰ (salinity 0.9–38) in the estuary of the Grand Rhône (NW-Mediterranean). For records made by Aguesse & Bigot (1960) in the Rhône delta see below, chapter on Neomysis integer. The above-listed own samples of M. slabberi from the deltas of the Rhône and Po Rivers are in the salinity range of 1.4–3.4. Pesta (1935) reported this species from S = 1.3 in a brackish drain at the Island of Corfu (NE-Mediterranean). None out of 85 Mediterranean samples inspected by us were below this value (S = 1.4–43; 1974–2012). In the Tamar River estuary at the E-Atlantic coast of SW-England, M. slabberi shows a less wide salinity distribution (S > 5) compared with the co-occurring N. integer; there is differential distribution of age stages versus salinity, and the population shows a down-estuary movement in winter (Moffat & Jones 1993, Moffat 1996). In accordance with this, M. slabberi tends to disappear in winter when salinity decreases to values below S = 5 and water temperature below 10°C in a warm-temperate estuary at the coast of Portugal (Azeiteiro et al. 1999). Its abundance along the Westerschelde estuary at the Dutch-Belgian border is primarily correlated with temperature (Rappé et al. 2011), whereas primarily with salinity in the Guadalquivir (SW-Spain; Fernández-Delgado et al. 2007) and Gironde (W-France; Castel 1993, David et al. 2005) estuaries. This species appears to be also rare in fresh-waters draining into the Black Sea. Dedju & Polischtuk (1968) listed it for fresh and brackish, near-deltaic waters of the Danube River, with main occurrence in the "polyhaline" (= probably 'mesohaline' according to the Venice System) reach. Similarly, Băcescu & Dumitrescu (1958) found masses in oligo- to mesohaline waters off Danube mouth branches. Upon our own sampling in Black Sea coastal waters, thousands of M. slabberi were found in oligo- to mesohaline coastal waters, but only one adult female in true fresh-water (NW-Black Sea, Danube Delta, Canal Tătaru, 45.0765N 029.6342E, sea distance 1.4 km, 0.3–4.7 m depth, S = 0.0–0.1, 376 µS/cm, snorkel diver operated net, 24 June 2008, leg. K. J. Wittmann). At that coast this mysid species has been found for almost a century (Borcea 1926; Bâcescu 1940, 1954; Teodorescu-Leonte 1977; MYSIDAE IN MEDITERRANEAN FRESH-WATERS Zootaxa 4142 (1) © 2016 Magnolia Press · 9 Begun & Gomoiu 2007) in the oligohaline waters of the perimarine Lake Sinoe (Danube Predelta). Begun & Gomoiu (2007) demonstrated great faunal changes with respect to the 1950–70s, and attributed this to freshwater input enhanced by humans. The mysids Mesopodopsis slabberi and Paramysis kroyeri (Czerniavsky, 1882) now tend to disappear during episodic freshening of the lake water, and inversely to recolonize this lake upon periods of seawater intrusion through a narrow canal. From their observations, Begun & Gomoiu (2007) concluded that both species are episodically found in fresh-water but are not capable of forming stable populations there. For the complex history of the taxonomy of M. slabberi see ‘Discussion’. FIGURE 2. Distribution of Hemimysis anomala G. O. Sars, 1907, and Mesopodopsis slabberi (Van Beneden, 1861), in tributaries and coastal waters of the Mediterranean and Black seas. Data original and from 135 literature sources. Hemimysini Czerniavsky, 1882, revalidated at tribe rank Fig. 3 Diagnosis. Mysinae with antennal scale setose except for a large proximal bare portion of the outer margin, this portion rostrally ending in a number of setae or in a subterminal series of articulate spines, scale without or with small apical segment; first thoracic endopod 7-segmented, with distinct endites on coxa, basis and ischium, and no endite or a less distinct one on the merus; ischium always well developed, its inner margin with more than half the length of that of the merus; carpopropodus of the thoracic endopod 6 with 3–6 segments; well-developed oostegites on thoracopods 7, 8, and rudimentary oostegite on thoracopod 6; all pleopods of females and pleopods 1, 2 of males rudimentary; male pleopods 3–5 each with well-developed sympod; male pleopod 3 with 1- to 2-segmented endopod and with short to minute exopod or without exopod at all; male pleopod 4 biramous, with 2-segmented sympod, with short, 1- or 2-segmented endopod, and with elongate, 6- to 7-segmented exopod, the latter with modified seta on both the penultimate and ultimate segment; male pleopod 5 biramous, with exopod and endopod, each 3- to 6-segmented; statoliths composed of fluorite or of vaterite; telson terminally truncate or more frequently with apical cleft, in any case medio-apically with > 10 spine-like laminae, no apical setae. Type genus. Hemimysis G. O. Sars, 1869 (9 species), only this genus known. Occurrence. Endemic in the NE-Atlantic, Mediterranean, and Ponto-Caspian, including the Baltic and Marmora Seas. For the anthropogenic expansion of Hemimysis anomala to the Baltic and to continental and coastal waters of western Europe and North America, see below. Several Hemimysis species are strongly sciaphilic, found in cave entrances and (micro)caves during the daytime; from here they emerge during twilight and night (Macquart-Moulin & Patriti 1966, Ledoyer 1989, Wittmann 2001, Delgado et al. 2013). 10 · Zootaxa 4142 (1) © 2016 Magnolia Press WITTMANN ET AL. Genus Hemimysis G. O. Sars, 1869 Hemimysis anomala G. O. Sars, 1907 Fig. 3 Hemimysis anomala G. O. Sars, 1907. Short selection from 306 references: Băcescu 1940; Daufresne et al. 2007; Porter et al. 2008; Wittmann & Ariani 2009; San Vicente 2010; Daneliya & Petryashev 2011; Daneliya et al. 2012; Golaz & Väinölä 2013; Wittmann et al. 2014; Roth 2015. Material examined (all samples from Mediterranean drainage, Rhône system, leg. K. J. Wittmann). 5 M ad. 5.0– 7.4 mm, 1 F subad., 5 imm., 4 juv., southern France, left bank of the river branch Grand Rhône at Port St. Louis, at Rhône km 323.2, 43.3827N 004.8073E, altitude 0 m, sea distance 6 km, 3–4 m depth, stones and mud, v = 0 m/s, S = 0.1–3.5, 426–6300 µS/cm, 22.5–21.7°C, pH 7.63–7.28, 7.35–6.55 mg O2/l, 11–12°d, 17–32 NTU, bottle traps exposed over night, 17/18 June 2009, NHMW reg. no. 25703; 1 M ad. 6.1 mm, 1 F subad., southern France, left bank of the river branch Petit Rhône at Tiki, 43.4512N 004.3977E, sea distance 150 m, 1.5–2.5 m depth, boulders with algae and gravel, strong wind-driven (Mistral) waves, v = 0–0.2 m/s, S = 1.5–1.7, 3040–3480 µS/cm, 23.7– 21.6°C, pH 7.93–7.65, 7.25–6.40 mg O2/l, 25–17 NTU, bottle traps exposed over night, 19/20 June 2009; 8 F ad. 6.4–8.6 mm, 14 M ad. 6.0–8.7 mm, 35 F subad., 6 M subad., 53 imm., 110 juv., southern France, right bank of Rhône River, ca. 2 km south of Beaucaire, at Rhône km 269.2, 43.7876N 004.6486E, altitude 8 m, sea distance 61 km, 2–2.5 m depth, v = 0.05–0.15 m/s, S = 0.1, 356 µS/cm, 21.5°C, pH 6.9, 6.16 mg O2/l, 9°d, 99 NTU, bottle traps exposed over night, 15/16 June 2007, NHMW 25702; 1 M ad. 7.9 mm, 1 F subad., 1 M subad., 3 imm., eastern France, right bank of Rhône River below weir near Wievroz, 45.8126N 005.0919E, altitude 179 m, sea distance ca. 352 km, 2–4 m depth, boulders and gravel, v = 0.2–0.3 m/s, S = 0.1, 294–304 µS/cm, 21.2–21.3°C, pH 7.32–6.78, 5.58–4.77 mg O2/l, 8°d, 24–19 NTU, bottle traps exposed over night, 29/30 June 2009; 1 M imm., eastern France, left bank of Rhône River at re-confluence with its small side branch near Chateaufort, 46.9299N 005.8247E, altitude 252 m, sea distance ca. 464 km, 0.2–3 m depth, bank macrophytes, Myriophyllum, detritus, soft bottom, v = 0.1–0.3 m/s, S = 0.1, 285 µS/cm, 19.8°C, pH 8.06, 5.85 mg O2/l, 8°d, 17 NTU, hand net, 30 June 2009, day; 175 juv., Switzerland, Lake Geneva, yachting harbour near Geneva, 46.2303N 006.1878E, altitude 371 m, sea distance ca. 531 km, 1.5–4 m depth, boulders with Myriophyllum and Zannichellia, S = 0.1, 269 µS/cm, 21.2°C, pH 8.29, 7.66 mg O2/l, 8°d, 0 NTU, hand net, 30 June 2009, day, NHMW 25704. Description (Fig. 3). Hemimysini with eyes well developed, cornea large, globular; eyestalks short (Fig. 3A). Antennal scale suboval, terminally rounded; inner and outer margins without spines, densely setose except for the smooth proximal half of the outer margin; terminal segment is only 3–7% scale length; antennal sympod with dorsal lobe projecting anteriorly above and shortly beyond basal segment of endopod (Fig. 3B). Rostrum very short, almost missing, its form sinusoid (Fig. 3C) or forming a terminally rounded, wide angle (Fig. 3A). Maxillary palpus with transversely flattened terminal margin bearing 4–6 setae and 10–13 articulate spines (spine-like setae) in between. Carapace almost smooth (Fig. 3C); neck, however, strengthened by cuticularized bars along median portions of the cervical sulcus; 4–6 cervical pores directly in front of this reinforcement; almost straight series of about 25–35 cardial pores transversely crossing above the heart. A framework of cuticularized bars strengthens the thoracic sternites (Fig. 3D); apart from this the sternites are smooth, except for a number of minute, spine-like scales on the (as usual) anteriorly directed lobe of the first thoracomere (Fig. 3D). First thoracic endopod without endite on merus; its epipod leaf-like, without seta (Fig. 3D); carpopropodus of endopods 3–8 with 5–6, (5)–6, 6, 6, 6, or 5–6 segments, respectively; claw very strong (Fig. 3D) in endopods 1 and 2, whereas weaker, but always distinct (Fig. 3E) in endopods 3–8. Basis of all thoracic exopods with well rounded outer corner; flagellum 8segmented in exopods 1 and 8, but 9-segmented in exopods 2–7, not counting the large intersegmental joint between basis and flagellum which may be mistaken as a segment. Penes normal, of moderate size, each with 8–11 curved setae around the ejaculatory opening. Male pleopods 3–5 each with large, 2-segmented sympod; male pleopod 3 with small, unsegmented endopod, and with rudimentary, minute, knob-like exopod (Fig. 3F); pleopod 4 with small, 2-segmented endopod and with long, 6-segmented exopod showing two large modified setae, one each on penultimate and terminal segment (Fig. 3G); pleopod 5 well-developed, setose, biramous, its exopod 3- to 4segmented, its endopod 2- to 4-segmented (Fig. 3H). Scutellum paracaudale well rounded, sinusoid in both sexes (Fig. 3J, K). Endopods of uropods with 6–9 spines in linear series from subbasal to subterminal portions of inner MYSIDAE IN MEDITERRANEAN FRESH-WATERS Zootaxa 4142 (1) © 2016 Magnolia Press · 11 FIGURE 3. Hemimysis anomala G. O. Sars, 1907, from the Rhône River at river-km 269, females with 9.2 mm (A) or 8.6 mm (J, L, M), and male with 8.7 mm (B–H, K) body length. A, habitus of female, dorsal; B, left antenna, dorsal; C, carapace expanded on slide, dorsal, pores not to scale; D, thoracic sternites 1–8 (ventral) with first thoracopod (caudal); E, 'tarsus' of left thoracic endopod 3, rostral; F, male pleopod 3, rostral; G, male pleopod 4, caudal; H, male pleopod 5, caudal; J, K, posterior margin of sixth pleonite in female (J) or male (K), lateral; L, right uropod, ventral; M, telson, dorsal. 12 · Zootaxa 4142 (1) © 2016 Magnolia Press WITTMANN ET AL. margin (Fig. 3L). Statoliths composed of vaterite. Telson rhombohedral, terminally truncate, i.e. without apical cleft (Fig. 3M); lateral margins each with 14–19 spines, not counting the pair of large latero-apical spines; terminal margin with 11–16 laminar, apically acute processes. Body colour bloody-red, pink to translucent, often with ivoryyellowish tinge; intensively red upon expanded chromatophores, whereas nearly translucent with numerous dark red spots upon contracted chromatophores. Apart from these short-term variations the colours become generally intensified with increasing body size (age) (Salemaa & Hietalahti, 1993). Bionomy. Adult body size mostly 6–11 mm, total range 5–17 mm; adult males on average slightly smaller than females. The species is strongly euryhaline, inhabiting a salinity range of S = 0–19, mostly in less than 10 m to a maximum of about 50 m depth (Băcescu 1954, Zhuravel 1960, Komarova 1991, Kelleher et al. 1999). During daytime it is strongly photophobic, (epi)benthic, gathering in swarms in shelters on the bottom. At night it swims more actively, at a greater distance from the bottom, often up to the surface. The diurnal vertical migration is clearly determined by the factor light (Borcherding et al. 2006). Near sunset, juveniles apparently emerge earlier at higher light levels compared to adults (Boscarino et al. 2012). Hemimysis feeds on detritus, algae, and zooplankton (Băcescu 1954, Borcherding et al. 2006). The mysids adapt their feeding behaviour to environmental conditions and to the availability of pelagic and benthic food sources (Marty et al. 2012). The portion of zooplankton consumed increases with increasing body size (Borcherding et al. 2006). The omnivorous habit of the species together with voracious predatory feeding of the adults may considerably affect the zooplankton composition (Ketelaars et al. 1999). Hemimysis is often found in the stomach of perch, pikeperch, and various other species of fish (Kelleher et al. 1999, Borcherding et al. 2006, Brooking et al. 2010, Lantry et al. 2012). Consumption by fish may show strong quantitative variations between seasons and years (Lantry et al. 2012). In the northern Black Sea and in the northern Baltic, H. anomala breeds from April to October (Băcescu 1954, Komarova 1991, Salemaa & Hietalahti 1993), in the alpine Lake Geneva from March to October (Golaz & Väinölä 2013). Numbers of eggs per brood are mostly in the order of 6–35, total range 2–70, with strong variations according to body size, season, and locality (Ketelaars et al. 1999, Borcherding et al. 2006, Pothoven et al. 2007, Marty 2008, Golaz & Väinölä 2013, Borza 2014). H. anomala is clearly a warm season breeder (Wittmann 1984) with young being produced from spring to autumn and the overwintering generation being born in autumn and reproducing in spring. In an artificial embayment of the Hungarian Danube, it produces four generations per year and shows very high fecundity, particularly by the overwintering generation with a mean of 43 young per breeding female (Borza 2014). Dumont & Muller (2009) reported that the species reproduces in waters of Alsace (NE-France) only three times a year, in March/April, June/July, and September/October. Here the overwintering population starts to reproduce in early March, when the water warms to 7–8°C. Distribution (Fig. 2). This Pontocaspian endemic was originally confined to the lower reach of rivers, to estuaries and to coastal 'marine' environments of the Caspian, Black, and Azov Seas (Băcescu 1937, 1954; Derzhavin 1939; Mordukhai-Boltovskoi 1960; Băcescu et al. 1971; Komarova 1991). By intentional transplantations and by non-intentional modes of areal expansion H. anomala reached tributaries and coastal waters of the NE-Atlantic, including waters of Ireland plus England, and tributaries of the Bay of Biscay, North Sea and Baltic. It also reached tributaries of the NW-Mediterranean, Lake Aral and even the Great Lakes of North America (see ‘Discussion’). Paramysini Wittmann, Ariani & Daneliya, new tribe Figs 4, 5, 7A, B, 8–10, 11A Diagnosis. Mysinae with antennal scale setose except for a large proximal bare portion of the outer margin, this portion rostrally ending in an articulate or a non-articulate spine, scale in most species with small apical segment; first thoracic endopod large, 7-segmented or only 6-segmented if there is no suture between coxa and basis, in each case with large endite on basis, and with endite of variable size on ischium, whereas endite absent or present on merus; ischium always well developed with more than half the size of the merus; carpopropodus of thoracic endopod 6 with 3–9 segments; well-developed oostegites on thoracopods 7, 8, and very small or rudimentary oostegite on thoracopod 6; all pleopods of females and pleopods 1, 2, 5 of males rudimentary; male pleopod 3 biramous, with 1- to 2-segmented endopod and with unsegmented rod-like exopod; male pleopod 4 biramous, its endopod short, 1- to 2-segmented, its exopod elongate, 5- to 7-segmented, terminally with modified seta(e); recent MYSIDAE IN MEDITERRANEAN FRESH-WATERS Zootaxa 4142 (1) © 2016 Magnolia Press · 13 statoliths composed of vaterite or fluorite, fossil statoliths composed of calcite (probably derived from the metastable vaterite by phase transformation during fossilization); telson variable, mostly with apical cleft, medioapically with at least two (as exceptional deviation with 0 or 1 in Katamysis and Paramysis s.str.) spine-like laminae (mostly > 15), no apical setae (not counting minute hairs such as found in Paramysis bakuensis G. O. Sars, 1895). Type genus. Paramysis Czerniavsky, 1882. Taxa included. Six genera: Caspiomysis G. O. Sars, 1907 (1 species), Katamysis G. O. Sars, 1893 (1), Paramysis Czerniavsky, 1882 (24), Praunus Leach, 1814 (3), † Sarmysis Maissuradze & Popescu, 1987 (3 fossil species), Schistomysis Norman, 1892 (6). Distribution. Endemic in coastal waters and tributaries of the NE-Atlantic, Mediterranean, and PontoCaspian, including White Sea, Barents, Baltic and Marmora Seas. Anthropogenic expansion of Katamysis warpachowskyi G. O. Sars, 1893 to inland fresh-waters of central and western Europe (Wittmann 2002, 2008; Hanselmann 2010). For expansion of Paramysis species to Lake Aral, etc., see below. Genus Paramysis Czerniavsky, 1882 Diagnosis. Paramysini with antennal scale showing a large, basal, smooth portion of the outer margin; this portion ending in a thorn (non-articulated spiniform process); setose apex bears a small apical segment with plumose setae (Figs 4B, 8H). Eyes well developed with large, dorsoventrally slightly flattened cornea (Figs 4A, 8A, B). Maxillary palp normal, its setose terminal segment laterally moderately (Fig. 9C) to strongly expanded, with anterodistal setae only or with additional 1–3 short, subapical spiniform setae (Paramysis s.str.). First thoracic endopod (Fig. 9D) large, 7-segmented, with endites on all segments from coxa to merus, the largest endite on the basis, endite of merus larger than endite of ischium. Thoracic endopods 3–8 (Figs 4H, 9G, H, 10A, D, F, H) each with carpus entire, short, stout, with brush of setae; propodus normally 3-segmented, but reduced with fewer segments in posterior endopods of a few species; small to minute dactylus with normal to very weak nail, this nail may appear seta-like on endopod 8. Penis normal, with caudolateral blade (Figs 5A, 9H). Male pleopods 3 (Figs 5B, 10O) and 4 (Figs 5C, 10P) each with two-segmented sympod; endopods 3 and 4 one- or two-segmented; exopod 3 rod-like, unsegmented; in contrast, exopod 4 is 6-segmented (Fig. 10P, Q) to 7-segmented (Fig. 5C, D) with one large, modified seta on both penultimate and ultimate segment. Male pleopods 1, 2, 5 (Figs 5E, 10M, N, R), and all pleopods of females, reduced, rod like with allusively to weakly developed pseudobranchial lobe. Uropods setose all around (Figs 5H, 10T), endopod shorter than exopod; statolith composed of vaterite or much less frequently of fluorite; variable numbers of spines below statocyst and often also more caudally along ventral face of endopod. Telson with spines along lateral margins; each lateral margin ends in a distinctly larger spine; terminal margin between these spines is transversely straight or excised, this margin furnished by at least two (as exceptional deviation with 0 or 1 in Paramysis s.str.), mostly many more acute laminae (Figs 5J, 10U, V). Type species. Paramysis Baeri Czerniavsky, 1882b: 56, as defined in Daneliya (2004). Distribution (Figs 6, 12). Endemic in the NE-Atlantic, Mediterranean, and Ponto-Caspian. Anthropogenic expansion of P. lacustris (Czerniavsky, 1882) and additional species to coastal waters and tributaries of the Baltic Sea, Lake Aral, and to inland water systems of northern and eastern Europe (Zhuravel 1950, Gasiunas 1968, Aladin et al. 1998, Wittmann 2007, Semenchenko et al. 2007, Zettler 2015). Subgenus Serrapalpisis Daneliya, 2004 Short diagnosis (characters for distinction of Mediterranean subgenera only; for full diagnosis see Daneliya (2004)). Median segment of mandibular palp with at least some (Fig. 4 D, E), mostly with many strong, serrated setae along its outer margin. Propodus of thoracic endopods 3–8 is 3-segmented (Fig. 4H). Telson with wide cleft that forms an angle of >120° (Fig. 5J). Laminae of cleft margin widened by wing-like expansions (Fig. 5K). Type species. Mesomysis lacustris Czerniavsky, 1882b: 42, as defined in Daneliya (2004). 14 · Zootaxa 4142 (1) © 2016 Magnolia Press WITTMANN ET AL. Paramysis (Serrapalpisis) kosswigi Băcescu, 1948 Figs 4, 5 Paramysis (Mesomysis) kosswigi Băcescu, 1948: Mordukhai-Boltovskoi 1979. Paramysis kosswigi: Băcescu 1954, 1966, 1985; Remane & Schlieper 1958; Mordukhai-Boltovskoi 1964a, b; Mauchline & Murano 1977; Komarova 1991 (partim); Kocataş et al. 2003 (partim); Daneliya & Petryashev 2011; Wittmann & Ariani 2011; ITIS 2014. Mysis relicta: Dügel & Kazanci 2004. Paramysis (Serrapalpisis) kosswigi: Anderson 2008; Audzijonyte et al. 2008a; Daneliya et al. 2012; Mees 2014. nec Paramysis kosswigi (Black Sea records): Şerban et al. 2000; Şerban 2004; Skolka 2005; Audzijonyte 2006; Özbek & Ustaoğlu 2006; Porter et al. 2008; Akbulut et al. 2009. Material examined (all from freshwater tributaries of the Aegean Sea in Turkey). 8 F ad. 8.7–11.1 mm, 29 M ad. 7.6–9.9 mm, 12 F subad., 8 M subad., 38 imm., 78 juv., Lake Işikli in Anatolia, 38.2667N 029.9254E, altitude 818 m, sea distance ca. 505 km, 0.5–1 m depth, over sand, S = 0.1, 320 µS/cm, 24.4°C, pH 8.1, 7.24 mg O2/l, 10°d, 55 NTU, hand net, 11 June 2006, leg. K. J. Wittmann, NHMW reg. no. 25198, BMNH 2011.1754-1763, MZUT; 9 ad. spec. 8.0–11.0 mm, same hydrological system, Işikli springs (type locality), 38.3225N 029.8507E, altitude 835 m, Sept. 1998, leg. C. Aygen, MZH HLA.150700, HLA.150701; 5 F ad., 9.5–10.5 mm, 4 M ad. 7.0–10.0 mm, 2 heads, drainage system of River Küçük Menderes, Oedemis [= Ödemiş, altitude ≈ 135 m], 38.24N 027.96E, 1971, leg. M. Băcescu, MGAB MYS 163. Supplementary description. All features within the ranges indicated above for the genus and subgenus. Head dorsally convex (inflated). Antennal scale (Fig. 4B) length is 2.6–3.6 times its maximum width; scale with smooth outer margin extending beyond terminal margin of antennular trunk in females but not so in males (Fig. 4A); setose apex is 20–27% length of scale; small apical segment with five plumose setae (Fig. 4B). Eyes large, dorsoventrally only weakly flattened; cornea hemispherical in dorsal view (Fig. 4A); maximum diameter of cornea is 42–49% antennal scale length; outer basal edge of eyestalks with field of scales. Wedge-shaped subrostral process extends straight forward well beyond the short, evenly rounded rostrum (Fig. 4A). This process is roughly triangular in dorsal view. Carapace (Fig. 4A) with medium-sized, subtriangular, basally rounded, mid-dorsal, posterior emargination; carapace leaves 1.5–2 posterior thoracic somites dorsally exposed; mid-dorsally it shows traverse row of 15–23 pores in cardial position (above the heart), arranged in two roughly symmetrical subgroups, and 9–14 pores in cervical position. Median segment of mandibular palp with several smooth and a number of serrated setae along its outer margin; serration along anterodistal portion of setae (Fig. 4D, E). First thoracic sternite with anteriorly directed, hairy median lobe contributing to the caudal closure of the mouth area in both sexes (Fig. 4F, G); sternites 2–5 of adult males with median, apically rounded processes of different size and shape, equipped with hairs (Fig. 4F); adult females without such processes but with hairs only on sternites 2–5 (Fig. 4G); sternites 6–8 smooth, without humps, ridges or hairs in both sexes (Fig. 4F, G). Thoracic endopods 3–8 each with 5-segmented tarsus showing very dense setation (Fig. 4H); propodus three-segmented throughout; dactylus minute, cylindrical, with weak, mostly straight, but terminally curved nail. Paradactylar setae of endopods 3–8 sickle-shaped, their armature with denticles is stronger in females (Fig. 4H, J) compared to males (Fig. 4K) and successively decreases in both sexes from endopods 3 to 8. Paradactylar setae of endopods 5–8 with denticles only on proximal half; extension of denticles more variable, generally longer, in endopods 3, 4 (Fig. 4H–K). Endopod 3 with inner (= rostral) paradactylar seta (Fig. 4H–K) showing a heavy armature of 5–8 denticles (i.e. stiff secondary projections) along its median portions; proximal denticles smaller and in part weakly serrated; outer (= caudal) paradactylar seta with much weaker armature (Fig. 4H) compared to the inner one. Merus of endopods 3–8 with 4–5 posterior bunches of setae. Penis with 3–4 short, barbed setae in subterminal position on caudolateral blade (Fig. 5A). Pleopods of both sexes (for males in Fig. 5B–E) as described above for the genus. Male pleopod 4 (Fig. 5C, D) with 7-segmented exopod and unsegmented endopod; second segment of its exopod with strong cuticularized ridge on rostral face. Such ridges are present in several species of Paramysis; instead there is a distinct lobus at this spot in P. arenosa (G. O. Sars, 1877). In P. kosswigi the third segment of exopod 4 shows strongly oblique segmental borders with both its neighbouring segments; this obliquity appears stronger in lateral (Fig. 5D) than in rostral (Fig. 5C) view. Scutellum paracaudale linguiform, biconvex; terminally undulate to serrated (Fig. 5F, G). Uropods (Fig. 5H) normal, endopod is 74–80% length of exopod; endopods ventrally with 3–6 spines near inner margin between statocyst and 24–39% endopod length from tip; spines increasing in length distally. Statoliths composed of vaterite. MYSIDAE IN MEDITERRANEAN FRESH-WATERS Zootaxa 4142 (1) © 2016 Magnolia Press · 15 FIGURE 4. Paramysis (Serrapalpisis) kosswigi Băcescu, 1948, from Lake Işikli (A–D, F–K) and its springs (E, type locality) in Anatolia. A, anterior body region of adult male with body length 8.7 mm, dorsal view (pores on carapace and scales on eyestalks not to scale); B, antenna of male 9.1 mm, ventral; C, left mandibular palp of male 9.1 mm, rostral; D, detail of (C) showing setae from the outer margin of the median segment; E, setae as in (D), female 10.5 mm; F, thoracic sternites 1–8 of male 8.9 mm, ventral; G, the same for female 10.1 mm; H, left 'tarsus' (carpopropodus plus dactylus) of thoracic endopod 3 in female 10.1 mm, rostral; J, anterior paradactylar seta in female 9.5 mm; K, the same for male 9.1 mm. 16 · Zootaxa 4142 (1) © 2016 Magnolia Press WITTMANN ET AL. FIGURE 5. Paramysis (Serrapalpisis) kosswigi Băcescu, 1948, from Lake Işikli in Anatolia. A, left penis of male with body length 8.2 mm, outer lateral; B, third pleopod in male 8.7 mm, rostral face; C, the same for fourth pleopod; D, exopod of fourth pleopod in lateral view, setae omitted, male 9.6 mm; E, fifth pleopod in male 8.7 mm, rostral; F, posterior margin of sixth pleonite in female 10.1 mm, lateral; G, the same for male 9.1 mm; H, uropods of male 8.7 mm, ventral; J, telson of same male, dorsal; K, detail of (J), showing right terminal spine and its neighbouring laminae. MYSIDAE IN MEDITERRANEAN FRESH-WATERS Zootaxa 4142 (1) © 2016 Magnolia Press · 17 Telson (Fig. 5J) subrectangular, slightly trapezoid, with very weakly curved, allusively S-shaped lateral margins; telson length is 1.8–2.1 times maximum width or 1.0–1.2 times length of sixth pleonite (measured along dorsal midline); lateral margins with 12–16 spines each, not counting the large apical spines; distal spine-free region between terminal and lateral spines extends over 14–21% length of lateral margins; telson terminally with wide subtriangular cleft that forms an angle of >120°; cleft is 7–10% telson length and does not reach the subterminal spines; cleft with convex margins on each symmetrical half; cleft lined by a total of 21–27 acutely pointed laminae. Occurrence (Fig. 6). Paramysis kosswigi is a stenoendemic in the freshwater lake Işıklı in Anatolia, including its springs (type locality) and its effluent Büyük Menderes Nehri (Great Maeander) to the Aegean Sea (EMediterranean), also occurring in the near, roughly parallel system of Küçük Menderes Nehri (Little Maeander). Records from altitudes of 135 m (orig.) to 1007 m (Kocataş et al. 2003) are acknowledged by the present authors. Previous records from the Turkish Black Sea coast appear doubtful (see ‘Discussion’). FIGURE 6. Distribution of Diamysis hebraica Almeida Prado-Por, 1981, D. lagunaris Ariani & Wittmann, 2000, and Paramysis kosswigi Băcescu, 1948, in tributaries and coastal waters of the Mediterranean Sea. Data are original and from Băcescu (1948, 1954), Almeida Prado-Por (1981), Ariani & Wittmann (2000), Kocataş et al. (2003), and Wittmann & Ariani (2011, 2012a). Subgenus Longidentia Daneliya, 2004 Occiparamysis Daneliya, 2004, syn. nov. Revised diagnosis. Anterior margin of carapace convex, smoothly rounded. Antennal scale with clearly produced distal part (Fig. 8B, H). Median segment of mandibular palp with normal, smooth or at most slightly barbed setae (Fig. 9A). Thoracic endopods 3–8 with 3-segmented propodus (Figs 9G, H, 10A, D, F, H). Penis with row of short, barbed setae in subterminal position on caudolateral blade (Fig. 9H). Telson roughly as long as last abdominal segment, with strong, subrectangular cleft (Fig. 10U). Cleft laminae without lateral wing-like expansions (Fig. 10V). Type species. Mesomysis Kröyeri Czerniavsky, 1882b: 46, as defined in Daneliya (2004). Remarks. The characters of P. adriatica sp. nov. take an intermediate position between the subgenera Longidentia and Occiparamysis, thus diminishing their diagnostic significance. In particular, being rather similar to P. agigensis Băcescu, 1940 (see details below), P. adriatica still lacks the most important distinguishing character of Occiparamysis—long and strong setae on penis outer lobe—rather having short, thin setae like in the species of Longidentia. Thus, upon synonymising Occiparamysis with Longidentia, we also propose the new, revised combination Paramysis (Longidentia) agigensis Băcescu, 1940. The redefined subgenus Longidentia is 18 · Zootaxa 4142 (1) © 2016 Magnolia Press WITTMANN ET AL. most similar to the subgenus Pseudoparamysis Băcescu, 1940, from which it differs by the 3-segmented propodus of thoracic endopods 3–8 (reduced, 1- or 2-segmented propodus of endopods 7–8 in species of Pseudoparamysis). Paramysis (Longidentia) adriatica sp. nov. Figs 7A, B, 8–10, 11A Mysis oculata relicta: Stammer 1932 (partim: station Tb9"). Paramysis helleri: Holmquist 1955; Ariani et al. 1993: Tab. I (partim: Split); Ariani 2004: fig. 3B. Type material. Holotype male with 9.1 mm body length (NHMW reg. no. 25694), 21 paratypes (16 F ad. 9.3–11.7 mm, 4 M ad. 7.9–8.9 mm, 1 F subad., NHMW 25695, MZH HLA.112100, MZUN, SMNH Type 8783, MFSNU) (co-occurring with eight Diamysis fluviatilis), NW-Adriatic coast, Adige River, 9 km from its mouth into the sea, 45.1160N 012.2684E, altitude 0 m, 0.4–0.8 m depth, among flooded bank vegetation after several days of rain, roots and branches of Salix, v = 0.05 m/s, S = 0.1, 305 µS/cm, 10.6°C, pH 5.48, 8.27 mg O2/l, 8°d, 37 NTU, 14 April 2012, hand net, leg. Ariani & Wittmann. Additional material examined (stations listed in clockwise order along Adriatic coasts). 4 F ad. 9.8–11.5 mm, 2 imm., NW-Adriatic coast, Po di Goro = mouth branch of Po River, at Mesola, 44.9251N 012.2310E, 27 km to the sea, altitude 2 m, 0.2–0.7 m depth, flooded bank vegetation, Salix, concrete, v = 0.03–0.06 m/s, S = 0.1, 442 µS/cm, 13.3°C, pH 6.43, 7.14 mg O2/l, 8°d, 81 NTU, hand net, 14 Apr. 2012, leg. Ariani & Wittmann, NHMW 25697; 1 F ad. 9.3 mm, 1 F imm. 8.7 mm, N-Adriatic coast, near Panzano, Canale Brancolo, 45.7804N 013.5129E, 4 km to the sea, altitude 0 m, benthic, 29 Sept. 1986, leg. F. Stoch & B. Zanolin, MFSNU; dissected parts of 1 M ad. 7.5 mm, 1 F ad. 8.5 mm with 2 postnauplioid larvae, on a total of 2 slides labelled "Paramysis helleri. Timavo. Det. Ch. Holmquist. prep. 64, 65", SMNH reg. nos. 140106, 140107, according to Stammer (1932) and Holmquist (1955: Tab. 1) this material was collected in 1928/29 by Hans Jürgen Stammer in the former right mouth branch of the Timavo, 45.7839N 013.5733E, Gulf of Trieste, NE-Adriatic; 1 F ad. 8.8 mm, 1 F imm. 6.6 mm, E-Adriatic coast, small freshwater creek at Split, roughly 43.515N 016.541E, 0–1 m depth, macrophytes, hand net, 26 Oct. 1976, leg. A. P. Ariani, NHMW 25698; 6 M ad. 8.3–9.4 mm, 4 F ad. 9.4–10.9 mm, 2 F subad., 1 F imm., E-Adriatic coast, lake Deran (= Deransko jezero), oligotrophic karstic lake, 43.04N 017.75E, altitude 0 m, total of about 31 km along its small effluent and following this along the Neretva River to the Adriatic Sea, sample labelled ”in clean and cold stagnant water in areas with mud bottom, specimens caught over the fine muddy sediment without any cover”, 0.5– 1 m depth, S = 0.2, 13–15°C, 9 July 2009, leg. D. Lučić & P. Tutman, NHMW 25696. Derivatio nominis. The species name is a Late Latin adjective with female ending, referring to the so far known exclusive occurrence of the new species in tributaries of the Adriatic Sea. Type locality. NW-Adriatic drainage basin, in fresh-water of the lower reach of the Adige River, 45.12N 012.27E (Fig. 12). This is the second-largest tributary of the Adriatic Sea. Occurrence (Fig. 12). In freshwater tributaries of the Adriatic Sea. So far known only from five running waters and one lake, in each case less than 5 m above sea level; maximum sea distance 31 km. This enumeration includes the former right mouth branch of the Timavo, now dry land. Whether Stammer (1932) found any Paramysis in other parts of the Timavo system cannot be judged currently because there is no Timavo material in the Stammer collection at the Department of Biology, University of Erlangen, Germany (Wolfgang Heimler, pers. comm.). The Swedish Museum keeps only those Timavo materials inspected by Holmquist (1955) and 60 years later by us. Inspection of Timavo springs and estuary by one of us (KJW) in Sept. 1987 yielded no Paramysis materials. Definition. Antennal scale length is 2.5–2.6 times its maximum width; scale with smooth outer margin ending slightly before or at the terminal margin of antennular trunk or more often extending beyond this margin (Fig. 8B), in any case ending at a non-articulated spiniform process; setose apex is 15–25% length of scale; apex bears a small apical segment (Fig. 8H). Eyes with large, reniform cornea (Fig. 8B); maximum diameter of cornea is 42–53% antennal scale length; ventral face of eyestalks with traverse stripe of hair-like scales. Wedge- to sickle-shaped, mostly upward-bent, subrostral process extends well beyond the short, evenly rounded rostrum (Fig. 8B, C). This process appears subtriangular in dorsal view (Fig. 8B). Carapace with medium-sized, subtriangular, basally rounded, mid-dorsal, posterior emargination (Fig. 8C). Median segment of mandibular palp with normal (i.e. not MYSIDAE IN MEDITERRANEAN FRESH-WATERS Zootaxa 4142 (1) © 2016 Magnolia Press · 19 microserrated) setae (Fig. 9A). Thoracic sternites 2–5 of adult males with median, apically rounded processes of different size, terminally equipped with hairs (hair-like scales); sternites 6–8 only with smooth, less distinct humps (Fig. 9D, E); no such processes in adult females. Thoracic endopods 3–8 with 5-segmented tarsus (Figs 9G, H, 10A, D, F, H); carpus as typical for the genus; propodus 3-segmented; dactylus minute, cylindrical, with small setalike nail. Endopod 3 with inner paradactylar seta (Fig. 10C, J) showing a heavy armature with 4–6 spinules (i.e. stiff secondary projections) along its median to subterminal portions; proximal spinules thinner, their (sub)terminal portions in part with secondary 'hairs'; outer paradactylar seta with much weaker armature (Fig. 10A, H). Penis with row of 4–6 short, barbed setae in subterminal position on caudolateral blade (Fig. 9H; same type of setae as in P. kosswigi, Fig. 5A). Pleopods of both sexes (of males in Fig. 10M–R) as described above for the genus; male pleopods 3 (Fig. 10O) and 4 (Fig. 10P) each with unsegmented endopod; male pleopod 4 with 6-segmented exopod (Fig. 10P, Q). Endopod of uropods with only 3–6 spines along inner margin between statocyst and 35–45% endopod length from tip (Fig. 10T). Statoliths composed of vaterite. Telson with weakly curved, allusively Sshaped lateral margins; margins with 13–15 spines, not counting the (not or) moderately larger apical spines (Fig. 10U, V). As an unusual state among species of Paramysis, shorter spines may alternate with longer ones on lateral margins of telson (Fig. 10U; this pattern distinct in 18 out of 30 specimens examined, indistinct or missing in the remaining ones; terminal spines not considered). Triangular terminal cleft is 9–12% telson length, cleft not or barely reaching the subterminal spines (Fig. 10U, V); cleft with almost straight margins on each symmetrical half, cleft armed with a total of 14–28 laminae of in part unequal size, even the longest laminae shorter than the subterminal or terminal spines, respectively. Description (types only). All features within the ranges indicated above for the genus. Appendage morphology and habitus of both sexes (Figs 7A, B, 8A, B) very similar to those of P. agigensis and P. helleri. Carapace leaves 1.5–2 posterior thoracic somites dorsally exposed. Carapace with traverse row of 16–22 pores in cardial position (above the heart), arranged in two roughly symmetrical subgroups, and 7–10 pores in cervical position (Fig. 8C–E). Eyes dorsoventrally flattened. The cornea is reniform in dorsal view (Fig. 8B), ovoid in lateral view (Fig. 8A). Male antennula with slender appendix masculina facing obliquely downwards in the vertical plane (Fig. 8A) and forwards in the horizontal plane (Fig. 8F). This appendix is 2.0–2.3 times the length of apical segment of antennular peduncle. Sympod of antenna produced into a large spiniform process at its outer margin (Fig. 8H). Antennal scale (Fig. 8B, H) is 1.1–1.5 times the length of antennular peduncle (Fig. 8A, B, F, G). Large non-articulated spiniform process (thorn) at 75–90% scale length from basis. Small terminal segment of scale with five large plumose setae. Basal segment of mandibular palp without setae, but with dense field of scales (Fig. 9A). Remaining two segments setose. Median segment with smooth setae in both sexes; with a minor exception in males only, insofar as the basal three setae on the outer margin bear up to four subbasal to median barbs each (Fig. 9A). Terminal segment half as long as the median one. Exopod of maxilla roughly fan-shaped, with 12–16 mostly long, plumose (Fig. 9C) setae; exopod not larger than distal segment of palpus (= endopod). This palpus with terminal segment laterally widened, about 1.7–1.8 times wider than long. First thoracic sternite produced into an anteriorly directed, distally hairy, medial lobe in both sexes (Fig. 9D). Unlike the sterna of males given in the definition above, the sternites 2–8 of females are medially smooth, without any humps or hairs. Basis of all thoracic exopods laterally expanded, blade-like; its distal outer corner with spinelike extension in thoracopods 1–7 (Fig. 9D), whereas well-rounded in thoracopod 8 (Fig. 9H). Flagellum of exopods 1 and 8 each with eight segments, whereas in exopods 2–7 normally with nine segments, not counting the large intersegmental joint between basis and flagellum which may be mistaken as a segment. In contrast, the flagellum of exopod 7 showed only eight segments in one out of ten females inspected in this respect. Thoracic endopods rather stout, moderately long. Total length increases from endopods 1 to endopod 3, and then decreases backward to endopod 8. Endopod 8, when stretched anteriorly, extends only up to basis of endopod 3. First thoracic epipod with groups of minute scales along its anterior margin and on intersegmental joint with its sympod (Fig. 9D). This sympod with additional groups of such scales and two plumose setae near insertion of epipod. First thoracic endopod with non-setose endite from coxa, and with large, setose endites from basis, ischium, and merus (Fig. 9D). Ischium shorter than merus in thoracic endopods 1 and 2, slightly longer in endopods 3 and 8, or distinctly longer in endopods 4–7. Paradactylar setae of endopods 3–8 weakly sickle-shaped, their armature with denticles is stronger in females (Fig. 10C) compared to males (Fig. 10J) and decreases successively in both sexes from endopods 3 to 8 (females Fig. 10C, E, G; males Fig. 10J–L). Penis as in Fig. 9H. Exopod of fourth male pleopod (Fig. 10Q): basal segment with strong, cuticularized ridge on its rostral face; this ridge more rugged than 20 · Zootaxa 4142 (1) © 2016 Magnolia Press WITTMANN ET AL. FIGURE 7. Paramysis (Longidentia) adriatica sp. nov. (A, B) and Troglomysis vjetrenicensis Stammer, 1933 (C). A, B, ethanol fixed adult female with body length 11 mm, from the Adige River, paratype, in dorsal (A) and in lateral (B) view (MZH reg. no. HLA.112100); C, ethanol fixed adult male 12.5 mm, from the type locality within the cave Donja Vjetrenica, lateral. A, B, photo M. Daneliya; C, photo A. P. Ariani. MYSIDAE IN MEDITERRANEAN FRESH-WATERS Zootaxa 4142 (1) © 2016 Magnolia Press · 21 FIGURE 8. Paramysis (Longidentia) adriatica sp. nov. from the Adige River, holotype, male with body length 9.1 mm (A), paratypes female 9.5 mm (B), male 7.9 mm (C–F, H, K), and female 10.9 mm (G, J). A, male habitus, lateral; B, anterior body region of female, dorsal; C, carapace expanded on slide, dorsal, details show cervical (D) and cardial (E) pore groups; F, antennula of male, dorsal; G, antennula of female, dorsal; H, antenna, ventral; J, labrum, ventral; K, maxillula, caudal. 22 · Zootaxa 4142 (1) © 2016 Magnolia Press WITTMANN ET AL. FIGURE 9. Paramysis (Longidentia) adriatica sp. nov. from the Adige River, paratypes, male with body length 7.9 mm (A–F, H), and female with 10.9 mm (G). A, mandibles, rostral; B, labium, caudal; C, maxilla, rostral; D, male thoracic sternites 1–8 (ventral) with left first thoracopod (caudal); E, lateral view on male sternal processes or humps in the sagittal plane; F, second thoracic endopod, caudal; G, sixth thoracic endopod with rudimentary oostegite, rostral; H, eighth thoracopod (caudal) with penis (postero-lateral blade of penis is accidentally shifted antero-laterally). MYSIDAE IN MEDITERRANEAN FRESH-WATERS Zootaxa 4142 (1) © 2016 Magnolia Press · 23 FIGURE 10. Paramysis (Longidentia) adriatica sp. nov. from the Adige River, paratypes, female with body length 10.9 mm (A–G, V), and males with 7.9 mm (H–P, R–U) or 9.1 mm (Q), respectively. A, female tarsus 3, i.e. tarsus of third thoracic endopod, rostral, details show a modified seta (B) of the propodus, and the rostral paradactylar seta (C); D, female tarsus 4, rostral, detail (E) as in (C); F, female tarsus 8, rostral, detail (G) as in (C, E); H, male tarsus 3, rostral, detail shows the rostral paradactylar seta (J); K, L, male rostral paradactylar setae of thoracic endopods 4 (K) and 8 (L); M–P, series of male pleopods 1–4, caudal (M, N) or rostral (O, P) view; Q, exopod of male pleopod 4 in lateral view, setae omitted; R, male pleopod 5, caudal; S, posterior margin of sixth pleonite, lateral; T, uropods, ventral; U, telson, dorsal; V, terminal portion of telson, dorsal. 24 · Zootaxa 4142 (1) © 2016 Magnolia Press WITTMANN ET AL. the homologous one found on the second segment in P. kosswigi (Fig. 5D); the second segment in P. adriatica (Fig. 10Q) with less strongly oblique segmental borders compared with the homologous third segment in P. kosswigi (Fig. 5D). Scutellum paracaudale subtriangular with rounded tip, margins sinusoid, lower margin inconspicuously undulate (Fig. 10S). Endopod of uropods is 77–84% length of exopod; spine size along inner margin of endopod and space between these spines increase, with some variations, distally (Fig. 10T). Exopod of uropod extends 38– 44% its length beyond telson. Telson trapezoidal (apart from cleft), its maximum width near basis is 2.1–2.7 times that at apex (Fig. 10U), its length is 1.9–2.5 times maximum width or 1.1–1.4 times length of sixth pleonite (measured along dorsal midline). Statoliths (Fig. 11A). The vaterite statoliths resemble ventrally flattened or truncated spheres, very similar to those of P. helleri (Fig. 4 in Ariani et al. 1993) and P. lacustris (Figs 4A, 6M in Wittmann et al. 1993); diameter 110–178 µm; statolith formula 2 + 3 + 1 + (5–9) + (11–16) = 22–31. Ambitus and tegmen with moruloid appearance, their surface resembling a wickerwork of lamellae. Such arrangements of lamellae often and so far exclusively found in vaterite statoliths. Ventrally the mantle does not cover the central part of the fundus, where a conical hole is typically visible (as in Fig. 11A). FIGURE 11. Vaterite (A) and fluorite (B) statoliths in Mediterranean freshwater mysids. A, Paramysis adriatica sp. nov. from small creek at Split, oblique ventro-lateral view on right statolith of adult female with body length 8.8 mm; B, Troglomysis vjetrenicensis from the cave Donja Vjetrenica (type locality), ventral view on right statolith of adult male 12.5 mm (statolith somewhat deteriorated upon treatments related to SEM study and X-ray diffraction analysis by Ariani et al. 1993). A, B, SEMmicrographs by A. P. Ariani. Colour. Not considering their less slender body and the larger eyes, freshly caught living specimens of the new species greatly resemble Diamysis mesohalobia mesohalobia Ariani & Wittmann, 2000 (colour photo in Ariani & Wittmann (2000): Fig. 2F) from Fiume Piccolo (see below). General appearance yellowish brown to dark brown depending on chromatophore expansion. Rich with chromatophores all over the body and appendages. The arborescent chromatophores give brown, yellowish, and to a lesser extent also rose markings. Mainly the brown components are preserved in the ethanol-fixed specimen shown in Fig. 7A, B. Comparison. The new species is most similar to P. agigensis known from brackish and marine waters of the Black, Marmora, and Aegean Seas, with which it shares a short terminal lobe of the antennal scale, heavily armed inner paradactylar seta on thoracic endopod 3, and the few spines on endopod of uropod. It shares scale-like hairs on rounded lobes of anterior (2–5) male thoracic sternites with P. intermedia, P. ullskyi, P. kessleri sarsi, and P. bakuensis (for lobe morphology in Paramysis see Wittmann & Ariani (2011)). In contrast, there are no hairs on the rounded lobes of P. helleri, P. festae, and P. nouveli, or on the less well-rounded lobes of P. kroyeri. There are also MYSIDAE IN MEDITERRANEAN FRESH-WATERS Zootaxa 4142 (1) © 2016 Magnolia Press · 25 smooth, but smaller, less distinct lobes in P. arenosa (G. O. Sars, 1877). The respective lobes are not rounded but subtriangular to sickle-shaped at least in large males of P. agigensis, P. pontica, and P. bacescoi. Additional differences of P. adriatica from its most similar (Ponto-)Mediterranean congenerics: from P. helleri by shallower telson cleft (15–24% telson length in the latter), stouter antennal scale (2.8–3.9 times as long as wide in P. helleri), strong, less numerous secondary spinules (about 20 thin distal setules in P. helleri) on the paradactylar setae in endopods 3–8, reminiscent of those of P. agigensis, and by fewer spines on the endopod of uropods (7–13 in P. helleri), also like in P. agigensis. The new species differs from P. festae by smaller eyes, a longer antennal scale, and a greater number of laminae in the telson cleft. It differs from P. agigensis by a shallower telson cleft (15–19% telson length in P. agigensis) not or barely reaching up to the penultimate lateral spines (almost reaching the antepenultimate spines in P. agigensis), by slightly stouter antennal scale (2.8–3.0 times as long as wide in P. agigensis), but slightly longer apical part of the antennal scale (16–18% scale length in P. agigensis), and by the presence of 4–6 short, barbed setae on caudolateral blade of penis (1–3 long, smooth setae in P. agigensis). Pleopods of two adult males each from shallow brackish waters at sandy beaches were dissected for P. helleri from the North Adriatic Sea (Spiaggia die Ponente, Caorle, Italy, 45.5814N 012.8513E, S = 17, leg. KJW) and P. agigensis from the Sea of Marmora (Büyükada Islands, Turkey, 40.8594N 029.1126E, again S = 17, leg. KJW). The fourth pleopod of both species differed from that in P. adriatica by two-segmented (versus unsegmented) endopod and 7-segmented (versus 6-segmented) exopod, the latter showing a weaker, less strongly ridge-like differentiation at its upper surface near the end of its second segment (corresponding to the basal segment in P. adriatica). FIGURE 12. Distribution of five species of Mysinae (Mysidae) in tributaries and coastal waters of the Adriatic Sea. In part modified from Wittmann & Ariani (2012b). 26 · Zootaxa 4142 (1) © 2016 Magnolia Press WITTMANN ET AL. Tribe Diamysini Wittmann, Ariani & Lagardère, 2014 Figs 13–15, 17–19 Diagnosis (extended from Wittmann et al. 2014). Mysinae with antennal scale setose all around, scale mostly with short apical segment (Fig. 18D); first thoracic endopod 7-segmented, with distinct endites on coxa, basis, and ischium, and indistinct endite up to a strong one on merus; ischium always well developed, its inner margin (but not always its outer one) more than half the length of that of merus; carpopropodus of thoracic endopod 6 with 2–3 segments (Fig. 18H); well-developed oostegites on thoracopods 7 and 8, rudimentary oostegite on thoracic endopod 6; pleopods rudimentary in both sexes, except male pleopods 3 and 4; male pleopod 3 reduced to small endopod, mostly fused with larger 2-segmented sympod (Fig. 13C), less frequently rudimentary as in female; pleopod 4 with large, 2-segmented sympod, its exopod longer, at least 2-segmented, frequently styliform, ending in 1 (2–3) large, modified setae, its endopod reduced to a small, 1- or 2-segmented rod often subbasally with outwards directed apophysis (Fig. 19H); endopod of uropod with 1 (0–2) spines on ventral face near statocyst (Fig. 19L), statoliths composed of vaterite, less frequently of fluorite; telson shorter than ultimate pleonite (Fig. 14A), mostly with short apical cleft (Fig. 17J) or at least apically truncate, spines (if present) on lateral margins arranged in continuous series, not arranged in groups of large spines with smaller spines in between; each lateral margin ending in a comparatively large spine, terminal margin including its cleft (if distinct) lined with spine-like laminae (Fig. 19M) or with small spines. Type genus. Diamysis Czerniavsky, 1882. Taxa included. Nine genera: Antromysis Creaser, 1936 (6 species), Diamysis (14), Gangemysis Derzhavin, 1924 (1), Indomysis W. M. Tattersall, 1914 (2), Limnomysis Czerniavsky, 1882 (1), Parvimysis Brattegard, 1969 (3), Surinamysis Bowman, 1977 (3), Taphromysis Banner, 1953 (3), Troglomysis Stammer, 1933 (1). Distribution. Tropical to temperate waters of the Atlantic, Mediterranean, Pontocaspian, and Indian Ocean, absent in the Pacific, Arctic, and Antarctic. In coastal to continental waters, mostly in brackish waters, also marine, showing a comparatively high frequency of freshwater and subterranean species. Genus Diamysis Czerniavsky, 1882 Figs 13–15, 17 Short diagnosis (modified from Wittmann & Ariani (2012a)). Diamysini with eyes normal. Antennal scale setose all around, with small apical segment (<= 20% scale length) surrounded by five plumose setae in both sexes (Figs 13A, D, E, 14A, 15A, K, 17A, K). Carapace with a pair of post-suborbital spines (Figs 13A, B, D, F, 14A, C, 15K, L, 17A). Carapace of females always without fringes, whereas potential presence of fringes (Figs 13B, 15L) is species-specific in males. Thoracic endopods 3–8 normal, with 2–4-segmented carpopropodus, dactylus small, with distinct claw (Figs 14E, 15B, M, 17D, L). All female pleopods and male pleopods 1, 2, 5 reduced to setose rods. Male pleopod 3 reduced to well-developed, 2-segmented sympod terminally fused with its small, setose, unsegmented endopod, exopod missing (Figs 13C, 14G). Male pleopod 4 biramous with 2-segmented sympod, with small, 1–2-segmented endopod, and with moderately long, rod-like, 2–3-segmented exopod bearing a modified, strong seta at tip (Figs 13G, 14H, 15D, E, N, 17E, N); the apical seta is much longer than any other seta on male pleopod 4. Endopod of uropod setose all around, with one (exceptionally two) spines below statocyst. Telson with superficial to deep apical incision; with spines on each lateral margin, these margins ending in a pair of larger, posteriorly directed apical spines; cleft lined by many laminae (Figs 13H, 14N, 15J, S, 17J, Q). Type species. Mysis bahirensis G. O. Sars, 1877, by subjective monotypy upon definition of the genus Diamysis by Czerniavsky (1882a, 1887: 84). Taxonomy. Interbreeding experiments by Ariani & Wittmann (2000) indicate failure of mutual interbreeding between the topotypical population of Diamysis bahirensis (G. O. Sars, 1877) from the El Bahira lagoon at the coast of Tunisia and a number of other Mediterranean populations of its genus. This and morphological differences point to a specific status of D. mesohalobia Ariani & Wittmann, 2000, and D. lagunaris Ariani & Wittmann, 2000. Distribution. Tropical to temperate waters of the E-Atlantic, Mediterranean, Pontocaspian, and W-Indian Ocean. Greatest species diversity in the Mediterranean (Figs 6, 12, 16). In coastal to continental waters, mostly in brackish waters, also marine, comparatively high diversity in fresh-water. Among the 14 species plus two nonnominotypical subspecies so far known, Diamysis pengoi (Czerniavsky, 1882), D. lacustris Băcescu, 1940, and D. MYSIDAE IN MEDITERRANEAN FRESH-WATERS Zootaxa 4142 (1) © 2016 Magnolia Press · 27 fluviatilis Wittmann & Ariani, 2012, are mainly found in fresh-water; whereas D. mesohalobia heterandra Ariani & Wittmann, 2000, is an essentially brackish-water taxon (Fig. 16), also found in fresh-water (Fig. 12). Bionomy. In D. lagunaris and in two subspecies of D. mesohalobia, the survival of brood pouch larvae was higher under mesohaline than euhaline conditions in the laboratory (Ariani & Wittmann 2000). Based on this, along with biomineralogical and morphological similarities of the statoliths with fossil ones from Miocene deposits of the brackish Paratethys, Ariani & Wittmann (2000) suggested a brackish-water origin of Diamysis, even of (mixo)euhalobious forms that may have returned to the sea from low-salinity environments. Remarks. Exhaustive data on D. lagunaris, D. lacustris, D. mesohalobia heterandra, and D. fluviatilis, are available in Wittmann & Ariani (2012a, 2012b). Data necessary for determination are given in Figs 13–15, 17. Additional remarks are in the ‘Discussion’. Diamysis fluviatilis Wittmann & Ariani, 2012 Fig. 13A–C Mysis oculata relicta: Stammer 1932 (partim). Diamysis bahirensis: Holmquist 1955; Minelli & Trevisanello 1985. Diamysis bahirensis ssp.: Ariani 1981b (partim). Diamysis mesohalobia: Cibinetto et al. 2006. Diamysis mesohalobia heterandra: Corazza et al. 2010. Diamysis fluviatilis Wittmann & Ariani 2012b: Mees 2014; Wittmann et al. 2014. Material examined. 23 samples from freshwater tributaries of the North Adriatic Sea, see Wittmann & Ariani (2012b). Additional materials are from the lower Timavo system, 45.79N 013.58E, Gulf of Trieste, NE-Adriatic: 4 slides labelled "Diamysis biharensis. Herzegovina & Timavo. Det. Ch. Holmquist, prep. 60–63"; the data given by Stammer (1932) and Holmquist (1955: Tab. 1) suggest that this material was collected in 1928/29 by Hans Jürgen Stammer in waters of the Timavo system: prep. nos. 60, 61 (SMNH reg. nos. 140111, 140112) are dissected parts of 1 M ad. 8.5 mm and of 1 F ad. 9.5 mm with nauplioid larvae from former fresh-waters in the meadows (now industrial port zone) between the Timavo estuary and the Monfalcone creek, 45.789N 013.581E; prep. nos. 62, 63 (SMNH reg. nos. 140113, 140114) are parts of 1 M ad. 8.5 mm and of 1 F ad. 8.5 mm with nauplioid larvae from still existing springs of the Monfalcone creek, 45.7984N 013.5701E. Diagnosis. Rostrum well rounded or forming a wide convex angle with broadly rounded tip (Fig. 13A). Male carapace with fringes arranged in two submedian stripes (Fig. 13B), no such fringes in females. Palpus of maxilla with subcircular terminal segment, armed with 9–19 denticles along distal margin. Basal segment of thoracic exopods 1–8 in males and exopods 1–3 in females with spiniform outer corner (as in Fig. 14D); this corner variable in exopods 4–8 of females. Thoracic endopod 3 with 3-segmented carpopropodus being longer than 5 times its maximum width; thoracic endopods 3–8 with long and slender claw. Penis with smooth setae only. Male pleopod 4 biramous with 2-segmented sympod, with small, 1–2-segmented endopod, and with moderately long, rod-like, 2segmented exopod bearing a modified, strong seta at tip and a smaller, smooth seta subterminally on the basal segment (as in Fig. 13G). Scutellum paracaudale well rounded to apically pointed, margins not or inconspicuously undulate. Endopod of uropod with one strong spine below large statocyst; statolith composed of vaterite. Telson subquadrangular, 0.7–0.9 times length of last abdominal somite; its apical cleft with slightly to strongly convex margins. Bottom of cleft concave, rounded. Cleft is 9–17% telson length, cleft lined by 15–48 laminae. Distribution (Fig. 12). In wells, springs, tributaries, canals, main courses, and estuaries of rivers flowing to the NE to NW coasts of the Adriatic Sea (Reno, Po, Mincio, Adige, Brenta, Sile, Livenza, Lemene, and Isonzo), including the Timavo system. Maximum observed sea distance 182 km, maximum altitude 16 m. Mostly in freshwater, also oligohaline; only in the lower Timavo estuary observed down to the mesohaline reach (S = 2–13). Diamysis lacustris Băcescu, 1940 Fig. 13D–H Diamysis bahirensis: Spandl 1926; Holmquist 1955 (partim). Diamysis bahirensis (?) mod. lacustris Băcescu 1940 (in footnote on p. 578). 28 · Zootaxa 4142 (1) © 2016 Magnolia Press WITTMANN ET AL. Diamysis pengoi: Genovese 1956 (partim); Komarova 1991 (partim); Daneliya & Petryashev 2011 (partim). Diamysis bahirensis mod. lacustris: Ariani 1981b; Wittmann & Stagl 1996. Diamysis lacustris: Wittmann & Ariani 2012b; Mees 2014. Material examined. 3 samples from freshwater Lake Scutari (= Skadarsko jezero = Liqen i Shkodres), 42.23N 019.16E, see Wittmann & Ariani (2012b). Diagnosis. Rostrum angular with narrowly rounded tip (Fig. 13E). Carapace without fringes in both sexes (Fig. 13D, F). Palpus of maxilla with subcircular terminal segment, armed with 10–16 denticles along distal margin. Males with spiniform outer corner of the basal segment in thoracic exopods 1–7 (Fig. 13D), this corner variable in exopod 8; females with this corner spiniform in exopod 1, subrectangular in exopods 2–7, or rounded in exopod 8. Thoracic endopod 3 with 3–2-segmented carpopropodus being longer than 5 times its maximum width (Fig. 13D); thoracic endopods 3–8 with long and slender claw. Penis with predominantly barbed setae on anterior face and in a semicircle close to the ejaculatory opening. Male pleopod 4 biramous with 2-segmented sympod, with small, 2-segmented endopod, and with moderately long, rod-like, 2-segmented exopod bearing a modified, strong seta at tip and a smaller, smooth seta subterminally on basal segment (Fig. 13G). Scutellum paracaudale triangular (Fig. 13D), with undulate margins ending in an acute tip. Endopod of uropod with one strong spine below large statocyst; statolith composed of vaterite. Telson subquadrangular (Fig. 13H), 0.7–0.8 times length of last abdominal somite; its apical cleft very shallow, only 7–12% telson length, cleft with almost straight margins forming an angle of 110–150°, cleft lined by 16–23 laminae. Occurrence (Fig. 12). Only known from the freshwater Lake Scutari at the SE-coast of the Adriatic Sea, altitude 5 m. The minimum distance of the lake to the sea is 43 km along its effluent or only 13 km as the crow flies. Diamysis mesohalobia Ariani & Wittmann, 2000 Figs 14, 15 Diagnosis (sensu lato: covering the three currently known subspecies). Appendix masculina 80–120% the length of terminal segment of antennular peduncle (Fig. 14B). Eyes normal; eyestalks with fenestra paracornealis weakly developed (Fig. 15K) or absent (Fig. 15A), in any case mostly not or poorly visible. Distal segment of maxillary palpus with 4–27 distinct denticles. Presence of fringes on male carapace varies between subspecies; no such fringes in females. Basal segment of thoracic exopods with outer corner spiniform or less frequently ending in an acute edge, occasionally rounded in the first and/or in some of the median and/or posterior exopods. All pereiopods with normal carpopropodus and slender, styliform claw (Figs 14E, 15B, M). Carpopropodus of thoracic endopods 3–8 with 3 (2; 4), 3–2 (4), 3–2, 2–3, 2–3 and 3–2 segments, respectively. Carpopropodus of endopod 3, if 3segmented, with basal segment not longer than remaining segments combined (Fig. 15M). Thoracic endopod 3 and often also endopod 8 with at least one among the four paradactylar setae distally pectinate in (most) females, smooth or pectinate in males. Male pleopod 4 biramous with 2-segmented sympod and with small, 2-segmented endopod (Figs 14H, 15N); its exopod 2–(3)-segmented, with large modified seta at tip. This exopod with basal segment bearing a smaller smooth seta (Figs 14H, 15D), in certain populations occasionally with an additional barbed seta (Fig. 15N). Endopod of uropod with one strong spine below statocyst, statolith composed of vaterite. Telson subquadrangular (Fig. 15S) to subtriangular (Fig. 15J); maximum width is 1.4–3.0 times that at apex; its apical cleft with 8–39 laminae; cleft is 5–26% telson length. Taxonomy. Interbreeding experiments by Ariani & Wittmann (2000) indicated mutual crossability between morphologically different Mediterranean populations of D. mesohalobia. The three main morphotypes distinguished were, therefore, described at subspecific level as D. mesohalobia mesohalobia Ariani & Wittmann, 2000, D. mesohalobia gracilipes Ariani & Wittmann, 2000, and D. mesohalobia heterandra Ariani & Wittmann, 2000; each of these subspecies are treated in separate subchapters below. Occurrence (Fig. 16). Marine, brackish and fresh (near)-coastal waters of the E-Mediterranean and Marmora Seas. MYSIDAE IN MEDITERRANEAN FRESH-WATERS Zootaxa 4142 (1) © 2016 Magnolia Press · 29 FIGURE 13. Diamysis fluviatilis Wittmann & Ariani, 2012 (A–C), and D. lacustris Băcescu, 1940 (D–H). Materials from the Sile River (A–C) and Lake Scutari (D–H). A, cephalic region of female with body length 8.9 mm, dorsal; B, carapace expanded on slide, male 9.5 mm, dorsal; C, third pleopod of male 6.6 mm, rostral aspect; D, male with body length 5.2 mm, lateral view; E, cephalic region of female 8.0 mm, dorsal; F, carapace expanded on slide, dorsal, same male as in (D); G, fourth pleopod of same male, rostral; H, telson of same male, dorsal. B, F, pores on carapaces not to scale. From Wittmann & Ariani (2012b). 30 · Zootaxa 4142 (1) © 2016 Magnolia Press WITTMANN ET AL. FIGURE 14. Diamysis mesohalobia mesohalobia Ariani & Wittmann, 2000, paratypes from the brackish spring Fiume Morello at the Adriatic coast of SE-Italy. A, dorsal aspect of male with body length 6.5 mm; B, antennula of male 6.3 mm, dorsal; C, carapace expanded on slide, male 6.0 mm, dorsal; D, exopod of first thoracopod, male 6.3 mm, caudal aspect; E, tarsus (i.e. carpopropodus plus dactylus) of fourth thoracic endopod, male 6.4 mm, rostral; F, right face of left penis, male 6.3 mm; G, third pleopod of same male, rostral; H, fourth pleopod of same male, rostral; J–M, posterior margin of sixth pleonite, lateral, in female 6.6 mm (J), female 6.8 mm (K), and two males, each with 6.3 mm (L, M); N, telson of male 6.3 mm, dorsal. A, C, pores on carapaces not to scale. A, B, D, E, G–N, modified from Ariani & Wittmann (2000); C, F, original. MYSIDAE IN MEDITERRANEAN FRESH-WATERS Zootaxa 4142 (1) © 2016 Magnolia Press · 31 Diamysis mesohalobia mesohalobia Ariani & Wittmann, 2000 Fig. 14 Diamysis bahirensis: W. M. Tattersall 1927; Ariani 1966 (partim), 1979 (partim), 1981a (partim); Ariani et al. 1981, 1982, 1983, 1993 (partim); De Matthaeis et al. 1982 (partim); Wittmann et al. 1990, 1993 (partim); Schlacher et al. 1992. Diamysis bahirensis ssp.: Ariani 1981b (partim). Diamysis sp.: Wittmann & Stagl, 1996 (partim); Ariani et al. 1999. Diamysis sp. A: Wittmann 1999. Diamysis mesohalobia Ariani & Wittmann, 2000: 2002, 2004; Ariani et al. 2000; Petrescu & Wittmann 2009; San Vicente 2010; Daneliya & Petryashev 2011; ITIS 2014. Diamysis mesohalobia mesohalobia Ariani & Wittmann, 2000: 2005; Kocataş et al. 2003; Ariani 2004; Özbek et al. 2004; Remerie et al. 2004; Özbek & Ustaoğlu 2006; Anderson 2008; Wittmann & Ariani 2010; Mees 2014. Material examined. 84 samples from mesohaline to mixoeuhaline coastal waters of the Adriatic, Aegean and Levantine Seas (Ariani & Wittmann 2000). Previously unpublished sample: 93 F ad. 5.5–7.1 mm, 223 M ad. 4.3– 6.3 mm, 22 subad., 56 imm., 112 juv., karstic spring with small salinity fluctuations, Fiume Piccolo, near Torre Canne, Adriatic coast of southern Italy, 40.8275N 017.4818E, 2–3 m depth, striped with hand net from brown algae, S = 15, pH 7, 18.5°C, 29 Nov. 2011, leg. A. P. Ariani. One specimen of this sample reminds of a gynandromorph of the 'fore and aft' type (Hollingsworth 1960): with pleopods as typical for adult males, but antennula without appendix masculina and also without plumose setae typical of males or females. Short updated description. The following data cover primarily the type population in the mesohaline spring Fiume Morello (Apulia, Adriatic Sea). Data from remaining populations, as far as different, are given in square brackets. Diamysis mesohalobia with short rostrum forming a wide convex angle with rounded tip (Figs 14A, C). Fenestra paracornealis poorly developed, rarely visible. Carapace without fringes (Fig. 14C) in both sexes. Palpus of maxilla with subcircular terminal segment, armed with 6–24 [5–24] denticles along distal margin. Basal segment of thoracic exopods with outer corner spiniform (Fig. 14D), less frequently ending in an acute or rounded edge, especially in posterior exopods. Pereiopods relatively short, endopod 8, when stretched anteriorly, extending to basis of endopod 1 or at most up to maxillae [mandibles]. Pereiopods stout (Fig. 14E) to moderately slender, with R6 = 4.4–6.1 [4.4–6.8]. Carpopropodus of thoracic endopods 3–8 with 3 (2), 3 (2), 3–2, 2–3, 2–3, and 3 (2) [3–2] segments, respectively. Thoracic endopod 3 with carpopropodus being longer than 5 times its maximum width; thoracic endopods 3–8 with long and slender claw. Penis with smooth setae only, arranged in a semicircle close to ejaculatory opening (Fig. 14F). Male pleopod 4 biramous with 2-segmented exopod bearing a modified, strong seta at tip and a smaller, smooth seta subterminally on basal segment (Fig. 14H). Scutellum paracaudale subtriangular, biconvex; tip pointed or less frequently rounded (Fig. 14J–M). Telson subquadrangular (Fig. 14N) [to subtriangular], 0.8–0.9 [0.7–1.0] times length of last abdominal somite; maximum width of telson is 1.6–2.0 [1.6– 3.0] times that at apex; lateral margins concave or rarely straight, armed with 7–12 [7–14] spines. Apical cleft of telson with convex (Fig. 14N) or rarely straight margins, bottom of cleft rounded, cleft is 12–16% [9–16%] telson length, cleft lined by 15–39 [8–39] laminae. Body length: Adult females 4.6–9.6 mm, males 4.1–7.7 mm. Distribution (Fig. 16). Eastern Mediterranean only: in Adriatic, Aegean, and Levantine Seas. Mostly in mesohaline karstic springs with small salinity fluctuations, also in mesohaline to mixoeuhaline lagoons and estuaries. Predominantly in the salinity range 10–38, locally down to S = 2. Samples of this subspecies were taken by Özbek et al. (2004) in the Köyceğiz lagoon at the Aegean coast of Turkey. Data by Akin et al. (2005) and own measurements at the northern shore (S = 2.3 in 0.5–2 m, 10 June 2006) suggest that the positive station was most likely from the oligohaline range within this large, oligo- to metahaline lagoon with complex salinity patterns. Diamysis mesohalobia gracilipes Ariani & Wittmann, 2000 Figs 15A–J Diamysis bahirensis: Ariani 1966 (partim), 1979 (partim), 1981a (partim); De Matthaeis et al. 1982 (partim); Wittmann 1992; Ariani et al. 1993 (partim). Diamysis sp.: Wittmann & Stagl, 1996 (partim). 32 · Zootaxa 4142 (1) © 2016 Magnolia Press WITTMANN ET AL. Diamysis sp. A ssp.: Wittmann, 1999. Diamysis mesohalobia gracilipes Ariani & Wittmann, 2000: 2004, 2005; Anderson 2008; Corazza et al. 2010; Wittmann & Ariani 2010; San Vicente 2010; Mees 2014. Material examined. 52 samples from brackish and marine waters of the Ionian and Adriatic Seas (Ariani & Wittmann 2000). Among these 52 positive samples, eight are from a karstic spring with constant salinity in the oligohaline range (S = 4), Fiume Chidro in the Gulf of Tarent, Ionian Sea, 40.3052N 017.6815E. Previously unpublished samples: 58 F ad. 4.8–5.5 mm, 14 M ad. 4.1–4.8 mm, 6 subad., 19 imm., 49 juv.; from type locality, mixoeuhaline embayment with lagoonal aspect, Mar Piccolo di Taranto, Gulf of Tarent, Ionian coast of Apulia, SE-Italy, 40.4689N 017.2982E, S = 38.5, 27°C in 0.5 m depth, 0.5–1.5 m, from sand and stones with Ulva, Enteromorpha, and Ceramium, 27 Aug. 2005, hand net, leg. Ariani & Wittmann, NHMW reg. no. 25705; 79 F ad. 4.5–5.1 mm, 82 M ad. 3.7–4.7 mm, 63 subad., 356 imm., 284 juv.; same data as before, 2–3 m depth, from mud and sand, a few Chaetomorpha, NHMW 25706; 1 F ad. 6.9 mm, 1 F subad. 5.8 mm, from the above-cited karstic spring Fiume Chidro, S = 4, 17.3°, 0.5–1 m, 29 Feb. 2012, hand net, leg. A. P. Ariani. Short updated description. The following data cover primarily the type population in the mixoeuhaline lagoon Mar Piccolo di Taranto (Gulf of Tarent, Ionian Sea). Data from remaining populations, as far as different, are given in square brackets. Diamysis mesohalobia with short rostrum forming a wide convex angle with rounded tip (Fig. 15A). Fenestra paracornealis feebly developed, often visible in specimens with expanded eyestalk pigment [not visible in certain populations]. Carapace without fringes in both sexes (Fig. 15A). Palpus of maxilla with subcircular terminal segment, armed with 7–19 [4–23] denticles along distal margin. Basal segment of thoracic exopods with outer corner spiniform at least in intermediate exopods, or rounded, especially in posterior exopods, partly also first endopod. Pereiopods long, thoracic endopod 8, when stretched anteriorly, extending up to labrum or even to sympod of antennae [or up to eyes]. Pereiopods slender, with R6 = 5.6–8.1 [4.9–9.1]. Carpopropodus of thoracic endopods 3–8 with 3 (2; 4), 3–2, 2, 2, 2, and 3–2 segments, respectively [3 (2), 3–2 (4), 2–3, 2 (3), 2–3, and 3–2]. Thoracic endopod 3 with carpopropodus longer than 5 times its maximum width (Fig. 15B); thoracic endopods 3– 8 with long and slender claw. Penis with smooth setae arranged in a semicircle close to ejaculatory opening; in the type population with 0–3 additional barbed setae on anterior face (Fig. 15C) [barbed setae less frequent, rare, or not found elsewhere]. Exopod of fourth male pleopod 2-segmented in small males (4–5 mm; Fig. 15D), with smooth seta at basal segment [and occasionally an additional small barbed seta]; or mostly 3-segmented in larger males (Fig. 15E), with distally barbed seta at median segment; distal segment in any case with a modified, strong seta at tip (Fig. 15D, E). Scutellum paracaudale biconvex [well-rounded, subtriangular; occasionally bifid (Fig. 15F) or rarely with subterminal suture (Fig. 15G)]; tip pointed (Fig. 15F–H) or less frequently rounded. Telson subquadrangular to subtriangular (Fig. 15J), 0.7–0.9 [0.8–1.0] times length of last abdominal somite; maximum width of telson is 1.9–2.7 times that at apex; lateral margins concave or occasionally straight, armed with 7–13 [7– 12] spines. Apical cleft of telson with straight or slightly convex margins, bottom of cleft rounded, cleft is 9–20% [5–21%] telson length, cleft lined by 9–24 [11–28] laminae (Fig. 15J). Body length. Adult females 4.6–8.5 mm, males 4.1–7.3 mm. Distribution (Fig. 16). Eastern Mediterranean only: in Ionian and Adriatic Seas. Marine coastal and in polyhaline to mixoeuhaline lagoons, also in estuaries and karstic springs. Salinity range 4–38, mostly in polyhaline to mixoeuhaline waters with small tidal variations; in marine environments with notable annual salinity fluctuations (S = 30–38 in the Gulf of Trieste). The karstic spring Fiume Chidro represents the only oligohaline station so far known; here the subspecies has always been found at constant salinity for more than three decades (S = 4; inspected 1980–2012). Diamysis mesohalobia heterandra Ariani & Wittmann, 2000 Fig. 15K–S Mysis oculata var. relicta: Zimmer 1927 (partim: Lake Deran). Diamysis bahirensis: Holmquist 1955; Avĉin et al. 1973 (partim); Matjašič & Štirn 1975 (partim); Ariani et al. 1983, 1993 (partim). Diamysis bahirensis ssp.: Ariani 1981b (partim). MYSIDAE IN MEDITERRANEAN FRESH-WATERS Zootaxa 4142 (1) © 2016 Magnolia Press · 33 FIGURE 15. Diamysis mesohalobia gracilipes Ariani & Wittmann, 2000 (A–J), and Diamysis mesohalobia heterandra Ariani & Wittmann, 2000 (K–S). Materials from Mar Piccolo in the Gulf of Tarent (paratypes: A–E, G–J, Ionian Sea), the Bay of Strunjan (F, Gulf of Trieste, NE-Adriatic Sea), Lake Deran (L, N, P–S, E-Adriatic), Limni Antinioti (K, M, Island of Corfu, Ionian Sea), and Lakes of Bacin (O, E-Adriatic). A, anterior body region of male with body length 7.2 mm, dorsal view on cephalothorax; B, tarsus (i.e. carpopropodus plus dactylus) of third thoracopod, male 6.5 mm, rostral; C, left face of right penis, male 7.2 mm; D, E, exopod of fourth pleopod, rostral, in males with 4.4 mm (D) or 6.5 mm (E) body length; F–H, posterior margins of sixth pleonite in female with 7.4 mm (F), in male with 7.2 mm (G), and in female with 4.8 mm (H) body length, lateral; J, telson of male 7.2 mm, dorsal; K, anterior body region of male 5.2 mm (paratype); L, carapace expanded on slide, dorsal, male 7.9 mm; M, tarsus of third thoracic endopod, rostral, male 5.2 mm (paratype); N, fourth pleopod of male 7.9 mm, rostral; O, P, exopod of fourth pleopod, rostral, in males with 5.5 mm (O) or 8.4 mm (P) body length; Q, R, posterior margins of sixth pleonite in female with 9.8 mm (Q) or in male with 6.0 mm (R) body length, lateral; S, telson of male 7.9 mm, dorsal. A, K, L, pores on carapaces not to scale. A, D, E, G–K, M, modified from Ariani & Wittmann (2000); L, N–S, modified from Wittmann & Ariani (2012b); B, C, F, original. 34 · Zootaxa 4142 (1) © 2016 Magnolia Press WITTMANN ET AL. Diamysis sp.: Wittmann & Stagl 1996 (partim). Diamysis mesohalobia heterandra Ariani & Wittmann, 2000: 2004, 2005; Anderson 2008; San Vicente 2010; Wittmann & Ariani 2010, 2012a, 2012b; Mees 2014. Material examined. 42 samples from oligohaline to metahaline lagoons and karstic springs in diverse parts of the eastern Mediterranean plus 13 samples from freshwater tributaries of the Adriatic Sea, see Ariani & Wittmann (2000) and Wittmann & Ariani (2012b); material studied by Holmquist (1955), previously unpublished reexamination: dissected parts of 1 M ad. 8 mm, 2 egged F ad. with body length 7 or 9 mm, respectively, on a total of 3 slides labelled "Diamysis biharensis. Herzegovina & Timavo, det. Ch. Holmquist, prep. 57–59", SMNH reg. nos. 140108–140110. According to Holmquist (1955) this material was collected by Janez Hoenigman in Lake Deran, 2–5 m depth, 27 Apr. 1954, 43.04N 017.75E, Herzegovina, altitude 0 m, sea distance 31 km as calculated along the small effluent and following this along the Neretva River to the east coast of the Adriatic Sea. Short updated description. The following data covers primarily the type population in an oligo- to mixoeuhaline lagoon with brackish spring, Limni Antinioti (Island of Corfu, Ionian Sea). Data from remaining populations, as far as different, are given in square brackets. FIGURE 16. Distribution of Limnomysis benedeni Czerniavsky, 1882, Neomysis integer (Leach, 1814), and the three subspecies of Diamysis mesohalobia Ariani & Wittmann, 2000, in tributaries and coastal waters of the Mediterranean and adjacent seas. Data original and from 79 literature sources. Diamysis mesohalobia with short rostrum mostly forming a wide convex angle with broadly rounded tip (Fig. 15K, L). Fenestra paracornealis weakly developed, mostly visible (Fig. 15K) in well preserved material [mostly visible in well preserved material from Lake Deran (near E-Adriatic coast) or rarely from Schiavetti Springs (Gulf of Trieste, N-Adriatic)]. Carapace of adult males with fringes arranged in two submedian stripes plus one subterminal stripe (Fig. 15K, L). The submedian stripes may be differentiated as two separate stripes each (Fig. 15K). Palpus of maxilla with subcircular terminal segment, armed with 8–27 denticles along distal margin. Basal segment of all thoracic exopods normally with spiniform outer corner, rounded only in some of the posterior exopods of small individuals (< 6 mm). Pereiopods of intermediate length, endopod 8, when stretched anteriorly, extending to maxillae or at most to labrum. Pereiopods stout to slender, with R6 = 4.5–6.7 [4.5–7.9]. Carpopropodus of thoracic endopods 3–8 with 3, 3 (2), 2–3, 2 (3), 2, and 3–2 segments, respectively [3, 3–2 (4), 2– 3, 2–3, 2–3, and 3–2]. Thoracic endopod 3 with carpopropodus longer than 5 times its maximum width (Fig. 15M); thoracic endopods 3–8 with long and slender claw. Penis with smooth setae only, arranged in a semicircle close to MYSIDAE IN MEDITERRANEAN FRESH-WATERS Zootaxa 4142 (1) © 2016 Magnolia Press · 35 ejaculatory opening. Male pleopod 4 biramous with 2-segmented exopod; apical segment with a modified, strong seta at tip and in large males (> 6 mm) often with a minute additional seta; basal segment subterminally with a smooth seta (Fig. 15N–P). Large males (> 7 mm) with 0–1 [0–4] additional small barbed [and/or smooth] seta (Fig. 15N, P) on terminal margin of basal segment of exopod. Scutellum paracaudale subtriangular, mostly biconvex [or with upper margin convex and lower margin concave]; tip pointed (Fig. 15Q, R) or less frequently rounded, rarely bifid. These margins mainly smooth in small specimens [or undulate in large ones (> 8 mm; Fig. 15Q)]. Telson mostly subquadrangular (Fig. 15S), but subtriangular in small specimens (< 6 mm), 0.7–0.9 [0.7–1.0] times length of last abdominal somite; lateral margins concave [to straight], armed with 8–13 [6–11] spines; maximum width of telson is 1.8–2.4 [1.4–2.4] times that at apex; its apical cleft with straight to strongly convex margins. Bottom of cleft angular to rounded. Cleft is 10–19% [10–26%] telson length, cleft lined by 12–31 [9–38] laminae (Fig. 15S). Body length. Adult females 3.7–9.7 mm, males 3.0–8.7 mm. Distribution (Figs 12, 16). In fresh and brackish waters of springs, estuaries, lagoons, and lakes all around the Adriatic Sea (Fig. 12), salinity range S = 0–42. Outside the Adriatic (Fig. 16) known only from oligo- to polyhaline waters on the coasts of the Ionian and Marmora Seas, so far not from fresh-water (Ariani & Wittmann 2000, Wittmann & Ariani 2012a, b). Diamysis lagunaris Ariani & Wittmann, 2000 Fig. 17A–J Mysis bahirensis G. O. Sars, 1877 (partim: material from La Spezia): Gourret 1897; Sudry 1910. Diamysis bahirensis: Băcescu 1941; Genovese 1956; Drake et al. 1997; Cunha et al. 2000; San Vicente & Munilla 2000; Goulletquer et al. 2002; Munilla & San Vicente 2005. Diamysis bahirensis ssp.: Ariani 1979 (partim: material from Lake Ganzirri). Diamysis sp. B: Wittmann 1999. Diamysis sp.: Wittmann & Ariani 2000. Diamysis lagunaris Ariani & Wittmann, 2000: 2004, 2005; Ariani 2004; Anderson 2008; Petrescu & Wittmann 2009; Petryashov 2009; Wittmann & Ariani 2009, 2010, 2012a; San Vicente 2010; ITIS 2014; Mees 2014; Wittmann et al. 2014. Material examined. Two samples from marine waters of the eastern Mediterranean, 32 samples from brackish and marine waters of the western Mediterranean, plus 3 from the E-Atlantic (Portugal): see Ariani & Wittmann (2000), Wittmann & Ariani (2012a). Among these 37 positive samples only one from the oligohaline reach (S = 3.4): 1 M subad. 4.7 mm from the Mediterranean coast of France, Canal d'Arles à Fos, 43.4663N 004.8338E; previously unpublished sample: 2 M ad. 5.4–5.6 mm, 1 F ad. 6.7 mm, among ~30,000 Mesopodopsis slabberi and 2 Limnomysis benedeni, Mediterranean coast of France, estuary of the Petit Rhône at Tiki, same sample as indicated above for M. slabberi, NHMW reg. no. 25707. Diagnosis (sensu lato: covering the known population range). Eyes normal, eyestalks dorsally with welldeveloped fenestra paracornealis (Fig. 17B), although not well visible in poorly pigmented eyestalks. Rostrum forms a wide convex angle with broadly rounded tip (Fig. 17A, B). Carapace without fringes in both sexes (Fig. 17A). Palpus of maxilla with distal segment subcircular, armed with 5–25 distinct denticles. Pereiopods of moderate length, eighth endopod extending to the maxillae or at most up to mandibles. All pereiopods with normal carpopropodus and slender, styliform claw (Fig. 17D). Basal segment of thoracic exopods with outer corner spiniform (Fig. 17C) or occasionally rounded in some of the posterior exopods, most often rounded in last exopod. Pereiopods poorly to markedly slender, with R6 = 4.8–8.1 (Fig. 17D). Carpopropodus of thoracic endopods 3–8 with 3–2 (4), 2–3, 2, 2, 2, and 2–3 segments, respectively; tarsus slender, with slender, in part feebly serrated claw; carpopropodus 3 longer than 5 times its maximum width (Fig. 17D). Exopod of fourth male pleopod 2-segmented with a large modified seta and often an additional minute seta at tip; basal segment with smooth seta and one (0–2) additional, small, barbed seta; endopod with distinct subbasal articulation (Fig. 17E). Scutellum paracaudale terminally well rounded or biconvex with rounded (rarely acute) apex (Fig. 17F–H), its lower margin occasionally almost straight. Endopod of uropod with one strong spine below statocyst, statolith composed of vaterite. Telson (Fig. 17J) subquadrangular to subtriangular, length 1.1–1.5 its maximum width or 0.7–1.0 times length of last abdominal somite; maximum width near basis 2.1–2.7 times that at apex; each lateral margin armed with 6–16 spines. Apical cleft 11–19% telson length, cleft lined by 9–23 laminae, its margins straight to convex. 36 · Zootaxa 4142 (1) © 2016 Magnolia Press WITTMANN ET AL. FIGURE 17. Diamysis lagunaris Ariani & Wittmann, 2000 (A–J), and D. hebraica Almeida Prado-Por, 1981 (K–Q). Materials from Lago di Caprolace (paratypes: A, C–E, G, J, Lazio coast of Tyrrhenian Sea), Étang de La Palme (B, Golfe du Lion, Mediterranean coast of France), coast of La Spezia (F, N-Tyrrhenian Sea), Lago di Ganzirri (H, Strait of Messina, Tyrrhenian Sea), and the coastal stream Nahal Tanninim (K–Q, Levantine Sea, coast of Israel). A, anterior body region of male with body length 5.6 mm, dorsal view on cephalothorax; B, eyes and anterior margin of carapace in female 5.3 mm, dorsal; C, exopod of first thoracopod in male 4.0 mm, caudal; D, tarsus (i.e. carpopropodus plus dactylus) of third thoracic endopod in female 6.1 mm, rostral; E, fourth pleopod of male 4.8 mm, rostral; F–H, posterior margin of sixth pleonite in females with 7.2 mm (F), 5.2 mm (G), or 5.1 mm (H) body length, lateral; J, telson of male 4.8 mm, dorsal; K, anterior body region of male 5.0 mm, dorsal view on cephalothorax; L, tarsus of third thoracic endopod, male 5.0 mm, rostral; M, exopod of sixth thoracopod in male 4.0 mm (paratype), rostral; N, exopod of fourth pleopod of male 5.0 mm, rostral; O, posterior margin of sixth pleonite, male 4.0 mm (paratype), lateral; P, the same for female 4.0 mm (holotype); Q, telson of male 5.0 mm, dorsal. A, K, pores on carapaces not to scale; A–J, from Ariani & Wittmann (2000); K–Q, original. MYSIDAE IN MEDITERRANEAN FRESH-WATERS Zootaxa 4142 (1) © 2016 Magnolia Press · 37 Body length. Adult females 4.1–8.1 mm, males 3.6–6.6 mm. Distribution (Fig. 6). Mainly in the western Mediterranean: along the coasts of the Tyrrhenian, Sardinian and Ligurian Seas, Golfe du Lion, Strait of Messina; rare in the eastern Mediterranean: Island of Crete in the Aegean Sea. The populations at the Atlantic coasts of southern Spain and Portugal may have originated from Mediterranean lagoons by transfer in ballast water (Cunha et al. 2000: as D. bahirensis), although an indigenous status of the Atlantic populations is not excluded (Wittmann & Ariani 2012a). Type locality is the mixoeuhaline to weakly metahaline lagoon Lago di Caprolace at the Lazio coast, Tyrrhenian Sea. The species is mostly found in mixoeuhaline to metahaline lagoons, also in marine coastal habitats as well as mesohaline to mixoeuhaline reaches of estuaries. Normal salinity range 14–49; so far only two positive samples from the oligohaline reach (S = 2–3), taken at different stations in the Rhône Delta on the Mediterranean coast of France. Diamysis hebraica Almeida Prado-Por, 1981 Fig. 17K–Q Diamysis bahirensis hebraica Almeida Prado-Por, 1981: Herbst & Mienis 1985; Por & Dimentman 1985, 1989; Ben-Eliahu 2008; ITIS 2014; Mees 2014. Diamysis hebraica: Ariani & Wittmann 2000: 2004; Ariani 2004; Daneliya et al. 2012; Wittmann & Ariani 2012a, b. Material examined. Holotype F ad. with body length 4.0 mm (HUJ: reg. no. ICr62, HUJINVCRUMYS3), one paratype M ad. 4.0 mm (HUJ: ICr63, HUJINVCRUMYS3), Levantine Sea, Mediterranean coast of Israel, brackish stream Nahal Tanninim and its springs, S = 0.7–1.8, 200 µm mesh hand net, Jan.–Feb. 1980, det. Almeida PradoPor; 8 M ad. 4.1–5.5 mm, 21 F ad. 4.9–6.2 mm, 1 F subad. 4.6 mm, three stations along the same coastal stream as before, 32.5491N 034.9146E to 32.5487N 034.9136E, 2.0–1.9 km to the sea, 0.1–1 m depth, among filiform green algae, S = 3–5, 18°C, very small hand net, 18/20 Dec. 2008, leg. K. J. Wittmann, NHMW 25708. Supplemented diagnosis. Cornea diameter large, 70–90% the length of eyestalk (Fig. 17K). Antennula of males with appendix masculina pointing obliquely forward to inward; this appendix 90–126% the length of the terminal segment of antennular peduncle. Rostrum well rounded or forming a wide convex angle with broadly rounded tip. Carapace without fringes (Fig. 17K) in both sexes. Palpus of maxilla with subcircular terminal segment, armed with 9–16 denticles along distal margin. Basis of all thoracic exopods terminally rounded (Fig. 17M). Thoracic endopod 3 with 2–3-segmented carpopropodus; this carpopropodus longer than 5 times its maximum width (Fig. 17L); thoracic endopods 3–8 with long and slender claw. Penis short, with smooth setae only. Male pleopod 4 biramous with 2-segmented sympod, with small, 2-segmented endopod, and with moderately long, rod-like, 2-segmented exopod (Fig. 17N) bearing a long modified seta at tip and a shorter, smooth seta subterminally on the basal segment. Scutellum paracaudale sinusoid (Fig. 17O) to triangular with blunt tip (Fig. 17P), i.e. not ending in a spiniform process as in the otherwise similar D. bahirensis (G. O. Sars, 1877). Endopod of uropod with one spine below large statocyst; statolith composed of vaterite. Telson (Fig. 17Q) subtriangular, 0.7– 0.9 times length of last abdominal somite; with lateral margins slightly sinusoid, maximum width is 2.0–2.7 that at apex. Apical cleft is 10–13% telson length, cleft with straight to slightly convex margins, bottom of cleft angular; cleft lined by 8–13 laminae (Fig. 17Q). Supplements to the description by Almeida Prado-Por (1981). Flagellum of thoracic exopod 1 with 8 segments, whereas exopods 2–8 each with 9 segments, not counting the large intersegmental joint between basis and flagellum which may be mistaken as a segment. Thoracic endopods 3 and 4 each with 2–3-segmented carpopropodus, endopods 5–8 with 2-segmented carpopropodus. Penis with 5–8 smooth setae close to ejaculatory opening. Male pleopod 4 with field of scales on inner face of terminal segment of sympod; basal segment of endopod with terminally setose exite; terminal segment of endopod is slender, with plumose setae all along its distal 60–75%. Endopod of uropods is 73–75% length of exopod. Statolith diameter 118–142 µm. Lateral margins of telson each with 6–10 spines, not counting the larger terminal spines. Distribution (Fig. 6). According to Herbst & Mienis (1985) known from three coastal streams leading to a stretch of about 40 km along the Mediterranean coast of Israel: Nahal Qishon, N. Daliya, and N. Tanninim. Type locality is the stream Tanninim, where the species lives in the salinity range of 0.7–5, mostly at 1–4. It was not found in the freshwater section of this stream (Herbst & Mienis 1985; inspection by one of us, KJW, in Dec. 2008). 38 · Zootaxa 4142 (1) © 2016 Magnolia Press WITTMANN ET AL. Genus Troglomysis Stammer, 1933 Fig. 18 Revised definition. Diamysini with cornea present as small external rudiments distally on the large eyestalks in both sexes (Fig. 18B, C). Antennal scale without spines, setose almost all around (Fig. 18A, D); scale with short apical segment, terminally rounded in both sexes. Carapace with a pair of post-suborbital spines, and without fringes in both sexes (Fig. 18A). Thoracic endopods 3–8 with 3–6-segmented carpopropodus, dactylus small, with distinct, in part seta-like claw (Fig. 18E–L); dactylus of endopod 8 flanked by a pair of minute paradactylar lobes (Fig. 18K) in both sexes. All female pleopods and male pleopods 1, 2, 5 reduced to setose rods (Fig. 18O). Male pleopod 3 reduced to well-developed, two-segmented sympod terminally fused with its small, setose, unsegmented endopod, exopod missing (Fig. 18M). Male pleopod 4 (Fig. 18N) biramous with 2-segmented sympod, with small, 2-segmented endopod, and with large, 3-segmented exopod bearing a modified seta at tip and a very long smooth seta at penultimate segment; seta from the penultimate segment extends far beyond the seta from the apical segment (Fig. 18N). Endopod of uropod setose all around, with only one spine below statocyst. Telson with small apical cleft; with spines on each lateral margin, these margins ending in a pair of large, posteriorly directed apical spines; cleft lined by a number of laminae (Fig. 18S, T). Type species. Troglomysis vjetrenicensis Stammer, 1933, by monotypy. Troglomysis vjetrenicensis Stammer, 1933 Figs 7C, 11B, 18 Troglomysis vjetrenicensis Stammer, 1933: 1935, 1936; Caroli 1937; Jeannel, 1949; Thienemann 1950; Karaman 1954; Gordan 1957; Riedl 1966; Holmquist 1972; Mauchline 1972; Mauchline & Murano 1977; Ariani 1981b, 2004; Schram 1986; Ariani et al. 1993; Müller 1993; Pesce et al. 1994; Kobusch 1999; Sket 1999; Anderson 2008; Daneliya et al. 2012; ITIS 2014; Mees 2014; Wittmann et al. 2014; Meland et al. 2015. Material examined. 6 F ad. 10.4–12.8 mm, mostly incubating nauplioid larvae, 1 F subad. 12.0 mm, 1 F imm. 9.7 mm, 1 M ad. 8.9 mm, plus fragments of 2 M ad. and 2 imm., subterranean freshwater lake in the karstic cave Donja Vjetrenica, 42.85N 017.98E, Herzegovina, Sept. 1960, leg. B. Sket, BUL; 5 F ad. 9.9–11.1 mm, mostly incubating nauplioid larvae, 1 F subad. 10.6 mm, 2 M ad. 9.4–9.9 mm, sampling data as above, 26 Sept. 1964, BUL; 1 M ad. 11.9 mm in 2 parts, sampling data as above, Oct. 1986, BUL; 1 M ad. 12.5 mm, completely dissected and mounted on slides, freshwater lake in same cave, Sept. 1981, leg. A. P. Ariani; 1 M ad. 10.6 mm, fresh-water in same cave, 4 Aug. 2000, leg. B. Trontelj, BUL. Short diagnosis. Troglomysis without eye pigment in both sexes, also body without any pigment (Fig. 7C). Apical segment is 6–9% total length of antennal scale (Fig. 18D). Total length of thoracic endopods, and of their carpopropodus, in particular, decreases strongly in series of endopod 3 to 7, and following this increases strongly to endopod 8; carpopropodus 3–8 with 5–6, 5, 4, 3–4, 3–4, and 4–5 segments, respectively (Fig. 18E–K). Telson with 10–13 spines all along each lateral margin, not counting the pair of apical spines; small sinusoid cleft between the apical spines, cleft lined by 5–8 laminae (Fig. 18S, T). Supplements and corrections to the description by Stammer (1936). Rostrum forms a wide convex angle with broadly rounded tip (Fig. 18A). Eyestalks distally with external rudiments of cornea (Fig. 18B, C) visible in all specimens with eyes well preserved (n = 19; including the smallest one: immature female with 7.3 mm body length). Rudiments of cornea not drawn and not reported by Stammer. Antennal scale with small apical segment bearing five plumose setae (Fig. 18D). In contrast, Stammer drew and explicitely stated an unsegmented antennal scale. Flagellum 8-segmented in thoracic exopod 1, and 9-segmented in each of exopods 2–8 (Fig. 18L). Stammer figured the exopods 1, 2 correctly, but counted a surplus of one segment for each of exopods 1–8, obviously by considering the large intersegmental joint between basis and flagellum as a distinct segment. Outer distal corner of basis always well rounded (Fig. 18L). Thoracic endopods 3–8 (Fig. 18E–L): tarsus (carpopropodus plus dactylus) with 6–7, 6, 5, 4–5, 4–5, and 5–6 segments, respectively (Stammer reported only 5, 5, 4, 3, 3, and 5 "tarsal" segments, respectively, presumably by not counting the minute dactylus not mentioned by him); dactylus of endopods 3–7 with long, slender claw (Fig. 18E–J); dactylus 8 with very slender, seta-like claw (Fig. 18K) in both sexes; paradactylar setae with weak, one-sided armature of small barbs. The previously unknown paradactylar MYSIDAE IN MEDITERRANEAN FRESH-WATERS Zootaxa 4142 (1) © 2016 Magnolia Press · 39 FIGURE 18. Troglomysis vjetrenicensis Stammer, 1933, from the cave Donja Vjetrenica (type locality). A, male with body length 9.9 mm, dorsal view on cephalothorax; B, left eye of male 12.5 mm, dorsal; C, right eye of female 12.0 mm, dorsal; D, antennal scale of male 12.5 mm; E–K, series of tarsi of thoracic endopods 3–8 in same male, setae omitted; L, eighth thoracopod with penis of same male; M–O, series of third to fifth pleopods in same male (M, O, views on outer = rostral face; N, inner = caudal face); P–R, posterior margin of sixth pleonite (lateral aspect turned by 90° to the left) in female 10.2 mm (P), female 10.4 mm (Q), and male 12.5 mm (R); S, telson of male 12.5 mm; T, distal third of telson in subadult female 12.0 mm. 40 · Zootaxa 4142 (1) © 2016 Magnolia Press WITTMANN ET AL. lobes (Fig. 18K) on endopod 8 represent an uncommon feature in Mysidae; endopods 1–7 without such lobes (Fig. 18E–J). Scutellum paracaudale triangular with rounded tip, upper and lower margins inconspicuously undulate (Fig. 18P–R). Telson cleft is lined by laminae (Fig. 18S, T) rather than by spines as otherwise counted and drawn by Stammer (1936: Fig. 17). Statoliths (Fig. 11B) composed of fluorite, diameter 96–157 µm; statolith formula 2 + (2–3) + (0–2) + (8–13) + (7–12) = 19–30. General form discoid, maximum height only about 50-70% maximum width. Flattened ellipsoidal in lateral view, subcircular in ventral view. Fundus and ambitus ventrally forming together a weakly vaulted structure. Occurrence (Fig. 12). Only known from subterranean freshwater lakes and canals in the karstic cave Vjetrenica, one of the world's highest biodiversity cave systems (Culver & Sket 2000), 42.85N 017.98E, near the village Zavala in the karstic plain Popovo polje, Herzegovina, 12 km from the coast of the Adriatic Sea. Type locality is the Donje jezero (Lower Lake) within the Donja Vjetrenica (Lower Vjetrenica) in this system. According to Stammer (1936) this lake is 234 m above sea level. Comparison. Already Stammer (1936) postulated some similarity but not a close relationship to the genus Diamysis Czerniavsky, 1882. Ariani (1979) noted that certain Diamysis from the Adriatic basin share with Troglomysis (referring to Stammer’s description) the highest numbers of carpopropodal segments (three or five, respectively) in thoracic endopods 3, 4, and 8, in contrast to fewer in endopods 5–7. Most of our new data make this morphological similarity appear closer: antennal scale with apical segment bearing five setae, and sinus of telson without spines but with laminae. However, the lobes flanking the dactylus of the eighth thoracic endopods represent a newly discovered feature (besides the already known ones), supporting the status of Troglomysis Stammer, 1933, as a separate genus. Genus Limnomysis Czerniavsky, 1882 Limnomysis benedeni Czerniavsky, 1882 Fig. 19 Short selection from 27 synonymy statements with a total of 416 references: Mysis relicta var. pontica Grebnitzky (in synonymy by Czerniavsky 1882a) (nomen nudum). Limnomysis Benedeni Czerniavsky, 1882a: 1887; Sars 1893; Chirica 1914; Colosi 1930. Limnomysis Brandti Czerniavsky, 1882a: 1887; Sowinsky 1904 (partim: 457). Limnomysis Schmankewiczi Czerniavsky, 1882a: 1887; Sowinsky 1895, 1904. Onychomysis mingrelica Czerniavsky, 1882a: 1887; Sowinsky 1904 (partim: 141, 338); ITIS 2014. Limnomysis benedeni: Zimmer 1915a; Derzhavin 1924; Komarova 1991; Kelleher et al. 1999; Daneliya 2002; Daneliya et al. 2012; Wittmann et al. 2014; Zettler 2015. Mysidella bulgarica Valkanov, 1935: 1936a, 1936b. Limnomysis brandti: ITIS 2014. Limnomysis schmankewiczi: ITIS 2014. Material examined (hand net, leg. K. J. Wittmann): Mediterranean drainage (southern France, Rhône system, 15–19 June 2009). 2 F ad. 7.3–8.3 mm, among ~30,000 Mesopodopsis slabberi and 3 Diamysis lagunaris, estuary of the Petit Rhône at Tiki, same sample as indicated above for M. slabberi; 1 F subad. 5.5 mm, 1 F imm. 4.7 mm, estuary of the Grand Rhône at Port St. Louis, Rhône-km 323.2, left bank, 43.3828N 004.8073E, sea distance 6 km, 0.5–4 m depth, among Potamogeton and on filiform algae on concrete walls and wrecks, v = 0 m/s, stratified salinity S = 0.1–3.5, 473–6300 µS/cm, near surface, 22.5°C, pH 7.63, 11°d, 7.35 mg O2/l, 17 NTU; 1 M ad. 6.0 mm, 1 F imm., 1 juv., (canal) Liaison Rhône— Fos, 100 m after branching off from Rhône River, 43.4217N 004.7470E, altitude 1 m, sea distance 13 km, 0.5–2.5 m depth, among Potamogeton, Elodea, and on boulders, v = 0 m/s, S = 0.2, 574 µS/cm, 23.0°C, pH 7.73, 11°d, 7.89 mg O2/l, 36 NTU; 6 F ad. 7.1–8.5 mm, 4 M ad. 6.0–8.2 mm, Canal d'Arles à Fos, in Arles, small basin at the Nouveau Pont, 50 m above the first lock after having branched off from the river at Rhône-km 283, 43.6725N 004.6185E, altitude 5 m, sea distance 47 km, 0.2–2 m depth, among bank vegetation, Trapa, Myriophyllum, and Carex, v = 0 m/s, S = 0.1, 390 µS/cm, 22.1°C, pH 7.44, 10°d, 6.67 mg O2/l, 20 NTU, NHMW reg. no. 25709; 1 M ad. 7.2 mm, Bras de l'Ardoise, Port 2—Port de la Plaisance, right bank at Rhône-km 214, 44.1051N 004.7058E, MYSIDAE IN MEDITERRANEAN FRESH-WATERS Zootaxa 4142 (1) © 2016 Magnolia Press · 41 altitude 28 m, sea distance 112 km, 0.3–2 m depth, among Myriophyllum, Elodea, and on filiform algae on stones, v = 0.1–0.3 m/s, S = 0.1, 434 µS/cm, 22.7°C, pH 7.43, 12°d, 6.68 mg O2/l, 16 NTU; 1 F ad. 8.1 mm, old branch of Rhône River at Le Pal de Fer (2.5 km upstream of Viviers), right bank at Rhône-km 163.5, 44.5078N 004.6881E, altitude 64 m, sea distance 161 km, 0.2–1.5 m depth, among bank vegetation and Myriophyllum, v = 0.1–0.3 m/s, S = 0.1, 395 µS/cm, 21.4°C, pH 7.83, 10°d, 9.44 mg O2/l, 5 NTU; 4 F ad. 6.5–8.3 mm, downstream of Meysse, at corner with Old Rhône, 300 m downstream of Barrage de Loriol, right bank at Rhône-km 135.8, 44.5923N 004.7229E, altitude 70 m, sea distance 169 km, 0.2–1.5 m depth, among Potamogeton and from filiform algae on stones, v = 0.1–0.3 m/s, S = 0.1, 356 µS/cm, 20.3°C, pH 7.68, 10°d, 7.52 mg O2/l, 20 NTU; 3 M ad. 5.6–6.3 mm, 1 M subad., 2 imm., great bay in Tournon-sur-Rhône, right bank at Rhône-km 90.4, 45.0729N 004.8255E, altitude 118 m, sea distance 234 km, 0.2–1.8 m depth, from filiform algae on stones, v = 0 m/s, S = 0.1, 303 µS/cm, 23.2°C, pH 7.78, 10°d, 8.59 mg O2/l, 11 NTU; 3 F ad. 7.7–8.5 mm, 3 M ad. 5.9–8.1 mm, 1 F subad., 2 juv., right bank of Rhône River at km 34, this is shortly downstream of the Barrage de Vaugris, 45.4949N 004.8247E, altitude 144 m, sea distance 287 km, 0.5–2 m depth, among Potamogeton and on boulders, v = 0.1–0.2 m/s (sheltered microhabitat), S = 0.1, 371 µS/cm, 20.6°C, pH 7.51, 10°d, 7.07 mg O2/l, 17 NTU; 3 F ad. 6.3–8.4 mm, 1 M ad. 7.9 mm, 2 juv., Lyon, Port Edouard Herriot at Rhône-km 3.3, 45.7056N 004.8430E, altitude 160 m, sea distance 320 km, 0.5–2 m depth, among Elodea, Potamogeton, and on stones with coat of algae, v = 0 m/s, S = 0.1, 393 µS/cm, 20.8°C, pH 7.79, 14°d, 8.38 mg O2/l, 6 NTU; 4 M ad. 4.9–7.3 mm, 4 imm., Villefranche-sur-Saône, right bank of Saône River at km 37, basin at the Chemin du Colombier, 45.9619N 004.7389E, altitude 166 m, sea distance 360 km, 0.3–1 m depth, among Elodea and Myriophyllum, v = 0 m/s, S = 0.1, 408 µS/cm, 22.7°C, pH 7.64, 16°d, 13.24 mg O2/l, 24 NTU; 2 M ad. 5.6–6.5 mm, Mâcon, right bank of Saône River at km 79.5, 46.2988N 004.8311E, altitude 165 m, sea distance 403 km, 0.5–2.5 m depth, from dirty Myriophyllum, Nuphar, and stones, v = 0 m/s, S = 0.2, 591 µS/cm, 22.1°C, pH 7.29, 16°d, 6.28 mg O2/l, 15 NTU; 2 F ad. 5.8–5.9 mm, 3 F subad., Chalons-sur-Saône, Zone Portuaire Sud at left bank of Saône River, at river-km 137, 46.7619N 004.8789E, altitude 172 m, sea distance 461 km, 0.3–2.5 m depth, from dirty Potamogeton and algae on stones, v = 0 m/s, S = 0.2, 601 µS/cm, 22.1°C, pH 7.29, 15°d, 9.74 mg O2/l, 12 NTU; 6 F ad. 5.7–7.5 mm, 1 M ad. 5.6 mm, 4 F subad., 1 F imm., Chalons-sur-Saône, yachting harbour at Saône-km 144, 46.7925N 004.8853E, altitude 173 m, sea distance 467 km, 0.3–2.5 m depth, from shore vegetation, Carex, algae on stones, and boulders, v = 0 m/s, S = 0.2, 583 µS/cm, 23.0°C, pH 7.01, 15°d, 8.05 mg O2/l, 9 NTU. Marmora Sea drainage (Turkey). 191 F ad. 6.0-7.6 mm, 141 M ad. 5.7-7.1 mm, 42 F subad., 28 M subad., 67 imm., 70 juv., NE-coast, (lagoon) Büyükçekmece Gölü, northern freshwater reach, 41.1041N 028.5614E, altitude 3 m, sea distance 10 km, 0–1.5 m depth, among roots of Salix and on boulders, v = 0 m/s, S = 0.2, 484 µS/cm, 21.5°C, pH 8.10, 10°d, 6.19 mg O2/l, 15 NTU, 14 June 2006, NHMW 25710; 1 F ad. 8.9 mm, 1 F subad. 7.0 mm, NE-coast, (lagoon) Küçükçekmece Gölü, 41.0105N 028.7700E, at about sea level, sea distance 5 km, 0.5 m depth, on algae, stones, and boulders, v = 0 m/s, S = 6, 10000 µS/cm, 26°C, 1 July 1988; 12 F ad. 5.7–8.6 mm, 5 M ad. 5.6–6.4 mm, 2 subad., 3 imm., NE-coast, small river near its mouth into (lagoon) Küçükçekmece Gölü, 41.0640N 028.7433E, altitude 2 m, sea distance 11 km, 0.3–0.5 m depth, among shore vegetation, v = 0.1–0.2 m/s, S = 0.3, 800 µS/cm, 27°C, 1 July 1988; 62 F ad. 5.9–8.9 mm, 11 M ad. 5.9–7.6 mm, 13 subad., 16 imm., 2 juv., S-coast, (lake) Uluabat Gölü, near Karaağaç, 41.1943N 028.6150E, altitude 4 m, sea distance 54 km, 0.5 m depth, among macrophytes, v = 0 m/s, S = 0.1, 500 µS/cm, 30°C, highly eutrophic aspect, 5 July 1988, MHNG; 183 F ad. 5.8–7.9 mm, 129 M ad. 5.5–7.9 mm, 96 subad., 146 imm., 72 juv., same lake as above, at Eskikaraağaç, 41.1889N 028.6087E, sea distance 51 km, 1–2 m depth, from mud, Myriophyllum and Potamogeton, v = 0 m/s, S = 0.2, 528 µS/cm, 19.9°C, pH 7.46, 8°d, 6.58 mg O2/l, 3 NTU, 13 June 2006; 2 F ad. 6.7–7.5 mm, 2 F subad., 2 imm., 3 juv., S-coast, small river Koca Dere, near Akçasusurluk, 40.2884N 028.4332E, altitude 4 m, sea distance 16 km, 0–1 m depth, among bank vegetation, v = 0.1 m/s, S = 0.1, 440 µS/cm, 27°C, 5 July 1988. NE-Atlantic drainage (from the canal network connecting the river systems of Seine, Meuse, and Rhine, 29 June 2009). 1 F ad. 6.3 mm, 2 M ad. 6.4–7.0 mm, 2 imm., Canal de l'Est, branche Nord, between Troussey and Vertuzey, shortly upstream of Écluse no. 3 Frâsnes, 48.7159N 005.6723E, altitude 236 m, sea distance 711 km (Meuse) or 731 km (Seine), respectively, 0.2–2.5 m depth, among Potamogeton, Myriophyllum, Elodea, and from filiform green algae on steel walls, v = 0–0.1 m/s, S = 0.1, 450 µS/cm, 23.5°C, pH 7.71, 13°d, 7.73 mg O2/l, 16 NTU; 2 F ad. 7.1–8.0 mm, 1 M ad. 7.2 mm, 1 F subad., 2 imm., Canal de la Marne au Rhin, near Pagny-sur-Meuse, at canal-km PK 115.0, 48.6904N 005.7171E, altitude 243 m, sea distance 717 km (Meuse) or 734 km (Seine), 42 · Zootaxa 4142 (1) © 2016 Magnolia Press WITTMANN ET AL. respectively, 0.1–0.5 m depth, from filiform green algae on steel walls, v = 0–0.1 m/s, S = 0.1, 454 µS/cm, 22.5°C, pH 7.50, 13°d, 8.93 mg O2/l, 29 NTU, NHMW 25711. FIGURE 19. Limnomysis benedeni Czerniavsky, 1882, from the freshwater reach of the Canal d'Arles à Fos, a tributary of the Rhône River; station is 47 river km from the Mediterranean coast of France; male with body length 6.7 mm (A, C, E–M) and females with 9.0 mm (B) or 8.1 mm (D). A, anterior body region of male, dorsal (pores on carapace not to scale); B, right eye, dorsal; C, male antennula, dorsal; D, antennal scale of female, dorsal; E, male antenna with posterior lobe containing end sac of antennal gland, dorsal; F, tarsus (carpopropodus plus dactylus) of third thoracic endopod; G, third male pleopod, outer = rostral face; H, fourth male pleopod, inner = caudal face; J, fifth male pleopod, outer = rostral face; K, posterior margin of sixth pleonite, lateral; L, uropods, ventral; M, telson, dorsal. MYSIDAE IN MEDITERRANEAN FRESH-WATERS Zootaxa 4142 (1) © 2016 Magnolia Press · 43 Description. Adult body size 5–12 mm. Eyes normal, length of cylindrical eyestalks without cornea is 1.1–2.0 times cornea diameter (Fig. 19B). Eyestalk dorsally with series of 2–5 ommatidia shifted a small distance proximally, i.e. away from cornea (Fig. 19B), similar to the findings in Diamysis fluviatilis (Fig. 13A) and D. lacustris (Fig. 13E). The eyestalk surface close to this ommatidia series is generally poorly pigmented but does not form a distinctly differentiated fenestra paracornealis such as found in D. lacustris (Fig. 13E). Antenna with large posterior lobe (Fig. 19E) containing end sac of antennal gland. Antennal scale 2-segmented, setose all around, terminally rounded in females (Fig. 19D), whereas pointed with bent, setose tip in males (Fig. 19A, E). Apical segment is 25–35% scale length. Carapace (Fig. 19A) anteriorly broadly rounded; no rostral and subrostral processes; dorsal surface with 5–10 cervical pores in about V-shaped arrangement; and with 8–18 cardial pores in about linear, slightly curved, transverse arrangement. Total length of thoracic endopods, and also of their carpopropodus, remain roughly the same in series of endopods 3 to 6, and then decrease by a total of 20–25% along endopods 7 and 8; carpopropodus 3–8 always 3-segmented as in Fig. 19F. Flagellum of thoracic exopod 1 with 8–9 segments, that of exopods 2–8 each with 9–10 segments, not counting the large intersegmental joint between the 1segmented basis and its multi-segmented flagellum; segmental counts tend to increase with increasing body size. Outer distal corner of basis always well rounded. All female pleopods and male pleopods 1, 2, and 5 reduced, fused, and styliform (Fig. 19J). Third male pleopod also terminally fused, but with larger, 2-segmented sympod (Fig. 19G). Fourth male pleopod with distinct sympod, endopod, and exopod (Fig. 19H). Sympod 2-segmented. Setose endopod subbasally with terminally setose exite. Exopod longer, distinctly or indistinctly 3-segmented, not reaching base of uropods. Its median segment with 2–5 small, conical, spine-like setae along inner margin, and with series of 3–6 minute spine-like setae in submedian position, at a small distance parallel to outer margin (Fig. 19H). Exopod usually stylet-like (Fig. 19H), but alternatively may appear bifid or even trifid, which is commonly attributed to regeneration. These last variants usually rare, but may account up to 30% of adult males in certain populations (Băcescu 1940, Kelleher et al. 1999). Scutellum paracaudale subtriangular, with slightly concave upper, and slightly convex lower margin; tip rounded (Fig. 19K). Endopod of uropod with one spine below statocyst (Fig. 19L). Statoliths composed of vaterite. Telson rather short and stout (Fig. 19M), subtriangular to almost subrectangular, with small apical incision; incision well rounded to rounded subtriangular, apart from being armed with 4–10 laminar processes. Lateral margins of telson with 7–14 spines each. Distribution (Fig. 16). Endemic in the Pontocaspian and in waters draining into the Marmora Sea (Băcescu 1954, Kelleher et al. 1999, Wittmann & Ariani 2009). It inhabits mostly fresh- and oligohaline waters of lakes, lagoons, and river mouths. Here it dwells mainly among vegetation in shallow water (0.1–2 m, less frequently down to 10 m); maximum depth 68 m in the Caspian Sea (Derzhavin 1939). Salinity optimum at S = 0–5, lower densities at S = 6–12 in river estuaries and lagoons along coasts of the Marmora and Black Seas (Kelleher et al. 1999), also found in coastal 'marine' waters of the Caspian Sea up to S = 12.8 (Sars 1907, Derzhavin 1939). It is potamophilous, avoiding current velocities > 0.5 m/s. Before human intervention, it penetrated only a few hundred kilometres into rivers (Băcescu 1940). For areal expansion of L. benedeni to tributaries of the North Sea, Baltic, and Mediterranean, see ‘Discussion’. Tribe Neomysini Wittmann, Ariani & Lagardère, 2014 Genus Neomysis Czerniavsky, 1882 Neomysis integer (Leach, 1814) Fig. 20 Short selection from 22 synonymy statements with a total of 622 references: Praunus integer Leach, 1814: Stebbing, 1893. Mysis integer: Leach 1815, 1817; White 1847, 1850; Bell 1853 (with query). Mysis entier [non-Linnean assignment]: Desmarest 1823, 1825; H. Milne Edwards 1837 (in remarks). Praunus flexuosus: Desmarest 1825 (in synonymy). Mysis vulgaris J. V. Thompson, 1828: H. Milne Edwards 1837; Kröyer 1861; Czerniavsky 1882a (partim: 2, 7); Ekman 1935; Muus 1967 (partim: Tab. 29). Mysis scoticus J. V. Thompson, 1828. Mysis commun [non-Linnean assignment]: H. Milne Edwards 1837. 44 · Zootaxa 4142 (1) © 2016 Magnolia Press WITTMANN ET AL. Neomysis vulgaris: Czerniavsky 1882a (partim: 15), 1882b, 1887; Needham 1937; Kinne 1955; Ten 1991; Mitina & Kharina 2011. Neomysis integer: W. M. Tattersall 1912; Băcescu 1941; Brun 1967; Wittmann & Ariani 2009; Remerie et al. 2004; Wittmann 2013; Zettler 2015. Material examined (hand net, leg. K. J. Wittmann, if not stated otherwise): Mediterranean drainage. 3 F ad. 11.6 mm (one dissected), 1 M ad. 9.1 mm, 1 M subad., 1 F imm., Canal d'Arles à Fos within the estuary of the Grand Rhône, S = 12.3, NHMW reg. no. 22995, for additional data see Wittmann & Ariani (2009); 1 M ad. 7.6 mm (dissected and mounted on slides), Canal du Rhône à Sète within the estuary of the Petit Rhône, 43.6432N 004.4298E, altitude 1 m, sea distance 30 km along canals or 42 km along the river Petit Rhône, 0.2–1.8 m depth, from shore macrophytes and from boulders with filiform algae, among masses of Atyaephyra desmarestii, v = 0 m/s, S = 0.4, 1010 µS/cm, 24.5°C, pH 7.92, 6.54 mg O2/l, 53 NTU, 19 June 2009. NE-Atlantic drainage. 1 F ad. 8.7 mm, 8 M ad. 7.6–8.7 mm, 4 F subad., 5 M subad., 15 imm., 2 juv., France, estuary of Gironde River at Cadou, river-km K.60, 45.3097N 000.7815W, altitude 0 m, sea distance 37 km, 0.5–3 m depth, from stone walls with filiform algae, v = 0.2–0.4 m/s, S = 4.1, 7620 µS/cm, 23.6°C, pH 7.67, 6.34 mg O2/ l, 222 NTU, 22 June 2009, NHMW 25712; 23 F ad. 8.7–11.6 mm, 31 M ad. 6.7–9.3 mm, 15 F subad., 15 M subad., 24 imm., 6 juv., France, Bay of Biscay, Bassin d'Arcachon, chenal de La Hume, 44.5921N 001.1237W, S = 1.2, 2 Aug. 1983, don. J. C. Sorbe (Arcachon), RNMH.CRUS.E.3782; 1 juv., The Netherlands, Oude Rijn, Katwijk, at Valkenburgseweg, 52.1877N 004.4274E, altitude 0 m, sea distance 4 km, 0.2–1 m depth, among Nuphar, S = 0.4– 1.0, 23 July 1998; 60 F ad. 9.2–11.8 mm, 5 M ad. 9.1–10.8 mm, 2 F subad., 1 M imm., The Netherlands, Den Helder, Noordhollands kanaal, 52.9638N 004.7667E, altitude 2 m, sea distance 5 km, 1–2 m depth, among macrophytes, S = 2–3, 22 June 1989, NHMW 25450; 26 F ad. 10.7–16.4 mm, 40 M ad. 9.0–12.0 mm, 2 F subad., 3 M subad., 49 imm., 23 juv., Scotland, Argyll, Loch Creran, near Inver, 56.5137N 005.3728W, altitude 0 m, sea distance about 26 km, 0.2–0.3 m depth, among algae, low tide, tidal current v = 0.3–0.6 m/s, S = 0–25, 11 Sept. 1983; 1 F ad., 1 M ad., 1 imm., Scotland, Argyll, Firth of Lorn, Dunstaffnage Bay, 56.450N 005.433W, 0.1–0.2 m depth, among algae and on mud, S = 28–33 (mixoeuhaline), 12°C, 13 Sept. 1983, NHMW 25449. Baltic drainage. 16 F ad. 12.0–16.7 mm, 61 M ad. 7.6–15.7 mm, 13 subad., 5 imm., 3 juv., Germany, FehmarnBelt, Puttgarden, 54.505N 011.222E, 0.3 m depth, sand, S = 14, 23 Aug. 1985, leg. K. J. Wittmann & A. P. Ariani, RNMH.CRUS.E.3783; 172 F ad. 10.7–16.7 mm, 35 M ad. 8.5–12.9 mm, 10 F subad., 10 imm., Sweden, Askö Island, 58.8288N 017.6366E, 0.3–1 m depth, among algae, S = 7, 22 Aug. 1985, NHMW 25713. Supplementary description. Adult body length 6–19 mm, eyes normal. Antennal scale strongly elongate, setose all around, terminally pointed in both sexes (Fig. 20A, B). Its two segments separated by a transverse, not always distinct suture; apical segment is 12–18% scale length. Carapace (Fig. 20A) ends anteriorly in an apically acute, triangular rostrum; no subrostral plates; lateral margins of carapace produced into a pair of anteriorly directed spiniform processes; dorsal face of carapace with 5–12 cervical pores in about linear, transversal arrangement; and with 12–29 cardial pores showing a transverse arrangement in two about symmetrical, slightly curved subgroups. Thoracic endopod 8 represents the strongest leg in both sexes by being always more stout and by being 0–30% longer compared with the subequal endopods 3–7; carpopropodus of endopods 3–8 with 5–6, 5–7, 5–7, 5–7, 5–7, or 6–8 segments, respectively (adult males tend to show more segments than same-sized adult females); these endopods ending in short, weak, slightly bent claws (Fig. 20E). Flagellum of thoracic exopods 1 and 8 each with 8 segments, that of exopods 2–7 each with 9 (10) segments, not counting the large intersegmental joint between the 1-segmented basis and its multi-segmented flagellum. Outer distal corner of basis always ending in a small spine. First thoracic sternite produced into an anteriorly directed, distally hairy (with minute setae), terminally nearly quadrangular, medial lobe in both sexes (Fig. 20C, D). Adult males (Fig. 20C): thoracic sternites 2–5 each with small median, terminally blunt processes each bearing 2–4 minute acute scales; sternite 6 medially with transverse cuticular ridge that may represent a vestigial process; sternites 7, 8 with smooth median portions. Adult females (Fig. 20D): thoracic sternites 2–6 medially with vestigial processes; a total of two large, finger-like processes project from thoracic sternites 7, 8 ventrally into the marsupium; these processes show densely set knob-like structures giving their surface a moruloid appearance, no scales, large longitudinal tube inside each process. Marsupium supported by a large, backwardly produced plate (Fig. 20D) from last thoracic sternite. This plate present only in females and represents an apron which overlaps part of first pleon sternite. Its alignment with anterior margin of pleon suggests a function as mechanical protection of marsupium upon flipping of pleon, which MYSIDAE IN MEDITERRANEAN FRESH-WATERS Zootaxa 4142 (1) © 2016 Magnolia Press · 45 FIGURE 20. Neomysis integer (Leach, 1814) from canals at the Mediterranean coast of France: adult male with body length 7.6 mm (A–C, E–J, L–N) from the freshwater reach of the Canal du Rhône à Sète, and incubating female 11.6 mm (D, K) caught in the mesohaline reach of the Canal d'Arles à Fos. A, anterior body region in dorsal view (pores on carapace not to scale); B, antenna, dorsal; C, thoracic sternites with median lobes in male, ventral; D, thoracic sternites in female (lobes forced into rostral plane by cover glass; in vivo projecting about ventrally), note large posterior plate from ultimate sternite; E, endopod of third thoracopod between merus and tip; F, right penis, lateral; G, third male pleopod, outer = rostral face; H, fourth male pleopod, inner = caudal face; J, posterior margin of sixth pleonite in male, lateral; K, the same for female; L, uropods, ventral; M, telson, dorsal, detail (N) shows hairs and spines on lateral margin. 46 · Zootaxa 4142 (1) © 2016 Magnolia Press WITTMANN ET AL. occurs most violently upon sudden escape. All pleopods reduced, fused and styliform (Fig. 20G) in both sexes, with exception of male pleopod 4 (Fig. 20H). This latter pleopod biramous, strongly elongate; its 2-segmented exopod reaches beyond base of uropods. Scutellum paracaudale mostly subtriangular with convex upper and concave lower margins (Fig. 20K), less frequently biconvex, and even less frequently sinusoid (Fig. 20J); tip blunt to well rounded. Endopods of uropods with 14–55 spines arranged in dense series along proximal half of inner margin (Fig. 20L). Statoliths composed of fluorite (according to Lowenstam & McConnell 1968; confirmed by own determinations on material from Canal d'Arles à Fos, chenal de La Hume, and Askö Island). Telson (Fig. 20M) entire, subtriangular, longer than ultimate pleonite; lateral margins all along with continuous series of spines, numerous fine hairs between the lateral spines (Fig. 20N); narrow, truncate apex with a pair of short median spines flanked by a pair of long latero-apical spines. Distribution (Fig. 16). Common in shallow coastal waters of the temperate to boreal NE-Atlantic from southern Spain to Norway, also common in the Baltic. Also found in the White Sea and the Barents Sea (Zimmer 1933, Holmquist 1972, Petryashov 2004). A presumably disjunctive population was detected by Băcescu (1941) in the NW-Mediterranean (Rhône Delta). This species occurs from anhaline to hyperhaline conditions, yet mainly in oligo- to polyhaline waters, where it has strong osmoregulatory capabilities (at S = 3–32 according to Vilas et al. (2006)). Freshwater records were made in waters pertaining to the drainage basins of the NE-Atlantic (Scott 1894, Elton 1937, Tattersall & Tattersall 1951, Hynes et al. 1960, Ketelaars et al. 1999, Lee & Bell 1999, Braune 2004) and of the Baltic (Gasiunas 1965, 1972; Zettler 1998). NE-Atlantic populations of N. integer are commonly found from oligohaline to polyhaline conditions, with main occurrence in the mesohaline reach, rarely in fresh-water. One own sample (listed above) of a juvenile was taken in almost fresh-water (S = 0.4–1.0) in an old Rhine branch at the coast of The Netherlands. The abundance of this species along the Westerschelde estuary at the Dutch-Belgian border is primarily correlated with salinity; here it lives at S = 0.6–27 throughout the year (Rappé et al. 2011). Similarly, Roast et al. (1998) observed a minimum of S = 1 for this species in a Cornwall estuary. There is a number of additional near-freshwater records (Castel 1993, Bernát et al. 1994, Köpcke & Kausch 1996, David et al. 2005, Vilas et al. 2009) as well as the above-cited true freshwater records along the Atlantic coasts of Europe. As a first freshwater record for the Mediterranean, we documented one adult male at S = 0.4 in a canal pertaining to the estuary of the Petit Rhône (Fig. 16; see material list above). Brun (1967) reported this species from the chlorinity range of 0.5–5‰ (salinity 0.9–9) in the estuary of the Grand Rhône. Wittmann & Ariani (2009) found it at S = 12.3 in the Canal d'Arles à Fos at a station within this estuary. Aguesse & Bigot (1960) reported N. integer together with Mesopodopsis slabberi from the Étang Baisse Salée (Camargue, Rhône delta). The salinities of this water body ranged from S = 0.8–40.1 (mostly 3–8) in 1955–1958. However, the exact range experienced by the mysids was not specifically indicated. For potential pathways of areal expansion to the Mediterranean, see ‘Discussion’. Discussion Taxonomy of the Mysinae tribes. The apparent morphological heterogeneity noted by Wittmann et al. (2014) for the tribe Mysini (subfamily Mysinae) is now reduced by detachment of the tribes Hemimysini and Paramysini. Both are characterized by a large smooth portion of the outer margin of the antennal scale, and distinguished from each other and from the remaining tribes of the Mysinae by features of male pleopods 3–5. The remainder after detachment of these two tribes is more homogeneous by showing an antennal scale setose all around, but remains heterogeneous in other important aspects. Among a total of ten genera within this remainder, four—Arthromysis, Parastilomysis, Hyperstilomysis, and Kainommatomysis—show a pair of plumose setae emerging from the telson cleft, a feature otherwise typical of the tribe Afromysini Wittmann, Ariani & Lagardère, 2014, within the subfamily Leptomysinae, and also often found in the subfamily Siriellinae. The 3-segmented carpopropodus of thoracic endopods 3–7 in Stilomysis, Parastilomysis, Hyperstilomysis, Kainommatomysis, and in certain species of Nanomysis is shared with many species of Leptomysinae. Upon first description of the genus Kainommatomysis, W. M. Tattersall (1927) discussed its great morphological affinities, regarding eyes and telson, with the Leptomysini (now Leptomysinae: Afromysini) genus Dioptromysis Zimmer, 1915, as a possible outcome of parallel evolution. In contrast, ribosomal gene sequence data in Porter (2005) suggest a rather close phylogenetic MYSIDAE IN MEDITERRANEAN FRESH-WATERS Zootaxa 4142 (1) © 2016 Magnolia Press · 47 relationship between Kainommatomysis and Dioptromysis despite their profound differences in male pleopods and resulting affiliation to different subfamilies. A more in-depth treatment of this enigma exceeds the limits of the present contribution. Within the tribe Paramysini, the monotypic Pontocaspian genus Katamysis bears certain outstanding characters. Its labrum has a long anterior spine, absent in all remaining genera of its tribe, but widespread among other Mysidae, for example in the tribe Neomysini and the subfamily Siriellinae. The telson of Katamysis is linguiform, with rounded posterior margin (similar to Siriellinae and Leptomysinae), while in other genera of its tribe it is more or less trapezoid with a cleft or with a truncated posterior margin. Although well fitting into its tribe in other respects, Katamysis would probably take a position somewhere close to the base of the group genealogy. Taxonomy of Mesopodopsis slabberi. Upon first description of Podopsis Slabberi, Van Beneden (1861) counted the genus Podopsis J. V. Thompson, 1829, among mysids, whereas Sars (1877) concluded that Thompson (1829) had actually described a macruran decapod. Sars proposed Macropsis G. O. Sars, 1877, as a replacement name at genus level, yet overlooked that this taxon was already preoccupied by the hemipteran genus Macropsis Lewis, 1834. Even more unfortunately, Czerniavsky (1882a, 1887) gave a completely new definition for the genus Podopsis by explicitly (1882a: 143) referring to crustaceans that differed strongly from those originally presented by Thompson. In the following years several attempts were made to repair the misidentification as Podopsis J. V. Thompson, and/or the synonymy with Macropsis Lewis: (1) upgrading of the subgenus Parapodopsis Czerniavsky, 1882a, to genus level (Butchinsky 1885, 1890; Nusbaum 1887; Kowalevsky 1889); (2) installation of the genus Leptocaris by Aurivillius (1898a, b) in order to allocate the species Slabberi Van Beneden; and (3) upgrading of the subgenus Mesopodopsis Czerniavsky, 1882a (Norman & Scott 1906, Norman, 1907). Ad (1), the genus name Parapodopsis was used for Black Sea populations mainly of the species P. cornuta (Czerniavsky, 1882a). It was almost never used for North Sea populations, except for the misspelled binomina Paropodopsis Göesii and Paropodopsis Slabberi by Sowinsky (1894). All these taxa are currently considered junior synonyms of Mesopodopsis slabberi. Ad (2), the binomen Leptocaris Slabberi was used only by Aurivillius (1898a, b). The otherwise disused Leptocaris Aurivillius, 1898, became a suppressed homonym of the later described copepod genus Leptocaris Scott, 1899, by precedence inversion according to ICZN (2000: case 3079). This decision has no effect on the status of Leptocaris Aurivillius, 1898, as a junior synonym of Mesopodopsis Czerniavsky, 1882a. Ad (3), Czerniavsky (1882a: 145–148) already used the binomen Mesopodopsis Slabberi, although he understood 'Mesopodopsis' at a subgeneric level. The use at generic level as Mesopodopsis slabberi started comparatively late (Norman & Scott 1906, Norman 1907, W. M. Tattersall 1922) due to the predominant use of the junior homonym Macropsis Sars up to the 1920s. The complex history of the taxon Mesopodopsis slabberi provides many examples that outdated synonyms or homonyms may persist in limited use over an additional 30–50 years. Taking this into account, most E-AtlanticMediterranean Mesopodopsis became well consolidated as the species M. slabberi in the 1930s up to the early 1990s. Following this, a caesura occurred when Wittmann (1992) described three new species by splitting off part of the south Mediterranean populations as M. aegyptia, West African as M. tropicalis, and South African as M. wooldridgei. Within the residual stock of M. slabberi, he found small, statistically overlapping morphological differences between NE-Atlantic, Mediterranean, and Black Sea populations. Remerie (2005) and Remerie et al. (2005) found high morphological diversity between NE-Atlantic and Mediterranean populations of M. slabberi, to some degree in concordance with genetic data. Sequencing of mitochondrial COI and 16S ribosomal RNA genes by Remerie et al. (2006) showed high diversity within and between NE-Atlantic and W-Mediterranean populations, indicative of cryptic speciation in M. slabberi. These findings suggest that the puzzling taxonomy of this species remains a challenge for future research. Distribution of Paramysis kosswigi. Băcescu (1948) gave a very short first description of Paramysis kosswigi from sources of the Great Maeander (= Büyük Menderes; Fig. 6) that drains into the Aegean Sea (EMediterranean). In 1954 he provided a more detailed description with figures and noted on page 103, lines 29–33 [transl.]: "Geographical distribution—Springs of the Great Maeander (Işliki in the south-west of Anatolia; col. Prof. C. Kosswig), nearly at the same latitude as the south of the Caspian Sea. Though located today in a river basin tributary of the Aegean Sea, Paramysis kosswigi is an authentic Ponto-Caspian relict of Miocene age, related to Paramysis lacustris turcica, that lives in Lake Beişehir, in the vicinity, in the full biocenoses of Dreissena 48 · Zootaxa 4142 (1) © 2016 Magnolia Press WITTMANN ET AL. polymorpha and European fish (Gobio gobio, Alburnoides punctatus). Its existence poses marked palaeogeographical problems". Komarova (1991) obviously referred to this text when she wrote on page 75, lines 21–25 [transl.]: "Distribution. On Turkish Black Sea shores, nearly at the same latitude as the south of the Caspian Sea. Although located in a river basin draining into the Aegean Sea, P. kosswigi is an indisputable Ponto-Caspian relict of Miocene age, a doubtless relative of P. lacustris turcica, which inhabits Lake Beişehir (Băcescu 1954)". She indicated no samples and no literature source other than Băcescu (1954). It is not clear if she had seen samples and/ or information about material from Turkish Black Sea shores. Due to a clear misunderstanding, Şerban et al. (2000) erroneously indicated in Tab. 1 (op. cit.) that Băcescu (1954) may have reported this species from the Black Sea, when in fact Băcescu did not do so. Şerban et al. (2001) claimed the absence of P. kosswigi from the Romanian part of the Black Sea. Without reference to any physical samples, Şerban (2004) and Skolka (2005) indicated a potential presence with query for the Black Sea. Without indication of species determination details, Akbulut et al. (2009) gave a first record for the Black Sea from brackish waters in the mouth of the Miliç River at the northern coast of Turkey. Excursions by one of us (KJW) to fresh and brackish waters of this coast in 1988 and in 2006 yielded only a closely similar species, P. lacustris (Czerniavsky, 1882), which in suboptimal material could be confounded with P. kosswigi by using the key of Băcescu (1954). According to our findings, P. lacustris is clearly distinguished from P. kosswigi by paradactylar setae of thoracic endopods 5–8 being 'serrated' over a longer portion of their length (see key to species below). In reasonably preserved material, it may also be distinguished by a less inflated dorsal surface of the head, and, according to Băcescu (1954), by eyes with more reniform cornea, and upon comparing specimens with expanded chromatophores, by a less intense, less dark pigmentation of the body. In conclusion, we consider the previous Black Sea records of P. kosswigi in part as being not clear or as misunderstandings, in part as based on outdated determination methods. This also makes the literature references to these records obsolete (see above for reference list of P. kosswigi). Areal expansion of Neomysis integer. Băcescu (1941) proposed two alternative pathways for the origin of the Neomysis integer population he detected in the Rhône Delta (NW-Mediterranean; Fig. 16): (1) the Rhône population originates from a transfer by ship through the Garonne River and the Canal-du-Midi from the abundant population in the Gironde estuary on the SW coast of France; and (2) immigration through the Strait of Gibraltar along the series of estuaries along the southern coasts of Spain and France. Kelleher et al. (1999) considered the Rhône population as disjunctive and, therefore, preferred alternative (1). Subsequent records by Munilla & San Vicente (2005) from the Mediterranean coast of Spain seemed to indicate a continuous population from the Atlantic to the Mediterranean along the coastline of Spain and, therefore, to support alternative (2) (Wittmann & Ariani 2009). Meanwhile, we were kindly informed by Jean Claude Sorbe (Arcachon; pers. comm. in Oct. 2008) and Carlos San Vicente (Creixell; pers. comm. in Sept. 2014) that the materials of Munilla & Corrales (1995) and Munilla & San Vicente (2005) were juveniles in poor condition, not determinable at species level. The cancellation of the previously supposed records at the Mediterranean coast of Spain makes the French population appear again as disjunctive, supporting alternative (1). Areal expansion of Limnomysis benedeni. This mysid was deliberately introduced into Lake Balaton, tributaries of the Baltic Sea, Lake Aral, and even in Turkmenistan to improve the food basis for fish (Woynárovich 1955, Mordukhai-Boltovskoi 1979, Leppäkoski 1984, Akmurdov et al. 2006). By artificial introduction into hydropower reservoirs of the Dnieper River and from there into reservoirs of the Neman River, and by subsequent expansion along waterways, it reached the Baltic coast at the Curonian Lagoon (Razinkovas 1996, Berezina et al. 2011). Additional, probably non-intentional, anthropogenic expansions proceeded from the lower to the upper reach of the Danube River and across the artificial Main-Danube-Canal down to the Main and Rhine Rivers. In 2007, Limnomysis had already reached fresh and brackish waters along the stretch of the Rhine from Lake Constance down to the North Sea (Wittmann & Ariani 2000, 2009; Bernauer & Jansen 2006; Fritz et al. 2006; Wittmann 2007; Lods-Crozet in CSCF 2008). The Odra estuary at the southern Baltic coast (Michels 2005, Wittfoth 2011) may have been reached via the Danube-Rhine drainage (Wittmann 2007, Wittmann et al. 2014). Mitochondrial DNA diversity suggests that this drainage was used for multiple invasion events (Audzijonyte et al. 2009). Additional expansions proceeded across the canal system in France to the Rhône River and therein down to its delta at the Mediterranean coast (Fig. 16), where Limnomysis was first detected in 2009 (Wittmann et al. 2014). The expansion to the Mediterranean is documented by part of the above sampling data. From species distribution MYSIDAE IN MEDITERRANEAN FRESH-WATERS Zootaxa 4142 (1) © 2016 Magnolia Press · 49 models, Gallardo & Aldridge (2013) derived a high potential of L. benedeni for future invasions of south-eastern parts of England and Ireland. Areal expansion of Hemimysis anomala. In order to enrich the food basis for fish, stocks from the Dnieper River were intentionally transplanted to the Kaunas reservoir in 1960 (Gasiunas 1968, Mordukhai-Boltovskoi 1979). From there the species expanded downstream to the Baltic Sea, where it widely dispersed in coastal waters up to Finland (Leppäkoski 1984, Salemaa & Hietalahti 1993, Arbačiauskas 2002, Berezina et al. 2011). Additional, probably non-intentional, expansions proceeded along two main routes (Audzijonyte et al. 2008b): a northern route from the Dnieper River to the Baltic Sea and from there up into the drainage system of the Rhine River, and a southern route from the Danube Delta up into Danube and across the artificial Main-Danube-Canal down to the Rhine Delta. These routes enabled it to reach fresh and brackish waters in vast areas of central and western Europe (Faasse 1998, Eggers et al. 1999, Wittmann 2007, Borza & Boda 2013, Golaz & Väinölä 2013). Mitochondrial DNA lineages from the Baltic and the one from the Danube River mixed secondarily in the Rhine Delta and in additional waters of NW-Europe (Audzijonyte et al. 2008b). The first record for the Mediterranean drainage was made in 2003 by Daufresne et al. (2007) in the middle course of the Rhône River. Already in 2007, H. anomala was found at many more stations within the Rhône system: in Lake Geneva (Lods-Crozet in CSCF 2008; Lods-Crozet et al. 2013) and along the rivers Saône and Rhône, down to brackish waters in the Rhône Delta at the Mediterranean coast of France (Fig. 2: Wittmann & Ariani 2009). Mitochondrial DNA sequencing by Golaz & Väinölä (2013) showed that the Lake Geneva population originates from the lineage that spread through the Danube to the Rhine. Overseas expansions led to limited areas in England (Stubbington et al. 2008) and Ireland (Minchin & Holmes 2008). From species distribution models, Gallardo & Aldridge (2013) derived a high potential of H. anomala for future invasions of additional areas covering most of England and almost entirely Ireland. Possibly ballast-water mediated, transoceanic expansion led to the Laurentian Great Lakes (Pothoven et al. 2007, Kipp & Ricciardi 2008) and additional waters in USA and Canada (Kestrup & Ricciardi 2008, Brooking et al. 2010, Brown et al. 2012). The invaders of England and in part of North America pertain to the Danubian lineage (Audzijonyte et al. 2008b, Brooking et al. 2010); but there are genes of Baltic lineages in North America as well (Questel et al. 2012). The above-listed samples taken in 2009 confirm the record by Hauer (2009) in Lake Traunsee, which is ca. 2201 km down the Traun and Danube Rivers from the Black Sea coast. They also corroborate records in the Marne River by Dumont & Muller (2009). Two of our samples represent the first records in the Seine River down to the Atlantic coast of France (Fig. 2). Finally, our 2009 samples complement previous findings (cited above) in Lake Geneva and along the Rhône down to brackish waters at the Mediterranean coast. Bioinvasion. Only three mysid species are so far known as non-indigenous inhabitants of fresh and oligohaline waters of the Mediterranean, altogether found on the Mediterranean coast of France: Neomysis integer since 1938/ 39 (Băcescu 1941), Hemimysis anomala since 2007 (Wittmann & Ariani 2009), and Limnomysis benedeni since 2009 (Wittmann et al. 2014). Neomysis is rather rarely found on the Mediterranean coast and, if so, in moderate numbers (see above), and may therefore be hardly termed an 'invader'. The two remaining species have both been found in inland waters of France since 1998 (Limnomysis) or 2005 (Hemimysis) (Wittmann & Ariani 2000, Dumont 2006). There they have meanwhile gained a wide distribution in the drainage systems of Rhine and Rhône Rivers, with the addition of H. anomala in the Seine. This and their potential ecosystem effects (below) justifies the term 'invaders'. The expansion dynamics are given above separately for H. anomala and L. benedeni, while potential expansion vectors and ecosystem effects of bioinvasion are discussed in the following: Potential vectors. The frequent occurrence of invasive freshwater mysids in European inland harbours suggests that the animals are carried by ship as the most important vector (Wittmann 1999). L. benedeni was found in hulls and in cooling water filters (Wittmann 1995, Reinhold & Tittizer 1997). The appearance of L. benedeni, H. anomala, and Katamysis warpachowskyi in drift samples points to a possible predisposition for dispersion by navigation (Wittmann et al. 1999). Gollasch (2007) concluded shipping as a key vector of species introductions into freshwaters of the Mediterranean. Overseas and transoceanic expansions occurred possibly in ballast water of transoceanic ships (Gollasch et al. 2002; Ellis & MacIsaac 2009; de Lafontaine et al. 2012). The mysids Mesopodopsis slabberi and Neomysis sp. were found in ballast water of such ships (Carlton 1985). Surprisingly, the sudden occurrence in inland waters of overseas areas also points to shipping traffic as a potential vector (Grigorovich et al. 2002, 2003; Ricciardi 2007). The appearance of mysids in poorly accessible or isolated lakes points to alternative ways of dispersion (Wittmann & Ariani 2009). Specifically, this involved systematic, 50 · Zootaxa 4142 (1) © 2016 Magnolia Press WITTMANN ET AL. intentional introduction of mysids as fish fodder, practiced mainly, but not exclusively, in Eastern European countries up to the 1980s. Examples for non-intentional introductions come from aquarist use, from overland transport of boats/ships containing ballast water, bilge water, and/or cooling water filters, from distribution of aquatic plants, and from inattentive stocking. Potential effects of bioinvasion. Stomach analysis by Borcherding et al. (2006) on H. anomala from a gravel pit lake connected to the Lower Rhine showed that larger specimens fed mainly on zooplankton, whereas smaller ones mainly on phytoplankton. Similar findings were made by Viherluoto (2001) on two species of Mysis from the Baltic Sea. In accordance with the mainly nocturnal activity of Hemimysis, the proportion of zooplankton consumed was highest during night and lowest during day. Feeding rates of this species on zooplankton of the Great Lakes of North America were in the order of 30–60% of body weight per day, depending on body size of the predator, light conditions, prey type, and prey density (Halpin et al. 2013). In a freshwater storage reservoir in The Netherlands, Ketelaars et al. (1999) observed a strong decline of Daphnia populations due to predation by recently invaded H. anomala. Wild carp taken in and near Vienna from waters along Danube River invaded by this mysid species showed markedly higher mercury levels compared to non-invaded waters (Wittmann et al. 2010). Ricciardi & Rasmussen (1998), Ricciardi (2007), and Kipp & Ricciardi (2008) forecasted food web disruptions and changes in nutrient and contaminant cycling as possible consequences of Hemimysis invasion in the Great Lakes. Koops et al. (2010) estimated the ecological risk of Hemimysis to the Great Lakes as high, and the risk to inland lakes as moderate to high. Dumont & Muller (2009) observed only minor impacts of H. anomala on the aquatic community of an Alsatian gravel pit lake (NE-France), in part due to decreasing population levels of the mysid during their three-year study. This shows the urgent need for long-term studies in view of the vast literature (based solely on short-term studies) assessing high ecological risk by H. anomala invasions. Unlike the generally more predatory H. anomala, the more day-time active L. benedeni is omnivorous, with strong herbivorous and detritivorous (Wittmann & Ariani 2000, Aßmann et al. 2009) but also carnivorous (Fink et al. 2012) behaviour. Fink & Harrod (2013) concluded from stable isotope analysis that carnivory and detritivory play an important role besides herbivory, depending on resource availability. According to Aßmann et al. (2009) this mysid also feeds on leaf litter and prefers different types of conditioned leaves, thus affecting leaf litter degradation in the invaded freshwaters. Outdoor mesocosm experiments by Fink et al. (2012) showed that the presence of phytoplankton lowered the predation pressure of L. benedeni on certain zooplankton taxa. Those authors concluded that the invasions by this mysid species may have complex impacts on the native zooplankton community and may potentially affect the whole aquatic ecosystem. Arbačiauskas et al. (2010) studied the effects of peracarid species introductions (in the 1960s) into Lithuanian lakes and water reservoirs to enhance fish production. Three species of amphipods and the mysids H. anomala, L. benedeni, and Paramysis lacustris were introduced. These species were found in fish stomachs but did not enhance fish production. Negative effects prevailed: changes in resident macroinvertebrate communities were induced and the local communities were detracted from naturalness. Biogeography of an- to oligohalobious mysids in the Mediterranean. The mysids considered here show quite different ecological ranges in the Mediterranean: In fresh-water only: Paramysis kosswigi, P. adriatica, Diamysis lacustris, Troglomysis vjetrenicensis. Freshwater, oligohaline, and beyond: Hemimysis anomala, Diamysis fluviatilis, D. mesohalobia heterandra, Limnomysis benedeni, Neomysis integer. Only oligohaline waters: Diamysis hebraica. Oligohaline and beyond: Mesopodopsis slabberi, Diamysis mesohalobia mesohalobia, D. mesohalobia gracilipes, D. lagunaris. Unlike the poly- to metahaline waters of the Mediterranean, the anhaline to mesohaline waters are inhabited by, if any, usually only one indigenous mysid species (Ariani & Wittmann 2004). These involve Mesopodopsis, Diamysis, Troglomysis, or Paramysis, or rarely a combination of 2 (3) species from different genera or by two different species of Diamysis. At the French coast we found Mesopodopsis slabberi frequently in combination with non-indigenous species, either Neomysis integer or Limnomysis benedeni. We expect increased frequency of such associations with the ongoing areal expansion by non-indigenous species. The available data basis is still insufficient to conclude any harm to indigenous taxa by this process. MYSIDAE IN MEDITERRANEAN FRESH-WATERS Zootaxa 4142 (1) © 2016 Magnolia Press · 51 The an- to oligohaline waters of the western Mediterranean yielded very few Diamysis populations (only two samples of D. lagunaris, Fig. 6). Here, the oligo- to mesohaline waters often yielded M. slabberi as the only mysid species (Fig. 2). From this, Ariani & Wittmann (2004) concluded that the Mediterranean forms (Wittmann 1992) of Mesopodopsis may be considered as vicariants of Diamysis in low salinity waters of the western Mediterranean, although co-occurring with several species of this genus in the eastern Mediterranean. Noteworthy in this context is that also the M. slabberi populations appear inhomogeneous: there are high levels of molecular diversity within and between Mediterranean populations (Remerie et al. 2006). From a strictly biogeographical point of view, the Mediterranean freshwater or oligohalobious Mysidae may be arranged in three major groups: A) indigenous Mediterranean species with detailed distribution range probably resulting from geohistorical events, in certain cases with ecological change towards less saline conditions; B) indigenous euryhaline Atlanto-Ponto-Mediterranean elements rarely found in freshwater or oligohaline environments; C) primary freshwater or oligohalobious (to mesohalobious) species that recently expanded outside their original geographical range by means of bioinvasion mechanisms (passive transport, man-made connections between waterways, etc.). Most species belong to group A, represented by eight species plus two non-nominotypical subspecies: Paramysis kosswigi, P. adriatica sp. nov., Diamysis fluviatilis, D. lacustris, D. mesohalobia mesohalobia, D. mesohalobia gracilipes, D. mesohalobia heterandra, D. lagunaris, D. hebraica, and Troglomysis vjetrenicensis. Only one species—Mesopodopsis slabberi—belongs to group B, based on its wide distribution plus several congeneric species in the E-Atlantic (Wittmann 1992) versus its absence in the Caspian and Aral Seas, on its general scarcity in fresh and oligohaline waters (see above), and on the precipitation of fluorite (CaF2) for the formation of statoliths (Ariani et al 1993; discussed below), which must be renewed upon each moult. Three species—Neomysis integer, Limnomysis benedeni, and Hemimysis anomala—are included in group C (see above for discussion of areal expansion in separate subchapters for each species). The present distribution of most species in group A, as well as of certain Mediterranean species of the genera Diamysis and Paramysis inhabiting more saline waters, can largely be explained in the light of two different kinds of geohistorical events. The more ancient, primary one, was recognized (Ariani 1981b; Wittmann & Ariani 1998, 2011; Ariani & Wittmann 2000) in the salinity crisis which affected the Mediterranean basin about 5.5 million year ago. The brackish waters of the Paratethys were drained into this basin, together with their faunistic elements (Hsü et al. 1977, Hsü 1978). Also certain Mysidae or their ancestors probably penetrated into the Mediterranean this way. This hypothesis is mainly based on the mineral composition of statoliths in all Mediterranean Diamysis and in almost all Mediterranean Paramysis species (Ariani et al. 1993; Ariani & Wittmann 2000, 2002; Wittmann & Ariani 1998, 2011, 2012a, b; present data). This includes P. kosswigi and now also P. adriatica sp. nov. These statoliths consist of calcium carbonate (CaCO3), as also found in the fossil ones from the Paratethyan Miocene (Voicu 1981) and in those from most Mysidae currently endemic to the Black Sea and the Caspian (residual Paratethyan basins). CaCO3 occurs as the crystal phase of vaterite in statoliths of living forms but as the crystal phase of calcite in fossil statoliths (Ariani et al. 1993). The stable calcite was probably formed from the metastable vaterite by phase transformation during or after the fossilization process (Ariani et al. 1993). In contrast to most species of the genera Diamysis and Paramysis, most of the remaining Recent Mediterranean Mysidae show fluorite (CaF2) statoliths, as do almost all Atlantic Mysidae examined in this respect (Ariani et al. 1993). This coincides with the fact that most Mediterranean faunal elements, including the above-discussed Mesopodopsis slabberi, are believed to have an Atlantic origin dating back to the early Pliocene flooding of the Mediterranean by Atlantic waters and to later events (Hsü et al. 1977, Taviani 2002, Emig & Geistdoerfer 2004). In line with this scenario, Băcescu (1940), following Ekman’s (1935) classification, envisaged a Mediterranean-Atlantic domain for a number of mysid species, and such biogeographical concepts appear still valid for the marine fauna in general (Gamulin-Brida & Span 1981). A Paratethyan origin of the Mediterranean Diamysis, as already hypothesized by Băcescu (1954) and Ariani (1981b), is supported not only by the evidence from biomineralogical data but also by the following points: (1) the extreme scarcity of Atlantic populations, whereas most other Mediterranean genera are mainly represented in the Atlantic; (2) the genus distribution roughly reminiscent (Ariani 1981b, Ariani & Wittmann 2000) of basins resulting from the reduction of the Paratethys and the formation of the Mediterranean Lago Mare (Hsü et al. 1977, Hsü 1978); (3) the ecology and distribution patterns of Diamysis in the Mediterranean, pointing to a colonization of this basin starting from the East, i.e. the above-cited Paratethyan drainage area(s), with adaptation to higher salinity 52 · Zootaxa 4142 (1) © 2016 Magnolia Press WITTMANN ET AL. upon expansion towards the West (Ariani & Wittmann 2004); 4) experimental results showing a higher survival of brood pouch larvae in three species of Mediterranean Diamysis at mesohaline compared with euhaline conditions, even in animals from populations living in marine or mixoeuhaline waters (Ariani & Wittmann 2000). The “regression model” proposed by Stock (1977, 1980) is considered by Wittmann & Ariani (2012b) to explain the following step from probably brackish (mesohaline?) towards the freshwater or oligohaline conditions characterizing certain Mediterranean Mysidae. In fact, the occurrence of all freshwater populations of Diamysis species (D. fluviatilis, D. lacustris, D. mesohalobia) so far known from the Mediterranean is restricted to tributaries of the Adriatic basin (Fig. 12), where transgressive-regressive events occurred in Pliocene-Pleistocene times. These events caused repeated alternations between marine and freshwater-terrestrial phases in the semi-enclosed Adriatic basin, for instance in the Venice area (Kent et al. 2002). The observed faunistic coincidence (detailed discussion in Wittmann & Ariani 2012b) can now certainly be extended to Paramysis adriatica sp. nov., and confirms the role of the Adriatic as the by far most species-rich drainage basin of fresh-waters inhabited by Mysidae within the Mediterranean (Fig. 12). This species richness appears mostly referable to effects of transgressive-regressive events, probably in combination with direct immigration of Ponto-Caspian elements, this last according to Băcescu (1985) and Por & Dimentman (1985) along hydrological connections between the Messinian Adriatic and the Paratethys reaching as far east as the Caspian and Aral basins. Bianco (1990) reported several emplacements of hypothetic ancient rivers where Paratethyan primary freshwater fishes might have survived. One of these emplacements roughly corresponds to Lake Scutari. This leads to an explanation of the distribution of D. lacustris along a drainage way from the Paratethys. This species is known only from Lake Scutari, and shows barbed setae on the penis instead of the exclusively smooth setae present in all other Diamysis species and subspecies examined in this respect (Wittmann & Ariani 2012b). Smooth setae are also present in Diamysis hebraica, inhabiting oligohaline waters on the Levantine coast of Israel (Fig. 6). Its actual distribution might be the result of a transgression-regression process, in analogy to the above hypothesis on Adriatic freshwater species. In fact, the sea deeply penetrated into the Levantine coast during the Lower Pliocene (Pasa 1953). Alternatively, a Levantine basin was already present much earlier in the Lago Mare and was directly derived from the Paratethys drainage (Hsü 1978). Among the freshwater mysids occurring in periadriatic localities (Fig. 12), Troglomysis vjetrenicensis shows a peculiar biogeographical situation. It is a troglobitic species today restricted to karstic cave waters more than 200 m above sea level, thus above the amplitude of the above-quoted transgression events. This species contributes, as a stenoendemic, to the outstanding biodiversity (Sket 1999) of the Vjetrenica cave in the Dinaric karst (BosniaHerzegovina). As possible explanations of this extraordinary biodiversity, Paar et al. (2014) stressed specific climatic and hydrological conditions in addition to lower glaciation levels in combination with diversified geomorphology and hydrology during the Pleistocene. Combined, these resulted in a great diversity of underground habitats. In biogeographical terms, T. vjetrenicensis can be regarded as a faunal element that survived in subterranean waters of the Dinaric karst like other endemics generally considered as living fossils. These include the serpulid Marifugia cavatica Absolon & Hrabĕ, 1930, and those few living species of Dreissenidae bivalves previously lumped together within the genus Congeria Partsch, 1835. The serpulid, recorded along with Troglomysis in Herzegovina, is thought to be a very ancient element of marine origin, whereby the area of origin does not coincide with the present Adriatic basin (Stammer 1935). The ancestors of Marifugia may have inhabited, like other endemic elements such as certain crustaceans, sea waters covering that region during the Cretaceous period (Remy 1937). This is reminiscent of a hydrographic situation preceding the Paratethys formation, i.e. pertaining to the Mesogeic Sea or Tethys. This points to an alternative scheme according to which also Troglomysis might have a Tethyan origin. This view is supported by the following considerations: 1) according to Băcescu (1954) Troglomysis may be an ancestor of Diamysis; with this relationship later regarded by Băcescu (1981) as incontestable; 2) Băcescu (1940, 1981) and Ariani (1981b) considered Diamysis or its immediate ancestors to be of Tethyan origin; 3) T. vjetrenicensis has fluorite statoliths as do D. frontieri H. Nouvel, 1965, from the Indo-Pacific, D. pengoi (Czerniavsky, 1882) from the Pontocaspian, and D. pusilla (G. O. Sars, 1907) from the Caspian (mineral composition of statoliths according to Ariani et al. (1993), and our own previously unpublished determinations in D. frontieri and D. pusilla). The latter three species are morphologically quite different from the Mediterranean Diamysis, mainly regarding the systematically important (Zimmer 1915a) fourth male pleopod. These species, particularly D. frontieri, may be assigned to different genera, thought to be derived together with Diamysis (sensu MYSIDAE IN MEDITERRANEAN FRESH-WATERS Zootaxa 4142 (1) © 2016 Magnolia Press · 53 stricto) and other closely related taxa (cf. Ariani 1981b) by splitting from Tertiary Tethyan ancestors. At any rate, D. pengoi and D. pusilla today inhabit basins directly derived from the more eastern portion of the Paratethys, in line with the hypothesis that ancient Tethyan elements could have survived the Mediterranean salinity crisis in the Paratethys and subsequently in the Black Sea (Por 1985, Por & Dimentman 1985). The ancestors of the fluorite(CaF2-) precipitating taxa may have survived outside the conditions by which certain lineages of mysids evolved (Ariani & Wittmann 2000) that today precipitate crystalline calcium carbonate (CaCO3 as vaterite) in their statocysts. In analogy to carbonatic statoliths as supposed indicators of freshwater or low-salinity ancestral environments, the evidence of fluorite statoliths suggests a marine origin. In fact, with the exception of a few species belonging to primitive taxa (Boreomysinae, Rhopalophthalminae) or living in deep-sea waters, which show non-mineralized (organic) statoliths, most marine mysids have fluorite statoliths (Ariani et al. 1993). Apart from its evolutionary meaning, the precipitation of large fluorite bodies upon each moult by freshwater animals such as T. vjetrenicensis and D. pengoi is highly surprising because of the high demand (Wittmann & Ariani 1996) versus poor availability of fluorine (Ariani et al. 1983) in their freshwater habitats. Key to the tribes of the subfamily Mysinae Haworth, 1825 Modified and updated from Wittmann et al. (2014) 1. 2a. 2b. 3a. 3b. 4a. 4b. 5a. 5b. 6a. 6b. 7a. 7b. 8. Mysidae with eyes well developed, eyestalks always separate, cornea rarely modified or reduced; antennal scale variable; females with two pairs of well-developed oostegites, rarely with three; pleomeres without projecting pleural plates; female pleopods reduced to simple, unsegmented, setose rods; male pleopods 1, 2 always uniramous, rudimentary as in females; male pleopod 4 and in certain taxa also pleopod 3, again less frequently also pleopod 5, biramous; both rami of the uropods undivided, exopods without spines, setose all around. Mysinae Haworth, 1825. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 Antennal scale setose except for a large proximal bare portion of the outer margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 Antennal scale setose all around . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 Bare portion of the outer margin of the antennal scale rostrally ending in a number of setae or in a subterminal series of articulate spines. Male pleopod 5 biramous with well-developed sympod . . . . . . . . . . . . . . . . . . . . . Hemimysini Czerniavsky, 1882 Bare portion of the outer margin of the antennal scale rostrally ending in a single articulate or non-articulate spine. Male pleopod 5 reduced to small, setose plate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Paramysini, new tribe Male pleopod 3 uniramous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 Male pleopod 3 biramous, with subdivided exopod . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 Male pleopod 3 reduced to small endopod fused with larger 2-segmented sympod. Telson shorter than ultimate pleonite, with short apical cleft or at least apically truncate . . . . . . . . . . . . . . . . . . . . . . . . . Diamysini Wittmann, Ariani & Lagardère, 2014. Male pleopod 3 reduced to small, setose plate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 Carpopropodus of thoracic endopod 6 with 4–20 segments; telson entire, linguiform to elongate subtriangular, its lateral margins with spines . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Neomysini Wittmann, Ariani & Lagardère, 2014. Carpopropodus of thoracic endopod 6 composed of only 1–3 segments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 Cornea bipartite, posterior part with large, backward-oriented ommatidium bearing a conical lens; telson with a pair of plumose setae at the bottom of its deep apical cleft (Mysini: Kainommatomysis) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 Cornea mostly normal, if bipartite then with small ommatidia only; telson devoid of setae or laminae; male pleopod 3 normally rudimentary as in females, rarely reduced to small endopod fused with sympod; uropods without spines; telson terminally rounded or with apical cleft . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Anisomysini Wittmann, Ariani & Lagardère, 2014. Carpopropodus of thoracic endopod 6 with 3–26 segments; statoliths composed of fluorite; telson variable, mostly with apical cleft . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Mysini Haworth, 1825 (see also ‘Discussion’). Key to the species of Mysidae in fresh and oligohaline waters of the Mediterranean All species indicated below belong to the subfamily Mysinae Haworth, 1825. Twelve species plus two nonnominotypical subspecies are so far known from fresh and oligohaline waters of the Mediterranean. In order to cover potential future invaders, this key includes three additional species that were previously encountered as invaders of warm temperate fresh-waters in any parts of Europe but so far not in the Mediterranean. 1a. 1b. 2a. Antennal scale setose all around . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 Antennal scale with setose margins except for a large proximal bare portion of the outer margin (and except for a thorn in certain genera) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 Outer margin of antennal scale with smooth portion ending in setae. Telson rhombohedral, terminally truncate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hemimysis anomala G. O. Sars, 1907 (Fig. 3) [expansive Ponto-Caspian endemic; the currently most expansive mysid species; invader of large parts of central, western, and 54 · Zootaxa 4142 (1) © 2016 Magnolia Press WITTMANN ET AL. 2b. 3a. 3b. 4a. 4b. 4c. 5a. 5b. 6a. 6b. 7a. 7b. 8a. northern Europe, in the Rhône system down to the Mediterranean coast (Fig. 2); overseas expansion to Ireland, England, and the Great Lakes of North America (USA, Canada); in fresh to brackish waters, sciaphilic, hiding on the substrate during daytime; in 0–10 m, down to 60 m depth] Outer margin of antennal scale with smooth portion ending in a thorn (= non-articulate spine). Telson variable. Katamysis, Paramysis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 Labrum with apical spine. Telson linguiform . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Katamysis warpachowskyi G. O. Sars, 1893 [expansive Ponto-Caspian endemic; indigenous in Caspian and Ponto-Azov basins; anthropogenic expansion to the mid and upper reaches of the Danube River and to Lake Constance; so far no Mediterranean records; mainly in rivers, also in lakes and estuaries, mainly in fresh-water, also in oligo- to mesohaline waters; mainly epibenthic, psammophilic, also among vegetation during daytime; in 0–4 m, down to 10 m depth] Labrum without apical spine. Telson terminally truncate or with terminal cleft. Genus Paramysis Czerniavsky, 1882 (Figs 4, 5, 7A, B, 8–10) [indigenous in the NE-Atlantic, Mediterranean, and Ponto-Caspian; anthropogenic expansion of several species to coastal waters and tributaries of the Baltic Sea, Lake Aral, and to inland water systems of northern and eastern Europe] . . 4 Telson terminally with strong, about rectangular cleft (Fig. 10U, V) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Paramysis (Longidentia) adriatica sp. nov. (Figs 7A, B, 8–10) [endemic in freshwater tributaries of the Adriatic Sea (Fig. 12); so far known only from five running waters and one lake, in each case less than 5 m above sea level; maximum sea distance 31 km; in 0–1 m depth among vegetation, also over bare muddy sediment] Telson with shallow cleft that forms an angle of >120°, bearing more than 15 laminae (Fig. 5J). Median segment of mandibular palp with at least some (mostly strong) serrated setae along its outer margin. Subgenus Serrapalpisis Daneliya, 2004 . . . . . 5 Telson without cleft; its wide, transverse terminal margin armed with numerous laminae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Paramysis (Mesomysis) intermedia (Czerniavsky, 1882) [expansive Ponto-Caspian endemic; indigenous in Caspian and Ponto-Azov basins; introduced into continental waters of eastern Europe, Uzbekistan, Kazakhstan, and Kirgizstan, particularly into lakes Aral, Balkhash, and Issyk-Kul; so far no Mediterranean records; in lakes, rivers, estuaries, mainly in fresh-water, also in oligo- to mesohaline waters; epibenthic, psammophilic; in 0–6 m, down to 18 (50) m depth] Paradactylar setae of thoracic endopods 5–8 'serrated' (by minute denticles) only along proximal half. Dorsal surface of carapace above head swollen (if well preserved). Eyes (if well preserved) with roughly hemispherical cornea in dorsal view (Fig. 4A) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Paramysis (Serrapalpisis) kosswigi Băcescu, 1948 (Figs 4, 5) [stenoendemic of the freshwater lake Işıklı in Anatolia, including its springs and its effluent Büyük Menderes (Great Maeander) to the Aegean Sea (E-Mediterranean), also found in the nearby system of the river Küçük Menderes (Little Maeander) (Fig. 6); up to 1007 m above sea level, epibenthic, psammophilic, in 0–1 m depth] Paradactylar setae of thoracic endopods 5–8 'serrated' over most of their length. Dorsal surface of carapace above head not swollen. Eyes (if well preserved) with about reniform cornea in dorsal view (as in Fig. 8B) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Paramysis (Serrapalpisis) lacustris (Czerniavsky, 1882) [expansive Ponto-Caspian endemic; indigenous in Caspian, Ponto-Azov, and Marmora basins, and in Lake Beyşehir in Anatolia; introduced into tributaries of the Baltic Sea and into waters of eastern Europe, Uzbekistan, Kazakhstan, and Kirgizstan, particularly lakes Aral, Balkhash, and Issyk-Kul; invader of parts of central and northern Europe, including the Gulf of Finland, so far no Mediterranean records; in lakes, rivers, estuaries, mainly in fresh-water, also in oligo- to mesohaline waters; mainly epibenthic, psammophilic, also among vegetation during daytime, may form dense aggregations close to the sediment; in 0–8 m, down to 48 m depth] Telson entire, subtriangular (Fig. 20M). Antennal scale terminally pointed in both sexes (Fig. 20B) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Neomysis integer (Leach, 1814) (Fig. 20) [NE-Atlantic endemic; indigenous in the E-Atlantic from N-Norway to Spain (Fig. 16), also in basins of the White Sea, Barents Sea, North Sea, and Baltic; probably non-indigenous at the Mediterranean coast of France; this coastal form penetrates deeply into river estuaries, populations with more than 100 km riverine distance from the sea may indicate anthropogenic dispersal; mainly in brackish water in 0–5 m depth, also marine in 3–30 m, down to 50 m depth, rarely in fresh-water] Telson rhombohedral to subrectangular, with apical cleft. Antennal scale terminally rounded at least in females, mostly also in males (pointed only in males of Limnomysis benedeni) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 Telson with large median lobe emerging from broad apical cleft. Eyestalks 3–4 times as long as cornea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Mesopodopsis slabberi (Van Beneden, 1861) (Fig. 1) [endemic in most European seas with exception of the Caspian; indigenous in the E-Atlantic from Scandinavia to Morocco, in the entire Mediterranean and the Black Sea (Fig. 2), including the North Sea, Baltic, and Marmora Seas; in marine to brackish coastal waters, found only at salinities of S >= 0.9 in the Mediterranean, but rarely though repeatedly found in true freshwaters pertaining to the Black Sea and NE-Atlantic; mostly in shallow water, often in masses in the brackish water off river mouths; may gather in large swarms that stay or move along closely over the substrate, such swarms disperse during the night] Telson without median lobe. Eyestalks less than 3 times length of cornea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 Apical segment is 25–30% antennal scale length (Fig. 19D, E). Scale terminally rounded in females (Fig. 19D), pointed in males (Fig. 19E). Exopod of fourth male pleopod 3-segmented, with only minute setae on penultimate (median) segment (Fig. 19H) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Limnomysis benedeni Czerniavsky, 1882 (Fig. 19) [expansive Ponto-Caspian endemic; indigenous in Caspian, Ponto-Azov, and Marmora basins (Fig. 16); introduced into Lake Balaton, tributaries of the Baltic Sea, Lake Aral, waters of Turkmenistan; invader of large parts of eastern, central, western, and northern Europe, coastal waters of the NE-Atlantic and the Baltic, in the Rhône system down to the Mediterranean coast; mainly in fresh-water, also brackish; mainly among vegetation during daytime, rarely observed to form swarms; in 0–2 m, MYSIDAE IN MEDITERRANEAN FRESH-WATERS Zootaxa 4142 (1) © 2016 Magnolia Press · 55 8b. 9a. 9b. 10a. 10b. 11a. 11b. 12a. 12b. 13a. 13b. 14a. 14b. 14c. down to 30 m depth] Apical segment is <=20% antennal scale length (Figs 13A, E, 14A, 15A, K). Scale terminally rounded in both sexes. Exopod of fourth male pleopod 2–3-segmented, with a medium-sized to large seta (Figs 13G, 14H, 15N, 17E, 18N) on penultimate segment. Troglomysis, Diamysis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 Eyes without pigment, cornea rudimentary (Fig. 18A–C). Seta from penultimate segment of exopod of fourth male pleopod extends far beyond the seta of the apical segment (Fig. 18N) . . . . . . Troglomysis vjetrenicensis Stammer, 1933 (Figs 7C, 18) [stenoendemic of subterranean fresh-waters in the system of the karstic cave Vjetrenica (Fig. 12) in Herzegovina, 12 km from the coast of the E-Adriatic Sea, at around 234 m above sea level] Eyes normal. The largest seta (mostly only one present) of the penultimate segment of exopod of fourth male pleopod is much shorter than the seta of the apical segment (Figs 13G, 14H, 15E, N, 17E). Genus Diamysis Czerniavsky, 1882 (Figs 13–15, 17) [mainly Mediterranean, also NE-Atlantic, Pontocaspian, and W-Indian Ocean; in coastal to continental waters, mostly in brackish waters, also marine, 3 out of 14 species known live mainly in fresh-water, an additional one to a minor extent in freshwater] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10 Rostrum angular with narrowly rounded tip (Fig. 13E, F). Carapace of males without fringes (Fig. 13F) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Diamysis lacustris Băcescu, 1940 (Fig. 13D–H) [stenoendemic of the freshwater Lake Scutari at the SE-coast of the Adriatic Sea (Fig. 12), altitude 5 m; sea distance of the lake >= 43 km along its effluent] Rostrum well rounded or forming a wide convex angle with broadly rounded tip. Carapace of males with or without fringes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11 Basal segment of thoracic exopods 1–8 terminally well rounded (Fig. 17M). Carapace of males without fringes (Fig. 17K) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Diamysis hebraica Almeida Prado-Por, 1981 (Fig. 17K–Q) [stenoendemic at S = 0.7–5 in brackish streams at the Mediterranean coast of Israel; Fig. 6] Basal segment of thoracic exopods 1–8, or at least in exopods 1–3, terminally ending in a spiniform projection (Fig. 14D). Carapace of males with or without fringes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12 Rostrum well rounded (Fig. 13A). Male carapace with fringes arranged in two submedian stripes (Fig. 13B) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Diamysis fluviatilis Wittmann & Ariani, 2012 (Fig. 13A–C) [in wells, springs, tributaries, canals, main courses, and estuaries of rivers flowing to the NE to NW coasts of the Adriatic Sea (Fig. 12); maximum observed sea distance 182 km, maximum altitude 16 m; mostly in fresh-water, also oligohaline to mesohaline (S = 0–13)] Rostrum less continuously rounded, forming a wide convex angle with rounded tip (Fig. 14A, C) . . . . . . . . . . . . . . . . . . . . 13 Eyestalks dorsally with fenestra paracornealis well developed (Fig. 17A, B), although not well visible in poorly pigmented eyestalks. Scutellum paracaudale well rounded (Fig. 17F) or biconvex with rounded (rarely acute) apex (Fig. 17G, H). Carapace without fringes in both sexes (Fig. 17A) . . . . . . . . . . . . . . . . Diamysis lagunaris Ariani & Wittmann, 2000 (Fig. 17A–J) [W- and E-Mediterranean, NE-Atlantic coasts of Portugal and southern Spain (Fig. 6); in oligo- to metahaline waters, mainly in poly- to mixoeuhaline lagoons, also coastal marine] Eyestalks with fenestra paracornealis poorly developed (Fig. 15K), mostly missing (Figs 14A, 15A). Scutellum paracaudale subtriangular, with acute (Fig. 15F–H) or rounded (Fig. 14J, M) apex. Carapace of males with or without fringes. Diamysis mesohalobia Ariani & Wittmann, 2000 (Figs 14, 15; species sensu lato; key to subspecies updated from Wittmann & Ariani, 2012a) [marine, brackish and fresh-waters of the E-Mediterranean and Marmora Seas; Fig. 16] . . . . . . . . . . . . . . . . . . . . . 14 Carapace without fringes in both sexes (Fig. 14C). Pereiopods relatively short, endopod 8, when stretched anteriorly, extending to basis of endopod 1 or up to mandibles. Pereiopods stout to moderately slender, with R6 = 3.9–6.8. Thoracic endopod 5 with 3- or less frequently 2-segmented carpopropodus. Exopod of male pleopod 4 always 2-segmented, with only one smooth seta on basal segment (Fig. 14H) . . . . . . . . . . . . . . . . . . . Diamysis mesohalobia mesohalobia Ariani & Wittmann, 2000 (Fig. 14) [Adriatic, Aegean, and Levantine seas (Fig. 16), in mesohaline karstic springs and in meso- to (mixo)euhaline lagoons and estuaries, also known from the oligohaline reach in a lagoon with complex salinity patterns] Carapace without fringes in both sexes (Fig. 15A). Pereiopods long, thoracic endopod 8, when stretched anteriorly, extending up to labrum or even eyes. Pereiopods slender, with R6 = 5.6–9.1. Thoracic endopod 5 with 2- or rarely 3-segmented carpopropodus. Exopod of male pleopod 4 normally 2-segmented, with smooth seta at basal segment (Fig. 15D); or occasionally 3-segmented in large males, with distally barbed seta at median segment (Fig. 15E) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Diamysis mesohalobia gracilipes Ariani & Wittmann, 2000 (Fig. 15A–J) [Adriatic and Ionian seas (Fig. 16), in marine embayments and in meso- to mixoeuhaline lagoons and estuaries; also known from one oligohaline spring] Carapace of males with numerous fringes arranged in at least two submedian stripes and in a traverse row shortly in front of the posterior margin (Fig. 15K, L). Pereiopods of intermediate length, endopod 8, when stretched anteriorly, extending to maxillae or at most to labrum. Pereiopods stout to slender, with R6 = 4.5–7.9. Thoracic endopod 5 with 2- or less frequently 3segmented carpopropodus. Exopod of male pleopod 4 always 2-segmented, with smooth seta subterminally on basal segment (Fig. 15O), in large males often with additional smooth and/or barbed setae terminally on this segment (Fig. 15N, P) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Diamysis mesohalobia heterandra Ariani & Wittmann, 2000 (Fig. 15K–S) [in anhaline to slightly metahaline waters of springs, estuaries, lagoons, and lakes all around the Adriatic Sea (Fig. 12), whereas only in oligo- to polyhaline waters on the coasts of the Ionian and Marmora Seas; (Fig. 16)] 56 · Zootaxa 4142 (1) © 2016 Magnolia Press WITTMANN ET AL. Acknowledgements Sincere thanks are due to Cem Aygen (Izmir), Karin Kronestedt (Stockholm), Davor Lučić (Dubrovnik), Boris Sket (Ljubljana), Jean Claude Sorbe (Arcachon), Fabio Stoch (Trieste), and Pero Tutman (Split) for providing materials for examination. We were kindly informed by Wolfgang Heimler that Stammer's (1932) material is not in the Stammer collection kept by the Department of Biology, University of Erlangen (Germany). Jean Claude Sorbe (Arcachon) and Carlos San Vicente (Creixell) kindly informed us about samples of Neomysis integer. We are also indebted to Iorgu Petrescu (Bucharest) for checking our translations of Romanian texts, to Victor Petryashov (St. Petersburg) for help with Ukrainian literature, and to Risto Väinölä (Helsinki) for depositing specimens at the Finnish Museum of Natural History. References Aguesse, P. & Bigot, L. (1960) Observations floristiques et faunistiques sur un étang de Moyenne Camargue: la Baisse Salée de la Tour du Valat. Vie et Milieu, 11 (2), 284–307. Akbulut, M., Çelik, E.Ş., Odabaşi, D.A., Kaya, H., Selvi, K., Arslan, N. & Odabaşi, S.S. (2009) Seasonal distribution and composition of benthic macroinvertebrate communities in Menderes Creek, Çanakkale, Turkey. 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