Mediterranean Marine Science

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Vol. 8, 2007

The freshwater ichthyofauna of Greece - an update based on a hydrographic basin survey

ECONOMOU A.N. Hellenic Centre for Marine Research, Institute of Inland Waters, P.O. Box 712, P.C. 19013, Anavyssos, Attiki GIAKOUMI S. Hellenic Centre for Marine Research, Institute of Inland Waters, P.O. Box 712, P.C. 19013, Anavyssos, Attiki VARDAKAS L. Hellenic Centre for Marine Research, Institute of Inland Waters, P.O. Box 712, P.C. 19013, Anavyssos, Attiki BARBIERI R. Hellenic Centre for Marine Research, Institute of Inland Waters, P.O. Box 712, P.C. 19013, Anavyssos, Attiki STOUMBOUDI M.ΤΗ. Hellenic Centre for Marine Research, Institute of Inland Waters, P.O. Box 712, P.C. 19013, Anavyssos, Attiki ZOGARIS S. Hellenic Centre for Marine Research, Institute of Inland Waters, P.O. Box 712, P.C. 19013, Anavyssos, Attiki http://dx.doi.org/10.12681/mms.164

To cite this article:

ECONOMOU, A., GIAKOUMI, S., VARDAKAS, L., BARBIERI, R., STOUMBOUDI, M., & ZOGARIS, S. (2007). The freshwater ichthyofauna of Greece - an update based on a hydrographic basin survey. Mediterranean Marine Science, 8(1), 91-166. doi:http://dx.doi.org/10.12681/mms.164

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Mediterranean Marine Science Volume 8/1, 2007, 91-166

The freshwater ichthyofauna of Greece - an update based on a hydrographic basin survey

A.N. ECONOMOUÅ, S. GIAKOUMIÅ, L. VARDAKASÅ, R. BARBIERIÅ, M. STOUMBOUDIÅ and S. ZOGARISÅ,Ç

ÅHellenic Center for Marine Research, Institute of Inland Waters, 46,7 km Athens-Sounio, 19013 Anavissos Attiki, Greece

Ç University of Ioannina, Dept. of Environment and Natural Resources Management, Laboratory of Ecology & Biodiversity Conservation, 2 Seferi St., 30100 Agrinio, Greece e-mail: [email protected]

Abstract

Distribution records (historical, contemporary) for native and non-native freshwater fish species from 105 hydrographic basin areas were compiled and analysed in order to develop a nation-wide inventory (including transboundary river basins). Overall, 162 species, including diadromous and euryhaline, with documented occurrence records in freshwaters, and taxa of unclarified taxonomic status, are accommodated in the distributional compilation. An annotated checklist summarises the confirmed ichthyofauna of Greek freshwaters (161 species); a provisional supplementary list contains species recorded in brackish waters (55 species). In comparison to the last published (1991) checklist of freshwater fish of Greece, the present checklist shows an increase in species number of 53% (56 species). This increase has resulted mainly from taxonomic re-evaluations of existing taxa on the basis of new information and adoption of a new systematic concept. The current trend, as reflected in recent ichthyological publications, is towards abandonment of the biological species concept (BSC) and adoption of the phylogenetic species concept (PSC) for the delineation of species boundaries. The practical implications of the change in species concept on biodiversity conservation and watershed management are discussed. An overview of the composition and characteristics of the freshwater fish fauna of Greece is provided, especially with regard to the native and introduced status of species, and the spatial patterns of species richness and endemism. This systematic inventory may assist in efforts to develop nation-wide surface water bioassessment tools within the demands of the Water Framework Directive (WFD); it may further promote biodiversity conservation and biologically-orientated fishery management approaches.

Keywords: Freshwater ichthyofauna; Fish distribution; River basin area; Endemism; Biodiversity; Greece.

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http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 09/03/2019 12:43:28 | Introduction ichthyological research over the past decades, from distributional surveys and Freshwater fish represent an impor- taxonomic work based on morphology, to tant component of the aquatic ecosystem studies of fish physiology, ecology, biology and are highly valued for their economic, and genetics. This change had a negative social and aesthetic importance. Fish are side-effect on the availability of fish distri- already involved in environmental policies butional data. Large-scale distributional as biodiversity and ecological quality indi- survey work was abandoned in favour of cators (KESTEMONT et al., 2000; regional or local scientific investigations, SCHMUTZ et al., 2007) and they have often focusing on single species or single been used successfully in biogeographical areas. It is characteristic that most of the studies (B N RESCU, 2004), ecore- information used to produce the checklist gion delineations (HAWKES et al., 1986; of freshwater fish of Greece published by ABELL et al., 2002), conservation evalua- ECONOMIDIS (1991) was derived from tions (MOYLE & RANDALL, 1998) and his earlier published catalogue of fish assessments of ecologically acceptable (ECONOMIDIS, 1973), which is based on water regime management (JOWETT, results of investigations undertaken before 1997). Greece has diverse inland water the 1970s. resources and hosts one of the richest According to the aforementioned freshwater ichthyofaunas in Europe. The review (ECONOMOU et al., 2004a) valu- number of species occurring in Greek able ichthyological information exists for waters is still an issue of active investiga- large rivers and lakes, however, the fish tion. The last published checklist of the fauna of small aquatic systems has been freshwater fish of Greece contains 105 poorly investigated. For some areas no native and introduced species (including information at all could be found, whereas euryhaline) plus five of doubtful occur- for other areas there are data shortages for rence (ECONOMIDIS, 1991). A recent most, except the larger water bodies. An re-evaluation gives 135 species, of which associated problem is that the distribution 89 are native exclusively freshwater, and 54 of alien fish is poorly known since species are endemic to the country or to the south- introductions or translocations are rarely ern Balkans (BOBORI & announced, and when they are, the data ECONOMIDIS, 2006). However, a com- are not always accessible. Quite often, plete nation-wide inventory of fish occur- information on the occurrence of non- rences by river basin has never been pub- native species can be obtained only from lished. grey literature or local authorities. A risk A review of publications concerning of misreporting and misidentification is the freshwater fish of Greece was recently inevitably inherent in this approach as the undertaken with the scope of assessing the taxonomy of the introduced species is not utility of available ichthyological informa- often verified by experts. Moreover, much tion for the implementation of the EU of the available information on species dis- Water Framework Directive tributions is spread out in various, often (ECONOMOU et al., 2004a). One of the obscure, publications which are not readi- outcomes of this review is that there has ly accessible to the scientific community, been a change in the thematic focus of especially to non Greek-speaking scien-

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http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 09/03/2019 12:43:28 | tists. This holds particularly true for disser- develop an updated and, to the degree that tation theses, works published in older or it is possible, complete inventory of the local journals, national conference pro- distribution of freshwater fish over Greek ceedings and project reports. It is obvious territory. Using a combination of data that a synthesis and compilation of rele- sources and historic or contemporary vant information is required to facilitate accounts, we compiled lists of the freshwa- knowledge access and sharing. ter fish occurring in 105 Greek hydro- Deficiencies in distributional data graphic basin areas and we characterized availability, coupled with taxonomic ambi- their provenance status (i.e. native or guities, have a negative impact on scientif- introduced in each basin area). The hydro- ic research and management applications graphic basin area was chosen as the spa- dependent on fish data. For example, plan- tial unit for this inventory because of its ning for biodiversity conservation in relevance to biogeography. Indeed, no sin- Greece is often hindered by lack of knowl- gle factor is more important in explaining edge of the distributional ranges of endan- large-scale distributional patterns of fresh- gered species and vagueness in ichthyolog- water fish than hydrographic basin limits ical nomenclature. These problems reflect (GILBERT, 1980). River basins have long the fact that the list of protected fish been considered as the operational unit for species for Greece included in the Annex- the biogeographical analysis of fish es of the Habitats Directive (92/43/EC) throughout the world (e.g. ABELL et al., contains species listed under invalid 2002; HIGGINS et al., 2005; THIEME et names, while many really threatened al., 2005; REYJOL et al., 2006). This is species are not listed. Recent taxonomic because both primary and many secondary revisions undertaken on a European scale freshwater fish are freshwater-obligate and the application of new criteria for organisms that cannot readily disperse defining species (KOTTELAT, 1997; across terrestrial boundaries or marine KOTTELAT & FREYHOFF, 2007a) areas. Effectively, the freshwater fish pop- have attempted to resolve problems relat- ulations within hydrological basins are ed to nomenclature and systematics but at reproductively isolated entities adapted to the same time they have complicated the their basin-scale ecosystems. River basins ichthyological faunal list of Greece by often exhibit a distinct fish composition renaming and ‘splitting’ many species. and can be seen as ‘biogeographical Thus, much confusion may be generated islands’ containing a specific pool of among environmental managers and non- species (HUGUENY, 1989). This island- specialist users of ichthyological data, as like character also renders the basin area some species appear under new names, an appropriate scale for studies of several new species have emerged, and the endemism and speciation (PETER, 2006). distributional ranges of many species are The main product of this work is a not clearly defined. This again reiterates hydrographic basin area-based compila- the need for a systematic treatment of tion of fish that live in the freshwaters of existing fish distributional records to pro- Greece. These data may assist in under- duce hydrographic basin area compilations standing regional assemblage structure incorporating these changes. and other faunistic attributes of relevance The aim of the present work is to to biogeography. In addition, the data are

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http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 09/03/2019 12:43:28 | of potential utility in resource and envi- last century (OECD, 2000) and many ronmental management, especially with aquatic ecosystems are threatened even in regard to ecological quality assessments, designated protected-areas. Data show biodiversity conservation and fishery that several fish populations have become exploitation. threatened or extirpated (ECONOMIDIS, From the perspective of ecological 1995, 1999, 2002; ECONOMIDIS et al., quality assessment, interest is centered on 1996; ECONOMOU et al., 1999; the successful implementation of the EU STOUMBOUDI et al., 2002; Water Framework Directive 2000/60/EC PERDIKARIS et al., 2005) and many (WFD). The WFD establishes an ecosys- more may face threats in the near future tem-based policy framework for water due to escalating pressures upon surface management and protection in Europe at freshwater resources. Organization of the the river basin scale. The Directive ichthyological information on a basin area demands, among others, that the member scale provides a practical approach for the states should develop monitoring pro- evaluation of conservation priorities and grammes and ecological status classifica- the design of restoration measures. Knowl- tion systems within river basin districts edge of fish distributions may also facili- using fish and other organisms as biologi- tate the monitoring of changes in the fish cal indicators. An important requirement faunas in relation to climatic change. for the establishment of the monitoring Lastly, freshwater fishing has a strong programmes is that reliable information socio-economic dimension, especially in on freshwater fish assemblages for each rural areas, where alternative employment river basin area is available, both for the possibilities are limited or highly seasonal. characterization of the undisturbed, type- The freshwater fisheries’ resources are specific ‘reference conditions’ and for the threatened by destructive or illegal fishing selection of the appropriate biological and, increasingly in recent years, by the metrics with which to measure ecological uncontrolled introduction of alien species degradation (ECONOMOU, 2002; (CRIVELLI et al., 1997; ECONOMIDIS ECONOMOU et al., 2003). et al., 2000a; ECONOMOU et al., 2001a). With regard to biodiversity conserva- The present compilation may provide pol- tion, the emphasis is on identifying areas icy-relevant information on the distribu- hosting species in need of protection tion status of native, exotic and translocat- according to the EU Habitats Directive ed species that may help in designing bio- (92/43/EC). Incomplete and/or inaccessi- logically-based fisheries management. ble information on fish species’ distributions has hindered the timely evaluation of Study Area areas of high conservation value and their inclusion in the NATURA 2000 protected- Greece’s land area (132 000 km2) sup- area framework. This is unfortunate, as ports remarkably varied inland water fea- Greece is one of Europe’s biodiversity tures. The country has a highly fragmented hotspots in terms of freshwater fish geography and various parts of the territo- endemism (MAITLAND & CRIVELLI, ry have very different environmental con- 1996). The country has already lost nearly ditions and biogeographical histories. 75% of its natural wetland areas during the 65.4% of the land exceeds 200 m in alti-

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http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 09/03/2019 12:43:28 | tude (CATSADORAKIS, 2003). The was being affected by the Pleistocene coastal morphology both on the continent glaciations (SKOULIKIDIS et al., 1998; and on the islands is diverse but it usually PERISSORATIS & CONISPOLIATIS, consists of a highly fragmented, narrow 2003). coastal zone with a varied relief. The most A proper classification of Greece’s extensive lowland areas exist in northern surface water systems has never been com- Greece where relatively large alluvial pleted and the variety of aquatic ecosys- plains are drained by the rivers Evros, tems and habitat types is certainly Nestos, Strymon, Axios and Aliakmon. immense. These water features range from In terms of inland water aquatic biota, large transboundary rivers, medium and Greece is at a biological crossroads among small perennial and intermittent streams Mediterranean, temperate European, to small endorheic karstic streams, spring Danubian-Black Sea, and Anatolian influ- systems, inland lakes, coastal lagoons, ences (B N RESCU, 2004). Although swamps and marshes. Water features have the biotic influences due to the country’s been poorly inventoried for their aquatic geography are unique, important biogeo- biota and little published information graphic barriers criss-cross the country and exists on regional and local scale species create even more heterogeneity. Long distributions; data for invertebrates and mountain chains, climatic rain-shadows, lower vertebrates is especially lacking wide marine gulfs and deep seas that were (LEGAKIS, 2004). Until recently, formal not drained due to sea level lowering at the inventory procedures have catalogued only glacial maxima, create inland water isola- the large wetland sites; one such compilation which has remained rather stable for tion lists 378 wetland sites covering millennia. More geographic idiosyncrasies approximately 2000 km2 in the entire are created by the country’s geological country (ZALIDIS & MANTZAVELAS, fragmentation and dynamics, its extensive 1994). Unfortunately, these published cat- coastline (16000 km), and a variety of cli- alogues of sites are a poor and incomplete matic zones. Climatically, Greece main- representation of the total number of tains remarkable extremes ranging from aquatic sites in the country (e.g. high-rainfall mountains receiving 2400 mm CATSADORAKIS & PARAGAMIAN, precipitation per year (Central Pindos), to 2006). In a recent regional-based survey, seasonally semi-arid areas (southern Atti- CATSADORAKIS & PARAGAMIAN ki) receiving less than 400 mm. Rain-shad- (2007) describe 352 wetland areas solely in ow areas in the southeast mainland and the Aegean islands (excluding Crete). the Cyclades islands create pockets of seasonally arid conditions with high evapo- Profile of Greek hydrographic basins transpiration rates and a long summer drought. Unlike most European countries It has been recently stated that limnol- Greece’s inland aquatic ecosystems are ogy and freshwater ecology in the Mediter- strongly characterized by geographical iso- ranean should not be based on temperate lation, both spatially as small river basin European paradigms, patterns and gener- areas, and temporally as long-standing rel- alizations (ALVAREZ-COBELAS et al., atively stable waterbodies which created 2005). Greek river basins differ markedly biotic refugia when much of the continent from most temperate European ones.

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http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 09/03/2019 12:43:28 | They usually comprise isolated hydro- lagoons; the number of these small water graphic basins, characterized by short, features is certainly in the thousands. Some steep fluvial systems that exhibit very ero- of these water features are associated with sive behavior, flashy, irregular flow lowland rivers and lowland or plateau lake- regimes and are influenced by varied geo- swamp environments. In contrast, many graphical, geological and climatic condi- small karstic spring-fed lentic systems espe- tions. Most rivers run through narrow cially in the limestone-dominated southern mountain valleys and descend abruptly to and western parts of the country are often the coast, usually lacking extensive lowland totally isolated from other surface-flowing sections and floodplain habitats. There are waters in the wider landscape. only eight large rivers in the country (with drainage areas larger than 6000 km2), Methods including five transboundary ones (SKOULIKIDIS et al., 1998). The number Our inventory approach takes the fol- of smaller rivers or streams with perennial lowing steps: a) selection of adequately- flowing segments is certainly in the hun- studied hydrographic basin areas (and dreds; a complete inventory of all hydro- other ‘isolated’ aquatic sites); b) compila- graphic basin delineations does not exist, tion of fish species occurrence for each despite attempts in the recent past (e.g. area; and, c) compilation of a provisional NTUA, 1994). ‘Lowland’ fluvial habitat annotated freshwater fish species checklist conditions are mainly encountered in the and a supplementary list of species record- plateaus and inland plains of the larger ed in transitional waters. rivers, which create unique biotic assemblages since their access to the river’s main Definition of studied basin areas stem may be blocked by gorges or other natural barriers including karstic phenom- The basic geographical unit to assem- ena. The deltas of Greek rivers are often ble our distributional compilation is extensive, although they are not affected defined as the hydrographic basin area. by estuarine tidal regimes (TSIOURIS & Only basin areas where documented fish GERAKIS, 1991). species occurrences have led to a ‘near- Most lakes in Greece are located with- complete’ ichthyofauna list are included in in river basins or have a historic relation the compilation. The size of each basin with their wider river basins. Some of the area was retrieved largely from the Hydro- older lakes have a unique history of isola- scope database of the National Technical tion and are centers of endemism (e.g. University of Athens (NTUA, 1994). This Lakes Prespa, Vegoritis, Pamvotis, Doira- geo-database delineates 737 river basins ni) (FROGLEY et al., 2001; GRIFFITHS and ‘wider river basin areas’ in Greece. et al., 2002; FROGLEY & PREECE, The NTUA delineations do not include 2004; ALBRECHT et al., 2007). Although some of the smaller hydrographic basin 56 major natural lakes have been areas we provide data for; their area was described (ZALIDIS & MANTZAVE- roughly estimated by us from maps (Table LAS, 1994; ZACHARIAS et al., 2002), 1). Each hydrographic basin area was cate- Greece has many smaller lentic bodies, gorized into one of six general biogeo- such as various wetlands, ponds and coastal graphical units, these roughly follow

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http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 09/03/2019 12:43:28 | Table 1 Hydrographic basins for which fish data are presented in this study. Basins are numbered according to their position on the map (see Fig. 1). For each basin the biogeographical region, the basin surface area and the included water bodies are given.

Estimated Biogeo-graphical No Name Given Area Included water bodies Area (km2) 1 Evros North Aegean 53000 Evros R. (Maritsa, Meric), Loutros R., Delta and lagoons. 2 Avas North Aegean 249 Avas R. (also known as Potamos R.) Filiouris R., Bospos R., Mitrikou (or Ismarida) L.; 3 Filiouri North Aegean 2107 adjacent coastal lagoons and Maronia R. 4 Kompsatos North Aegean 596 Kompsatos R. 5 Vistonis North Aegean 3200 Vistonis L., Porto Lagos Lagoons. 6 Kossinthos North Aegean 435 Kossinthos R. Laspias (or Laspopotamos) R., Avdira wetlands and 7 Laspias North Aegean 138 surrounding wetlands and lagoons. 8 Nestos North Aegean 6200 Nestos (Mesta) R., reservoirs, Delta wetlands and lagoons. 9 Marmaras North Aegean 235 Marmaras R. 10 Nevrokopi North Aegean 473 Streams in Nevrokopi basin. 11 Strymon North Aegean 17000 Strymon (Struma) R., Kerkini L., reservoirs. 12 Ladopotamos North Aegean 25 Ladopotamos, Agion Oros Peninsula. 13 Mavrolakas North Aegean 80 Mavrolakas R. 14 Asprolakas North Aegean 91 Asprolakas R. 15 Rihios North Aegean 2090 Rihios R. Volvi L., Koronia (or Aghiou Vassileiou or Lagada) L., 16 Volvi North Aegean 1903 tributary streams. Doriani (Dorjan) L. (within the broader basin of the Axios 17 Doirani North Aegean 420 R.), Megalo Rema and other tributaries. 18 Axios North Aegean 22250 Axios (Vardar) R., reservoirs. 19 Anthemountas North Aegean 428 Anthemountas R. 20 Gallikos North Aegean 1022 Gallikos (or Echedoros) R. 21 Loudias North Aegean 1409 Loudias R. Vegoritis L., Cheimaditis L., Petron L., Zazari L., 22 Vegoritis North Aegean 752 tributary streams. Kastoria (or Orestias) L. (within the broader basin of the 23 Kastoria North Aegean 264 Aliakmon R.), tributary streams. Alkiakmon R., Almopeos R., Tripotamos R., reservoirs, 24 Aliakmon North Aegean 8677 Delta wetlands. 25 Mavroneri North Aegean 815 Mavroneri (or Itamos) R. 26 Pinios The North Aegean 9500 Thessalian Pinios R. Mikri and Megali Prespa L. (within the broader basin of 27 Prespa South Adriatic 1383 the Drin R.), Ag. Germanos R. 28 Aoos South Adriatic 6710 Aoos (Vjose) R. and reservoir near Metsovo. 29 Kalamas Ionian 1831 Kalamas (or Thyamis) R. 30 Zaravina Ionian 13 Zaravina L. 31 Pamvotis Ionian 330 Pamvotis (or Ioannina) L. 32 Paramythia Ionian 138 Small Lakes of Paramythia (basin of Margaritiou). 33 Kalodiki Ionian 69 Kalodiki L. (continued)

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http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 09/03/2019 12:43:28 | Table 1 (continued)

Estimated Biogeo-graphical No Name Given Area Included water bodies Area (km2) 34 Acheron Ionian 752Acheron R and coastal wetlands. 35 Ziros Ionian 10 Ziros L. 36 Louros Ionian 983 Louros R. 37 Arachthos Ionian 2009 Arachthos R., reservoirs and Delta. 38 Vouvos Ionian 205 Vouvos R. (Kombotiou R. basin). 39 Vlychos Ionian 45 Vlychos spring and Myrtari Lagoon. 40 Voulkaria Ionian 74 Voulkaria L. 41 Astakos Ionian 80 Astakos R. Acheloos R., Agios Dimitrios (Lesini) R., lakes, 42 Acheloos Ionian 6329 reservoirs, Delta and lagoons. 43 Evinos Ionian 1112Evinos R. and reservoir. 44 Mornos Ionian 998 Mornos R., reservoir, Delta wetlands and Gouvos Spring. Insular 45 Kerkyra Ionian Kerkyra (Corfu) Island water features. drainages Insular 46 Lefkas Ionian Lefkas Island water features. drainages 47 Assopos Pel Ionian 286 Assopos (Peloponnese) R. 48 Dervenios Ionian 65 Dervenios R. 49 Krios Ionian 130 Krios R. 50 Krathis Ionian 155 Krathis R. 51 Vouraikos Ionian 273 Vouraikos R. 52 Keronitis Ionian 98 Keronitis R. 53 Selinous Ionian 373 Selinous R. 54 Meganitis Ionian 107 Meganitis R. 55 Phoenix Ionian 97 Phoenix R. 56 Volinaios Ionian 55 Volinaios R. 57 Glafkos Ionian 142Glafkos R. 58 Piros Ionian 577 Piros R. 59 Tsivlos Ionian 10 Tsivlos L. (within the Krathis R. basin). Prokopos Lagoon, Lamia Swamp and adjacent springs and 60 Prokopos Ionian 280 streams. 61 Kotychi Ionian 266 Kotychi Lagoon and Vergas R. 62 Pinios Pel Ionian 868 Peloponnesian Pinios R. and reservoir. 63 Alfios Ionian 3658 Alfios R. and reservoir in the Ladon tributary. 64 Neda Ionian 287 Neda R. 65 Yiannousagas Ionian 48 Yiannousagas R. and adjacent Yalova Lagoon. 66 Peristeras Ionian 184 Peristeras (or Kalo Nero or Miras) R. 67 SW Messinia Ionian N/A Streams of SW Messinia, south of Kyparissia. 68 Pamissos Ionian 728 Pamissos R. and Aris tributary. 69 Kandila Ionian 216 Kandila spring and former wetlands. 70 Feneos Ionian 233 Doxa reservoir in Feneos Plateau and streams. 71 Stymphalia Ionian 216 Stymphalia L. 72 Taka Ionian 102Taka L. 73 Evrotas Ionian 1738 Evrotas R. (continued)

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Estimated Biogeo-graphical No Name Given Area Included water bodies Area (km2) 74 Vassilopotamos Ionian 14 Vassilopotamos canals within the Evrotas Delta Area. 75 Smynous Ionian 192Smynous (or Arniotiko) R. 76 Ardeli Ionian 78 Ardeli (or Ardelolaggado) R. 77 Lerni East Peloponnese 20 Lerni Spring. 78 Kato Almyri East Peloponnese N/A Kato Almyri Spring. 79 Erassinos Arg. East Peloponnese 22 Erassinos R. (in Argolis) 80 Vouliagmeni Attiko-Beotia N/A Vouliagmeni Karstic Lake, Attiki. 81 Erassinos Vra. Attiko-Beotia 25 Erassinos R. (in Vravron, Attiki). 82 Rafina Attiko-Beotia 90 Rafina (Megalo Rema) R. 83 Kato Souli Attiko-Beotia 40 Kato Souli (or Schinias Marathon) Wetland. 84 Marathon Attiko-Beotia 114 Marathon reservoir, Charadros stream and other tributaties. 85 Kifissos Att Attiko-Beotia 420 Attikos Kifissos R. (in Attiki). 86 Assopos Beo Attiko-Beotia 724 Beotian Assopos R. 87 Kifissos Beo Attiko-Beotia 1958 Beotian Kifissos R. (in Beotia). 88 Yliki Attiko-Beotia 494 Yliki L. and Paralimni L. 89 Thermopyles Attiko-Beotia 71 Thermopyles Springs. Sperchios R., Gorgopotamos and other tributaries, Delta 90 Sperchios Attiko-Beotia 1828 wetlands. 91 Cholorema Attiko-Beotia 192Cholorema R. (Pagasitikos Gulf). 92 Kireas Attiko-Beotia 441 Kireas-Nileas R. (Kirinthos), Euboea. 93 Manikiotiko Attiko-Beotia 158 Manikiotiko (or Monodriotiko) R., Euboea. 94 Dystos Attiko-Beotia 57 Dystos L., Euboea. 95 Rigia Attiko-Beotia 29 Rigia & Lala river, Karystos plain, Euboea. Insular 96 Samothraki Aegean islands Samothraki Island water features. drainages Insular 97 Lesvos Aegean islands Lesvos Island water features. drainages Insular 98 Samos Aegean islands Samos Island water features. drainages Insular 99 Rhodos Aegean islands Rhodos Island water features. drainages 100 Almyros Aegean islands 20 Heraklion Almyros spring and stream, Crete. 101 Koutsoulidis Aegean islands 578 Koutsoulidis R. (Yeropotamos Basin); including Zaros reservoir, Crete. 102 Kourtaliotis Aegean islands 109 Kourtaliotis R., Crete. 103 Kourna Aegean islands 20 Kourna L and adjacent water features, Crete. 104 Agia Aegean islands 166 Agia reservoir within the Platanias basin, Crete. 105 Tavronitis Aegean islands 131 Tavronitis R., Crete.

B N RESCU (2004): 1) Northern ed here by the Aoos R. and the Lake Pres- Aegean (includes both the Thrace and pa), 3) Ionian (includes the Ionian islands Macedonia-Thessaly zoogeographic and nearly all of the Peloponnese except regions), 2) Southern Adriatic (represent- the eastern part 4) East Peloponnese

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http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 09/03/2019 12:43:28 | (rain-shadow coastal area only), 5) Attiko- included as ‘isolated’ water features, Beotia (includes Euboea and Fthiotis in despite present or past drainage con- eastern central Greece), and 6) Aegean nections with wider river basins (e.g. islands (includes Crete). Lakes Kastoria, Doirani and Pamvo- The hydrographic basin areas cited are tis). In a few instances even very small defined by the traditional watershed lakes, semi-isolated spring-fed wet- boundaries of the entire catchment, taking lands, and other karstic surface water- into account hydrographic idiosyncrasies bodies are delineated in isolation, and historical drainage connections. In this either because they represent unique context, lakes with a present or past con- geological features or they have other nection to large rivers were usually incor- surface water attributes which are porated into the wider river basin area. worthy of particular conservation This also includes small rivers lying in the interest (e.g. Lakes Tsivlos, Vouliag- vicinity of the deltaic depositional zone of meni and Zaravina, Kato Almiri larger rivers. For instance, the Acheloos R. Spring). basin area is defined here to include the b. Some assemblages of separate small drainages of the natural lakes Trichonis, stream basins on the mainland and on Lyssimachia, Ozeros and Amvrakia; the the islands are incorporated into single artificial reservoirs Kremasta, Kastraki, artificially defined ‘basin areas’. This is Stratos and Tavropos; the entire deltaic done in cases where complete infor- wetland area, including the small Aghios mation on species composition for Dimitrios (or Yeroporos) R. with which each separate stream basin is lacking, the Acheloos R. had a direct connection but incidental records among the before the draining of the deltaic Lake group of proximate basins is consid- Meliti. The hydrographic basin areas of ered adequate to warrant the wider hydrologically isolated lakes are consid- areas’ inclusion (i.e. SW Messinia ered to contain the drainages of associated includes several small streams south of smaller lakes, as well as the drainages of Kyparissia-Messini, Peloponnese). In rivers discharging into them. For example, some other cases, surface hydrology is the hydrographic basin area of Lake complicated and semi-connected wet- Vegoritis contains the watershed of Lakes land conditions or artificial canals blur Vegoritis, Cheimaditis, Zazaris and Petron watershed boundaries; hence nearby and all streams discharging into these streams and wetlands are included lakes. Expert judgment based on criteria of (e.g. coastal wetland and isolated geographical distinctiveness and surface canal features such as the Laspias- hydrological connectedness was used to Avdira wetlands near Xanthi, Thrace). define the areal extent of the hydrograph- In the case of small islands, the entire ic basin area unit considered in this inven- network of an island’s inland water tory. In certain cases (14 basin areas), sev- features is presented as an artificially eral exceptions and violations to the use of defined ‘hydrographic basin’ area. traditionally defined hydrographic basin This treatment is of course provisional areas were made: and was followed simply because in a. Certain older lakes or lake groups most insular systems ichthyological within major river catchments are research is still inadequate. Only for

100 Medit. Mar. Sci., 8/1, 2007, 91-166

http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 09/03/2019 12:43:28 | two of the largest Aegean islands and all relevant publications up to mid (Crete and Euboea) are specific river 2007. KOTTELAT & FREYHOF (2007a) basin area compilations provided. provide a major revision of taxonomic units in Europe and include information Compilation of fish species occurrence for on the distribution of the taxa occurring in each area Greece. Some of the taxa described are not given separate species status while dis- We compiled a species-hydrographic tributional information for some species is basin area dataset hosting the freshwater vague or incomplete. For the sake of con- fish species occurring in Greece and, in the sistency and practical policy-relevant case of transboundary rivers, in neighbour- applications, we adopt the taxonomic ing southern Balkan countries (Albania, changes given by KOTTELAT & Bulgaria, FYR Macedonia, Turkey-in- FREYHOF (2007a) and we critically com- Europe). This compilation is based on an ment on particular difficulties in the taxon- extensive bibliographic study that began in omy and distribution of certain taxa in the 2003 (ECONOMOU et al., 2004a, 2006). checklist (Appendix I). In some cases we Unpublished survey data from HCMR carefully document divergence in certain field surveys were also utilized when the taxa distributions from that presented by available material was collected through the above authors. the participation of the authors in field The following species inclusion and sampling projects. The authors have been nomenclature premises were made: involved in wide-ranging site-based sam- a. All species, native or introduced, that pling surveys particularly in western, spend their entire lives or a significant northwestern and southern Greece portion of their life-cycle in freshwa- (including 73 of the presented hydrograph- ters were considered in the basin area ic basins areas); much of this work remains distributional compilation, i.e. primary unpublished in refereed journals (ECO- freshwater species (intolerant to salt- NOMOU et al., 1999, 2004a, 2007; ZO- water) and secondary freshwater GARIS et al., 2004, 2006). species (species that now live exclu- Attention was paid to documenting sively in freshwater but were once able and ascertaining the quality of each partic- to tolerate saltwater). Diadromous ular record for each basin area. The occur- species and euryhaline species for rence of a species in a basin area had to be which adequate distributional data provided by at least one reliable source exist are included, but so-called ‘spo- (publication, technical report) or to have radic species’ or marine stragglers that been confirmed by the authors during field seem indifferent to salinity or have a surveys. Some information was received definite marine life-history, are not from competent ichthyologists through included. Information is largely lack- personal communications. On some occa- ing in order to confirm the regular sions substantiated uncertainty of the qual- presence of certain marine species in ity of an accepted record is provided with a freshwaters, although proof of several question mark notation beside the record. marine species’ residence in brackish Fish systematic taxonomy was based transitional waters exists (e.g. KOU- on KOTTELAT & FREYHOF (2007a), TRAKIS et al., 2000).

Medit. Mar. Sci., 8/1, 2007, 91-166 101

http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 09/03/2019 12:43:28 | b. Certain euryhaline species known to confer) between the genus name and be regularly present in freshwaters but the species name to show that the for which consistent records are miss- species in question is similar to a ing are excluded from the distribution- named species but there is uncertainty al compilation. However, these species about its taxonomic status, or that this are placed within the summary check- species may represent a distinct list of freshwater fish in the appendix unnamed species. Our overall objec- and should be considered as important tive was to produce an operational and elements of the freshwater fauna. complete distributional compilation of c. Valid species names are given but in the freshwater fish of Greece rather some instances tentative operational than a taxonomically more accurate taxa names are given for species that but incomplete distributional account. present identification problems, strict- e. Notations are given where uncertainty ly following KOTTELAT & FREY- exists about the native or introduced HOFF (2007a). In addition, in five iso- status of a population and where the lated cases, we provide only our own population is presumed extirpated or operational name for unidentified possibly extirpated. species known only to genus level (i.e. two Eudontomyzon spp., Knipowitschia Annotated freshwater species checklist (in sp. Squalius sp1, Squalius sp2). This is Appendix) not a diversion from any accepted taxonomy, but it is required to show the A critical assessment of the ichthy- presence of an unidentified species in ological data assists in the production of an a particular basin area. updated checklist of species known to d. Species of undefined taxonomic status inhabit freshwaters in Greece. As previ- are included with a notation to indi- ously mentioned, certain migratory and cate taxonomic uncertainty. For exam- transient marine species regularly present ple, KOTTELAT & FREYHOF in freshwaters are included but most (2007a) provide the controversial marine species are excluded, since evi- taxon Salmo sp. Louros and we dence is lacking on how frequent these include this poorly described taxon in species are in the freshwater parts of the our compilation, but with a notation to basin areas. This checklist is still a prelimi- show ‘doubtful taxonomic status’. Fur- nary contribution for use in practical con- thermore there are several instances servation-relevant applications. Sweeping where KOTTELAT & FREYHOFF taxonomic name changes have drastically (2007a) refer to species occupying par- altered previously given names and abol- ticular hydrographic basins but give no ished the use of sub-species names, so a information on presumably the same practical updated list linking the previous or closely related fish that exist in and the current taxonomy is needed to nearby basins. In such cases of uncer- clarify confusion. In this respect, notation tainty we have used available distribu- on previous species names and new additional data and expert judgment to tions to Greece’s species list are given. For assign these fish to a specific taxon. conservation purposes the level of We used the acronym ‘cf’ (Latin for ‘endemicity’ of each species according to

102 Medit. Mar. Sci., 8/1, 2007, 91-166

http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 09/03/2019 12:43:28 | the scale of the Greek territory is summa- included in each area and the estimated rized. Endnotes present special taxonomic total surface area. The basin areas are problems and distributional uncertainties. arranged numerically according to their It must be made clear that in some cases position on the orientation map of Greece the available data is often of poor quality which is given in Figure 1. Most of the and uncertainties exist; consequently, basin areas covered in this survey lie on the unreliable records and poorly validated western, more humid side of the country taxonomic problems are inevitable. The (50 areas). Northern Greece covers a very checklist provides notes concerning our large region with the predominance of professional opinion on these difficulties. large or very large hydrographic basins (26 areas). Relatively few basins from the drier Supplementary list of species recorded in rain-shadow parts of the country (Central- transitional waters (in Appendix) Eastern Greece, Eastern Peloponnese, the Aegean islands) are accommodated in the A separate supplementary list of dataset, reflecting both the scarcity of larg- species recorded in transitional waterbod- er perennial waterbodies and poor sam- ies is also attempted (this includes river pling effort (29 areas). mouths, coastal lagoons and the brackish reaches of lowland rivers). This is certainly Composition and characteristics of the a very rough provisional contribution ichthyofauna of Greece needing further research. References are given for each species that is included in All species known from freshwaters, the list. This list does not include species native or introduced, are recorded in the cited in the aforementioned checklist of checklist of the freshwater fish fauna of freshwater species; therefore, many fresh- Greece (Appendix I). The supplementary water species that have been recorded in list of other euryhaline and marine species transitional waterbodies are not included. that have been recorded in the published This supplementary list attempts to pro- literature from transitional brackish waters vide a more complete picture of the hydro- is provided in Appendix II. graphic basin-based species assemblage, since transitional waters are definitely a The distributional compilation of part of the river basin area unit. Some of species occurrence data in 105 hydrograph- these species are potential candidates for ic basin areas is presented in Tables 2-6. inclusion in the freshwater checklist when The compilation includes all species con- further evidence is gathered. tained in the checklist, plus seven species that have been reported from sections of Results transboundary rivers outside Greek territory; however, it excludes 9 euryhaline Hydrographic basin areas considered in species contained in the checklist for which the survey the distributional accounts in freshwaters are incomplete. Among these, mugilids Table 1 lists the hydrographic basin (five species) have a widespread occur- areas considered in this study along with rence in estuaries and lowland sections of their main synonyms, the waterbodies rivers but their presence is not regularly

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http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 09/03/2019 12:43:28 | Fig.1: Map of Greece showing the location of the hydrographic basin areas considered in the ichthyofaunal distributional compilation. reported or given at species level. The same 2001; ECONOMOU et al., 2001b). Two holds for Atherina boyeri, which is com- more species excluded from the compila- monly found in coastal lagoons, estuaries tion are Dicentrarchus labrax and Zosteriss- and the lower reaches of rivers. Notably, esor ophiocephalus, which are not a fre- there are records of three landlocked quent target of faunistic investigations, and sandsmelt populations in Greece, in Lakes their occurrence in transitional and adjoin- Voulkaria, Kourna and Trichonis respec- ing freshwater waterbodies often goes tively (LEONARDOS, 2001; TIGILIS, unreported.

104 Medit. Mar. Sci., 8/1, 2007, 91-166

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Pinios The Pinios c

1 1

Mavroneri

Aliakmon

(continued)

Kastoria

Vegoritis

Loudias

Gallikos

Anthemountas Axios

11111 11111111 1111

Doirani c 1 11 1 1 1 1 1 1 Volvi

1 1

Rihios

Asprolakas

Mavrolakas Ladopotamos

2 Strymon a c

b,c

1 2 (1) Nevrokopi

Table 2

Marmaras

Nestos c a

1111 1111 1 1 (2) 1 1

Laspias

Kossinthos

Vistonis c

1 1

Kompsatos Filiouri

1111

Avas Evros a,b,c c a b,c 111 1 11111111 1?1111 1 1? 1 1 1 1? 1 1 1? 1 1 1 1 1 1 1212 1 1 11111 1 1 1 (1) 1 1 1 1 1 1 1 1 1 (1?) Fish faunistic lists for the hydrographic basins of Northern Greece. The position of the basins is shown in Figure 1. sp. Volvi 1 1 1 Alburnus vistonicus Alburnus volviticus Alosa fallax Alosa vistonica Aphanius fasciatus Alosa macedonica Aspius aspius Barbatula barbatula Anguilla anguilla Acipenser gueldenstaedtii Acipenser stellatus NumberAbramis brama Acipenser sturio Alburnoides bipunctatus Alburnus macedonicus 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 Alburnus alburnus Alburnus Alburnus thessalicus Barbus cyclolepis Barbus macedonicus Carassius auratus Barbus balcanicus Barbus sperchiensis Barbus strumicae

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1 1 1

Mavroneri b

Aliakmon a a

2 1 (continued) Kastoria

211 22

Vegoritis a b

Loudias

Gallikos

Anthemountas

Axios

Doirani

11 1 1 1 1 11 Volvi

222 22222 2

Rihios

Asprolakas

Mavrolakas

Ladopotamos

Strymon b 1 1

Nevrokopi

Marmaras Nestos a a Table 2 (continued)

(2) (2)

Laspias

Kossinthos Vistonis

221

Kompsatos b Filiouri

21? 1111 1 1 11?1

Avas Evros b b,c 111 1 1 111 11 1 1111111 1 11 1 111 1 11 1 11 11 1 1 11 12 12 2 2 2?1 1111111 22 1 222 11 1 111 21111111 2 2222 11 22(2?) 22 1 111 1 (1) 1 1 1 1 1 1 1 1 2 (2) sp. albula lavaretus peled cf. cf. cf. sp. (2) Chondrostoma vardarense Cobitis stephanidisi Cobitis puncticulata Cobitis punctilineata Cobitis strumicae Cobitis vardarensis Coregonus Eudontomyzon hellenicus Eudontomyzon Cyprinus carpio Gobio bulgaricus Huso huso Carassius gibelio Gasterosteus gymnourus Gobio feraeensis Ictalurus punctatus Ictiobus Knipowitschia caucasica NumberCarassius carassius 1 2Coregonus 3Coregonus 4Ctenopharyngodon idella 5 6 7 8Gambusia holbrooki 9 10 11 12 13Hypophthalmichthys molitrix 14 15 16 17 18 19 20 21 22 23 24 25 26 Esox lucius

106 Medit. Mar. Sci., 8/1, 2007, 91-166

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a,c 1

Mavroneri

Aliakmon a

2 1 (continued)

Kastoria Vegoritis

Loudias a

Gallikos

Anthemountas Axios a

111 111 111 111 1111

1 (1)

Doirani a

11 1 11? 1 11 11 1 1 1 1 11 11 1 21 1

Volvi c

Rihios

Asprolakas

Mavrolakas

Ladopotamos

Strymon a

1 1

Nevrokopi

Marmaras a

Table 2 (continued) Nestos a a

2 1 1

Laspias

Kossinthos Vistonis

11111 1 1 Kompsatos

11 1 1 11 11 1 1 1

Filiouri a

1 Avas

a Evros a 11 122222222 11111 1 2222222 1211 11 2222222 1 1111 1 11 1 1 11 1111 111111111 111111 11 1 1?1 1 11 111?11? 1 (1) (2?) (2?) (2) phoxinus macedonicus cf. cf. Salmo farioides Salmo pelagonicus Salmo Salaria fluviatilis Lepomis gibbosus Leucaspius delineatus Misgurnus fossilis Oncorhynchus kisutch NumberKnipowitschia thessala 1 2 3 4 5Phoxinus strymonicus 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 Oncorhynchus mykiss Perca fluviatilis Proterorhinus semillunaris Oxynoemacheilus bureschi Pseudorasbora parva Pungitius platygaster Pachychilon macedonicum Rhodeus amarus Rhodeus meridionalis Romanogobio elimeius Rutilus rutilus Sabanejewia balcanica Petroleuciscus borysthenicus Petromyzon marinus Phoxinus

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Pinios The Pinios

Mavroneri

Aliakmon Kastoria

1111 211

Vegoritis a a b

2 2

Loudias

Gallikos

Anthemountas Axios

111 1 Doirani ated, as follows: c

11 1 1 11 y.

Volvi a

2 1

Rihios a

Asprolakas

Mavrolakas

Ladopotamos

Strymon

Nevrokopi

Marmaras

Table 2 (continued) Nestos

Laspias

Kossinthos

Vistonis

Kompsatos

Filiouri

Avas Evros 111111 111111111 2 12 111 1 111 11111111 1 1111?1111 1? 1 (1?) 1 1 1 1 1 1 1 1 1 1 trutta cf. = Presumably native, reported but unconfirmed presence; = Reported but unconfirmed introduction. Tinca tinca Vimba melanops SUM 47 6 22 13 25 15 10 35 7 2 42 2 1 1 17 25 20 37 3 12 30 20 13 38 13 29 Zingel balcanicus NumberSalmo Salvelinus fontinalis 1Squalius vardarensis 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 Sander lucioperca Silurus glanis Scardinius erythrophthalmus Silurus aristotelis Squalius orpheus = Native, confirmed presence in river basin; = Introduced to the basin; : Doubts on native or introduced status; : Doubtful taxonomic status of population; : Extirpated or possibly extirpated population. 1 1? 2 2? The above symbols placed in parenthesis indicate occurrence only in sections of transboundary rivers outside the Greek territor a b c Notations are further given where taxonomic and native status uncertainty exists or where the population may be presumed extirp

108 Medit. Mar. Sci., 8/1, 2007, 91-166

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Lefkas Kerkyra

(continued)

Mornos

Evinos Acheloos c

Astakos

Voulkaria

Vlychos

Vouvos

Arachthos a

2 1

Louros a

Ziros Acheron

Table 3

Kalodiki

Paramythia

Pamvotis a

2 22 2

Zaravina

Kalamas a b,c

11 1 1?1 111 1 1 Aoos c c a

22 22 11? 1 1 1 1

(1) 1 1? 1? 1 (1) 1 1 1 1 1 1 1 1 (1) (1) Prespa a 1 1 111111?1 1 1111 1 11111 1 22221 2 2 2? Fish faunistic lists for the hydrographic basins of Western Greece. The position of the basins is shown in Figure 1. scoranza cf. NumberAcipenser baeri Acipenser gueldenstaedtii Acipenser naccarii Acipenser ruthenus Acipenser sturio Alburnoides bipunctatus Alburnoides prespensis 27 28 29Alosa fallax Anguilla anguilla 30 31 32 33Carassius auratus 34 35 36 37Cobitis hellenica 38Cobitis meridionalis 39 40 41 42 43 44 45 46 Alburnus belvica Alburnus Aphanius fasciatus Barbus rebeli Cobitis ohridana Barbus peloponnesius Barbus prespensis Carassius gibelio Clarias gariepinus Chondrostoma prespensis Chondrostoma vardarense Cobitis arachthosensis

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Lefkas Kerkyra c 1

(continued)

Mornos

Evinos

Acheloos a

2 2 1 1 2 2

Astakos

Voulkaria

Vlychos

Vouvos Arachthos

2?2 2?

Louros a

111 1?111

Ziros b

Acheron

1 11 Kalodiki

Table 3 (continued)

Paramythia

Pamvotis

Zaravina Kalamas

1 21 11 1 1111 1 111 1

Aoos a

1 1 Prespa 222 2 22 22 2 2 222?2 1 (2?)222 22(2)2 22 222 2 2 2 2 2 2 2 2 2 2? 2 2 2 sp. Louros 1 sp. 1 21 1? 1 1 1 1 1? lavaretus cf. sp. (1) skadarensis cf. Oreochromis niloticus Micropterus salmoides Oncorhynchus kisutch Oncorhynchus mykiss Oxynoemacheilus pindus Eudontomyzon Coregonus Esox lucius Knipowitschia Lampetra Lepomis gibbosus NumberCobitis trichonica Ctenopharyngodon idella 27 28Gambusia holbrooki 29 30Hypophthalmichthys molitrix 31Ictalurus punctatus 32 33Knipowitschia milleri 34 35 36 37 38 39 40 41 42 43 44 45 46 Cyprinus carpio Economidichthys pygmaeus Gasterosteus gymnourus Hypophthalmichhtys nobilis Economidichthys trichonis Gobio Knipowitschia goerneri Luciobarbus albanicus

110 Medit. Mar. Sci., 8/1, 2007, 91-166

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Lefkas a Kerkyra

(continued)

Mornos

Evinos

Acheloos a

1 1 1 2 2 2

Astakos

Voulkaria

Vlychos

1 11111 1

Vouvos Arachthos

1111 Louros a,c a,c a

1 1

1? Ziros

a,b

Acheron a Kalodiki

Table 3 (continued)

Paramythia

Pamvotis a

21 21? 11 2 2? Zaravina a

1? Kalamas

11 1111 11111 1 Aoos a a

1 111 1

(1) Prespa a 1 222 2 1 2 1 1 sp. (2?) sp. Louros Salaria economidisi Rutilus prespensis Salaria fluviatilis Salmo dentex Silurus aristotelis Rhodeus amarus Petromyzon marinus Poecilia Pseudorasbora parva Polyodon spathula Perca fluviatilis NumberPachychilon pictum Parabramis pekinensis Pelasgus prespensis Pelasgus thesproticus 27 28 29 30 31Rutilus ohridanus Rutilus panosi 32 33 34 35Salmo farioides 36 37 38Salvelinus fontinalis 39 40 41 42 43 44 45 46 Pelasgus epiroticus Pelasgus stymphalicus Scardinius acarnanicus Salmo peristericus Salmo Salmo letnica

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Lefkas c

Kerkyra c a

Mornos a

Evinos a

Acheloos a

1 1

Astakos

Voulkaria a

Vlychos

Vouvos a

Arachthos a

Louros a

11 1 11 1 1 Ziros a,b b

Acheron a

1 Kalodiki

Table 3 (continued)

Paramythia

Pamvotis

Zaravina

Kalamas a

111111111 11 11 1

Aoos Prespa 2222 22 2 221 2 . peloponnensis cf sp. Aoossp. Evinos 1 Tropidophoxinellus hellenicus Tinca tinca Valencia letourneuxi NumberSilurus glanis Squalius pamvoticus Squalius prespensis Squalius Telestes pleurobipunctatus 27 28 29SUM 30 31 32 33 34 35 25 36 24 37 20 38 9 39 23 40 1 41 42 2 43 15 44 5 45 22 46 20 3 8 8 4 39 13 10 10 5 Squalius Squalius Notes as in Table 2

112 Medit. Mar. Sci., 8/1, 2007, 91-166

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Erassinos Arg Erassinos a

1 Kato Almyri Kato

1 Lerni a

(continued)

Adreli a

Smynous a

Vassilopotamos

Evrotas

Taka

Stymphalia a

Feneos

Kandila Pamissos

12 11 SW Messinia SW a

Peristeras a

Yiannousagas a

Neda c Alfios c

22? 22 2 1111 1 2 2 2? 2? 1? 1?

Pinios Pel Pinios

22 2 12?

Kotychi

1111 Prokopos

11 1

Table 4 Tsivlos

b 2222?222 22222 22 22221111 2 2 1111111 1

Piros

Glafkos

Volinaios

Phoenix

Meganitis

Selinous

Keronitis

Vouraikos Krathis

11 1 11 Krios

1? 1 1 1 1 1? 1 1 1 1 1 1 1 1 1 1 1 1?11 11 1 11111 1111 11111?11 1? 1 1 1? 1 1 1 1 1 1 1 1 1 Dervenios

22 2 2 2 2 Assopos Pel Assopos c 1 Fish faunistic lists for the hydrographic basins of the Peloponnese. The position of the basins is shown in Figure 1. sp. 1 1 1 peloponnensis trutta cf. cf. Aphanius fasciatus Barbus peloponnesius Carassius gibelio Ctenopharyngodon idella Cyprinus carpio Gambusia holbrooki Gasterosteus gymnourus Hypophthalmichthys molitrix Knipowitschia Leponis gibbosus Luciobarbus albanicus Oncorhynchus kisutch Oncorhynchus mykiss Pelasgus laconicus NumberAcipenser sturio Anguilla anguilla Aphanius almiriensis 47 48 49 50 51 52 53 54 55 56 57 58 59 60 61 62 63 64 65 66 67 68 69Pelasgus stymphalicus 70Perca fluviatilis 71 72 73 74 75 76 77 78 79 Squalius Petromyzon marinus Salaria fluviatilis Salmo farioides Salmo Silurus glanis

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Erassinos Arg Erassinos

Kato Almyri Kato

Lerni

Adreli

Smynous

Vassilopotamos Evrotas

Taka

Stymphalia c

Feneos

Kandila Pamissos

SW Messinia SW

Peristeras

Yiannousagas Neda

11111 11

Alfios c Pinios Pel Pinios

1 11 Kotychi

111

Prokopos

Tsivlos Piros

Table 4 (continued)

Glafkos

Volinaios

Phoenix

Meganitis

Selinous

Keronitis

Vouraikos

Krathis

Krios

Dervenios Assopos Pel Assopos hellenicus Squalius moreoticus NumberSqualius keadicus Telestes pleurobipunctatus 47 48 49 50 51 52 53 54 55SUM 56 57 58 59 113441 421114557132255651252437622523 60 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 76 77 78 79 Tropidophoxinellus Tinca tinca Tropidophoxinellus spartiaticus Valencia letourneuxi Notes as in Table 2

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http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 09/03/2019 12:43:28 | Table 5 Fish faunistic lists for the hydrographic basins of Central-Eastern Greece. The position of the basins is shown in Figure 1. Yliki Rigia Kireas Rafina Dystos Sperchios Marathon Cholorema Kato Souli Kifissos Att Assopos Beo Kifissos Beo Thermopyles Manikiotiko Vouliagmeni Erassinos Vra

Number 80 81 82 83 84 85 86 87 88 89 90 91 92 93 94 95 Alburnoides bipunctatus 1a Anguilla anguilla 11 1 1 11 1 1 1 1 1 1 Aphanius fasciatus 1 Barbus euboicus 1 Barbus sperchiensis 111 Carassius gibelio 22 Ctenopharyngodon idella 2?2 Cyprinus carpio 222 2 Gambusia holbrooki 2 22 2 2 2 22 Gasterosteus gymnourus 1 Hypophthalmichthys molitrix 2 Hypophthalmichthys nobilis 2 Knipowitschia caucasica 11 Luciobarbus graecus 11 1 Oncorhynchus kisutch 2 Oncorhynchus mykiss 22a Oreochromis niloticus 2 Pelasgus marathonicus 1c 11 11c 11 11 Poecilia cf. latipinna 2a Pungitius hellenicus 1 Rutilus sp. Sperchios 1 Rutilus ylikiensis 11 Salaria fluviatilis 1 Salmo salar 2? Scardinius erythrophthalmus 1a Scardinius graecus 11 Silurus glanis 1?b Squalius sp.1 1c Squalius sp. Evia 1a 1 Squalius vardarensis 1a 1a Telestes beoticus 1c 11 SUM 2 3 1 3 3 3 4 11 11 1 17 4 3 3 2 3

Notes as in Table 2

The distributional compilation indi- from hydrographic basins within the Greek cates the presence of 162 species (including territory. Five of the species present in the doubtful occurrences) in 105 basin areas. North Aegean region, all introduced, do Of these species, 155 have been recorded not occur in Greece and are known only

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http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 09/03/2019 12:43:28 | Table 6 Fish faunistic lists for the hydrographic basins of the Aegean Islands. The position of the basins is shown in Figure 1. Agia Samos Kouna Lesvos Rhodos Almyros Tavronitis Samothraki Kourtaliotis Koutsoulidis

Number 96 97 98 99 100 101 102 103 104 105 Alosa fallax 1 Anguilla anguilla 11111 111? Aphanius fasciatus 11 Barbus pergamonensis 1 Carassius auratus 2?a Carassius gibelio 2 Ctenopharyngodon idella 22 Cyprinus carpio 2?2 Gambusia holbrooki 2222?2 2 22 Hypophthalmichthys molitrix 22 Knipowitschia caucasica 111 Ladigesocypris ghigii 1 Oncorhynchus kisutch 2 Oncorhynchus mykiss 2222 Oxynoemacheilus theophilii 1 Petroleuciscus smyrnaeus 1 Salaria fluviatilis 11 1 Salmo salar 2 Squalius cf. cii 1 Squalius sp.21 SUM 2 10 5 7 4 4 2 6 2 3

Notes as in Table 2 from Bulgarian sections of the rivers Evros, areas of their occurrence; these are Strymon and Nestos (Coregonus albula, C. referred to as aliens to the country (anoth- peled, Ictalurus punctatus, Misgurnus fossilis, er two species were reported but their pres- Ictiobus sp.). Some characteristics of the ence requires confirmation). Ten of these fish assemblages in the designated biogeo- aliens have been established through natu- graphic regions are summarised in Table ral reproduction; the presence of the 7a. The North Aegean region has the high- remaining 14 species in the wild depends est (85 species) and the East Peloponnese on stocking and aquaculture escapes. region has the lowest (6 species) species Finally, 19 species have native populations richness. The total number of native in Greece but are introduced in one or species with confirmed occurrence in the more hydrographic basins outside their examined basins is 130. 25 species were native range. These species are referred to invariably assigned as introduced in all as translocated, though there are instances

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http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 09/03/2019 12:43:28 | Table 7a Summary data of the ichthyofauna of Greece. Fish occurrences in the hydrographic basin areas included in the distributional compilation.

Designated regions All South East Attiko- Aegean Attributes North Aegean1 Ionian3 regions Adriatic2 Peloponnese4 Beotia5 Islands Number of basins areas 26 2 48 3 16 10 105 Number of fish taxa (species) 85 45 65 6 31 20 162 Species within Greek territory 80 37 61 6 29 19 155 (confirmed occurrences) Species recorded only in neighbouring countries 5 6 - - - - 7 (confirmed occurrences) Native species 65 29 37 5 19 11 130 (confirmed occurrences)* Introduced species 20 14 24 1 10 8 25 (confirmed occurrences)** Regional endemics (confined 29 18 28 1 9 1 86 to the biogeographic region) Proportion of regional endemics to the native fish 44.6 62.1 75.7 20.0 47.4 9.1 66.1 fauna of the region (in %) Average basin area species 18.0 23.0 6.9 3.0 4.4 4.2 9.1 richness (confirmed occurrences) Average basin area native species 14.0 15.0 4.8 2.3 3.1 2.3 6.9 richness (confirmed occurrences) * Species recorded as native in at least one basin area of the biogeographic region. ** Species always recorded as introduced in the region.

1: Northern Greece, from Evros R. to Pinios Thessaly R.; Table 2. 2: Prespa L. and Aoos R.; Table 3. 3: Western Greece, from Kalamas R. to Evrotas R.; Tables 3 (mainland western Greece and Epirus) and Table 4 (Peloponnese). 4: The eastern coasts of Peloponnese (Kato Almyri Spring, Erassinos R., Lerni Spring); Table 4. 5: Central Eastern Greece, from Cholorema R. to Vouliagmeni L. including Euboea Island; Table 5.

Table 7b: Summary data of the ichthyofauna of Greece. Fish contained in the checklist of freshwater fish species (Appendix I) and the supplementary list of transitional water species (Appendix II). Attributes Number Total number of fish species confirmed in freshwaters (Appendix I) 161 Typical freshwater species (do not readily enter seawater conditions) (Appendix I) 138 Brackish water or marine species that spend part of their lives in freshwater conditions (Appendix I) 23 Number of Greek endemics (species confined exclusively to Greece) (Appendix I) 47 Number of Balkan endemics (species with a distribution restricted south of the Danube R.) (Appendix I) 28 Number of "near endemic" species (species found along the frontiers of Greece) (Appendix I) 14 Total number of fish species recorded in transitional waters but not confirmed 55 as residents of freshwaters (Appendix II) Total number of fish species recorded in fresh and transitional waters (Appendices I+II) 216

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http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 09/03/2019 12:43:28 | where the introduced specimens were presence in freshwaters. The supplemen- imported from abroad, rather than being tary list (Appendix II) contains 55 species translocated from another Greek basin. with a recorded presence in transitional Taking into account the distributional waters, arranged in 22 families. ranges of the native species, 86 species are Table 8 shows the ten native and the recorded as regional endemics (their dis- ten introduced species with the highest fre- tribution is confined to only one biogeo- quency of occurrence in the examined graphical region). Some endemics are basins. Of the native species, Anguilla known from single or very few basin areas. anguilla is the most widespread, reported This category of range-restricted endemics from 74 basins, followed by Salaria fluvi- includes species of high conservation pri- atilis, which is known from 32 basins. First ority, such as Aphanius almiriensis, Alosa in the list of introduced species are Gam- vistonica, Barbus euboicus, Cobitis busia holbrooki, with confirmed occurrence stephanidisi, Eudontomyzon hellenicus, in 52 basins and Oncorhynchus mykiss, with Pungitius hellenicus and Squalius keadicus. occurrence in 27 basin areas. The latter A number of species have a distribution species has not yet been reported as repro- confined to the frontiers of Greece with ducing in Greece and its occurrence neighbouring countries (Lake Doiraini: depends on stocking programmes and one species; Lake Prespa: nine species). aquaculture escapes. Two alien species, All regional endemics of the East Pelo- Pseudorasbora parva and Lepomis gibbosus, ponnese, Attiko-Beotia and Aegean are highly invasive and their distributional Islands are entirely within Greek territory. range is expanding. The Ionian region has the highest A comparison of fish assemblage com- endemicity level (75.7%) and the Aegean position among the six defined biogeo- Islands the lowest (9.1%). graphical regions reveals that only 15 Table 7b summarises the fish data native species, of the 130 native species appearing in Appendices I and II. The total recorded, have joint presence in two or number of fish species recorded in the more regions (see Table 9). Seven of these fresh and transitional waters of Greece is species (Aphanius fasciatus, Knipowitschia 216. The checklist of freshwater fish species caucasica, Salaria fluviatilis, Anguilla (Appendix I) contains 161 species that live anguilla, Alosa falax, Gasterosteus gymnu- in freshwaters, arranged in 28 families. Of rus and Petromyzon marinus) are second- these, the family Cyprinidae strongly dom- ary freshwater or peripheral fish with pre- inates with 80 species that comprise 49% of sumed ability to utilise the marine route the total number of species. Another five for their dispersal. One species, Cyprinus families (Acipenseridae, Cobitidae, Sal- carpio, with paired presence in the Adriat- monidae, Mugilidae, and Gobidae) are ic region (Lake Prespa) and the North represented by 5 or more species. Twelve East Aegean regions has been ommitted families are represented by single species. because its native status in Lake Prespa is Judging from spatial occurrence informa- not certain. The high degree of faunistic tion, 139 taxa are provisionally classified as dissimilarity among regions can only partly typically freshwater species (recorded pri- be attributed to the fine-level taxonomy marily from freshwaters) and 23 are classi- adopted in this study. The most probable fied as euryhaline species with a confirmed reason of the dissimilarity is the presence

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http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 09/03/2019 12:43:28 | Table 8 The most widespread native and introduced fish species in Greek freshwaters (ranked from 1-10). Introduced species highlighted in grey are recorded as translocated and may occur as native in some hydrographic basins.

Top 10 Native species Top 10 Introduced species Species Catchments % Species Catchments % Anguilla anguilla 74 70.5 Gambusia holbrooki 5249.5 Salaria fluviatilis 3230.5 Oncorhynchus mykiss 2725.7 Squalius cf. peloponnensis 2120.0Carassius gibelio 20 19.1 Barbus peloponnesius 2120.0Cyprinus carpio 18 17.1 Aphanius fasciatus 2120.0Ctenopharyngodon idella 11 11.4 Pelasgus stymphalicus 20 19.1 Hypophthalmichthys molitrix 10 10.5 Cyprinus carpio 19 18.1 Lepomis gibbosus 10 10.5 Knipowitschia caucasica 18 17.1 Pseudorasbora parva 10 10.5 Rutilus rutilus 16 15.2 Tinca tinca 7 6.7 Gobio bulgaricus 15 14.3 Oncorhynchus kisutch 5 4.8 of geographic barriers preventing faunal species. Basin area species richness is exchanges among regions. determined by a multiplicity of factors representing local and regional scales. Local Fish species richness and endemism influences on species richness include factors determining habitat diversity and Hydrographic basin area species rich- environmental stability, such as basin sur- ness (number of fish species per area) is face area, discharge level and variability, generally low in Greece. The number of mean elevation and slope, presence of native freshwater fish species per area var- lakes or floodplains in the basin, etc. Here ied from 0 to 32 (Evros) and that of intro- we restrict ourselves to the examination of duced species varied from 0 to 18 (Pamvo- the relationship between basin native tis and Acheloos). Only 53 hydrographic species richness and basin surface area basins, mostly small, had no record of (see Fig. 3). Despite the high scatter of introduced species. The areas with the points, the data show a clear increase of highest total basin richness (number of richness with basin area. The small size of native and introduced species) are Evros most basins explains, at least partly, the low (41), Strymon (40), Acheloos (38) and Ali- average richness in Greek waterbodies. akmon (38). Figure 2 shows the distribu- On the regional scale, basin area tion of richness in the basin areas exam- species richness reflects the pool of ined, where richness was calculated sepa- species that occur in a biogeographic rately for native, introduced and all region, as determined by a combination species. More than half of the examined of historical factors and contemporary basin areas (59) host up to five species, environmental influences. Other condi- with only five basin areas hosting more tions being equal (e.g. when basins of sim- than 35 species. If only the native species ilar size are compared), basin species are considered, the number of basin areas richness is associated with regional hosting up to five species rises to 65. Only species richness (defined as the number ten basin areas host more than 20 native of fish species known to occur within a

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http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 09/03/2019 12:43:28 | biogeographic region). Inspection of the extinctions and the depauperation of the data shows that the basins in the North local fish communities. Aegean contain on average more species Figure 5 shows the spatial distribu- than basins of similar size in the other tion of fish endemicity. Within Greece regions. This is largely a reflection of the endemicity is very high (47 species found richer ichthyofauna in the North Aegean exclusively in Greece; or 35% of its native region in comparison to other regions fish fauna). In some basin areas (e.g. Ach- (see Fig. 4). The regional species richness eloos, Evrotas, Beotian Kifissos), the pro- is particularly low in the Aegean islands, portion of endemic fish exceeds 75% of which are comprised of very small insular the total number of native fish. The gen- basins. Low annual precipitation and the eral trend is towards an increase of the frequent occurrence of prolonged proportion of endemics westwards and droughts may have contributed to species southwards. This trend is opposite to that

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http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 09/03/2019 12:43:28 | Fig. 3: Relationship between hydrographic basin surface area (km2) and native species richness.

Fig. 4: Regional species richness (native, introduced and total fish species) for the six designated biogeographical regions. Figures above columns indicate number of basin areas. of species richness, which increases east- slightly beyond the Greek territory wards and northwards. Another 14 (specifically in the Butrint basin in Alba- species (10.3%) are considered ‘near- nia). Finally 28 species (20.6%) have a endemic’ since they inhabit isolated wider distribution in the Balkan Peninsu- waters on the borders of Greece (Prespa la south of the Danube, considered here and Doirani) or their range extends as Balkan endemics.

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http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 09/03/2019 12:43:28 | Fig. 5: Endemicity of the fish fauna relative to the territorial boundaries of Greece. Endemics confined to Greece include 47 species (ENWE, ENNO, ENCE, ENAEG). ‘Near-endemics’ include 14 species (NENNO, ENWE+). 28 species are restricted to the Balkans (ENBAL). See Appendix I - Legend for codes descriptions.

Discussion 1972; ECONOMIDIS et al., 1981; ECO- NOMIDIS & SINIS, 1982; KOKKI- Sources of bias in compiling the basin NAKIS et al., 1999; TIGILIS, 2000; area species lists and checklists – data KOUTRAKIS et al., 2000; DAOULAS et availability, knowledge gaps and unmet al., 2001; ECONOMOU et al., 2001b, needs 2004b; TACHOS, 2003; KOKKINAKIS, 2006; STOUMBOUDI et al., 2006; The freshwater fish of Greece have LEONARDOS et al., 2007) or describe been studied for more than 150 years. His- the distributional ranges of species and torical ichthyological information and species groups (e.g. ECONOMIDIS, 1989; early records of commercial catches (e.g. ECONOMOU, 2000; ECONOMIDIS et VALENCIENNES, 1844; HELDREICH, al., 2000b; BOBORI et al., 2001; 1878; APOSTOLIDIS, 1883, 1892; DAOULAS, 2003; KALOGIANNI et al., ATHANASSOPOULOS, 1917, 1923, 2006). An even smaller number of publica- 1925; KOLLER, 1927) provide a sound tions take a synthetic approach to fish dis- basis for ascertaining the native distribu- tribution treating all species and/or com- tion of many species. However, the distri- piling basin-specific species lists over wide bution of some species is still insufficiently geographical areas (STEPHANIDIS, known and their native or introduced sta- 1939, 1950, 1971; ECONOMIDIS, 1974; tus in some waterbodies is uncertain. DAGET & ECONOMIDIS, 1975; Despite the relatively large number of ECONOMOU et al., 1999, 2001a; publications dealing with freshwater fish ECONOMIDIS et al., 2001; BARBIERI et (ECONOMOU et al., 2004a), few only al., 2002). Two of the aforementioned provide complete fish faunistic lists in indi- works, namely the thesis dissertations of vidual drainages (e.g. KATTOULAS, STEPHANIDIS (1939) and ECONO-

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http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 09/03/2019 12:43:28 | MIDIS (1974), deserve special mention Another problem with our dataset – for comprehensive taxonomic work and that again addresses unmet data needs – is detailed accounts of species occurrences in the unresolved issue of each basin area’s a large number of freshwater bodies. A fish assemblage completeness. The data catalogue of the fish of Greece produced provided here sometimes refer to very by ECONOMIDIS (1973) forms a land- large basin areas, thus much local informa- mark in the ichthyological research of tion has been amalgamated, and important Greece for reporting all freshwater fish site-specific data is not presented (i.e. fish occurrences known at that time. A popu- assemblages along a particular river seg- larized check-list was later published by ment). We are almost certain that some the same author (ECONOMIDIS, 1991). species, especially introduced ones, are It is inevitable that some of our basin missing from even the larger river basin area’s fish compilations may have errors of area accounts in our dataset. Also, some omission. It is remarkable that with few smaller river basins, such as the Acheron, exceptions many smaller isolated aquatic Kalamas, Kireas, and Lerna, do not yet sites, especially along Greece’s Aegean have complete species lists. This is either coastline and its islands, have never been because sampling efforts have been few or properly surveyed for fish. For example, the sampling conditions are especially dif- our compilation provides data for only ficult due to deep-water, non-wadeable eight islands although wetland habitats reaches which have never been surveyed exist on a very large number of Greek with appropriate tools (i.e. boat-based islands. In a recent inventory of wetland electrofishing, multi-mesh gill-nets etc). sites (CATSADORAKIS & PARAGA- Lastly, there has been no recent sampling MIAN, 2007), fish presence was recorded in some basin areas. For example, the most in 72 of 352 small wetlands in the Aegean reliable account of the Mornos R. fish islands; unfortunately, however, these fish dates back to 1972 (KATTOULAS, 1972) were rarely identified to species level. A and is based on data obtained with old- stark example of the extent of the unex- fashioned sampling techniques that may plored areas is given by the extent of wet- have been ineffective for small-bodied land exploration on the large island of species likely to occur in the lower parts of Euboea (3.685 Km2), which hosts a unique the river. In addition, this investigation freshwater fish fauna that includes two took place before the huge wave of alien local endemics. CATSADORAKIS & species’ introductions that occurred from PARAGAMIAN (2007) provide descrip- the 1970’s through the 1990’s. Thus, the tions of 39 wetlands on Euboea and they apparent absence of introduced species in note that this number is far from a com- this system may reflect lack of recent fau- plete inventory; we compiled ichthyofau- nistic information. nal data of only four Euboean wetlands, Apart from these distributional ambi- two of which are not listed in the afore- guities, there are also problems with occur- mentioned inventory. The number of wet- rence records of some species appearing in lands on Euboea is certainly very large, old publications but are not verified by and sadly we have only the slightest knowl- recent surveys. For example, we decided to edge of freshwater fish distributions on note our doubt for the occurrence of this island. Alburnoides bipunctatus in the Acheron R.,

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http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 09/03/2019 12:43:28 | despite the fact that its presence in this Moreover, there are unresolved taxo- river has been mentioned by STE- nomic issues with respect to the identity of PHANIDIS (1939) on the basis of a single some populations. Due to the difficulties in specimen collection. This species has not resolving taxonomy rapidly, many past been found in the Acheron R. since then records of occurrence are difficult to sub- and, more importantly, it is absent from all stantiate. For example, the species Pseu- other river basins of the Ionian region, dophoxinus stymphalicus has recently been which substantiates our doubt. However, split to five species under the new genus the Acheron R. is far too deep to be sam- name Pelasgus (see KOTTELAT & pled efficiently with conventional sampling FREYHOFF, 2007b). All previous pub- techniques, so the presence of A. bipuncta- lished records referring to the nominate tus in this river cannot be excluded with species P. stymphalicus are difficult to be certainty. On similar grounds, we decided included in a geographical compilation to express doubt for the occurrence of because the distributional limits of the Barbus peloponnesius in the Louros R., newly described taxa are not yet complete- mentioned in older surveys, because this ly defined. Are the specimens from the east- species is persistently absent in the collec- ern coast of the Peloponnese P. stymphali- tions of extensive recent surveys. cus, P. laconicus or P. marathonicus? Also, In addition, the bibliography is rife what is the translocated Pelasgus species in with ambiguities concerning site-specific Lake Pamvotis? Challenging problems information that would be of use to practi- with the systematics and the distribution of cal management and conservation applica- many species remain to be resolved, such as tions (ECONOMOU et al., 2006). Difficul- confirming the taxonomic validity of some ties in ‘deciphering’ locality information unnamed and undescribed taxa within the from the bibliography are widespread and genera Squalius and Alburnus for which they may also be a major reason for poten- morphological characters alone do not tial errors. There is even difficulty with allow reliable identification. Molecular interpreting the basin-specific location of studies have greatly contributed to unravel- data for many species. For example, ing the genetic and the underlying phyloge- ECONOMIDIS (1991) mention the exis- netic relationships of some species. Ongo- tence of Pseudophoxinus (now Pelasgus) ing genetic work and other biological inves- stymphalicus on the island of Euboea with- tigations within the next few years should out giving specific locality information. focus on sorting relationships in taxa with Other survey compilations have also disjunct or fragmented distributions. encountered difficulty with unconfirmed or Finally, there is a lack of published erroneous identifications of fish site-based ichthyofaunal surveys on transi- (MAURAKIS et al., 2003). Unfortunately tional waters. This is a serious problem it is very difficult to verify distributional because a large number of species use information, and errors can easily creep estuarine areas, rivermouths, lagoons and into a dataset if the inventoried data are brackish waters in deltas as rearing areas not confirmed with recent field observa- or seasonally as transient habitats. Our tions. This shows that more field work is bibliographic search showed that many needed to confirm and monitor the pres- species have been recorded in transitional ence of species in many river basins or sites. waters but many surprising records of

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http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 09/03/2019 12:43:28 | marine species need verification (including differences of diagnostic importance could marine stragglers such as Epinephelus not be established. For instance, aeneus, Scorpaena scrofa and Gaidropsarus KOTTELAT & FREYHOFF (2007) uti- mediterraneaus). We must reiterate the lized evidence of genetic distinctiveness provisional and tentative nature of our provided by BOHLEN et al. (2006) to supplementary list of species recorded in establish Rhodeus meridionalis that other- brackish waters since 36 (56%) of the wise would not be easily recognized as a species that we present had been submit- distinct species on the basis of classical fea- ted by a single published bibliographic ref- tures such as morphology. Another cause erence. On the other hand, our list con- of this increase of species number has been tains several species which possibly enter the introduction of several alien species to freshwaters regularly. In fact, some species many waterbodies for aquaculture and in this list were located in the lists of fishery enhancement (e.g. CRIVELLI et ‘freshwater’ species by ECONOMIDIS al., 1997; ECONOMIDIS et al., 2000a; (1991) and BOBORI & ECONOMIDIS LEONARDOS et al., 2007). (2006). Pending accurate detailed infor- The present checklist of the freshwater mation on these and other species’ resi- fish of Greece (Appendix I) compiles the dence in freshwaters we retain them only fish data in a standardized way comparable within this provisional supplementary list. to ECONOMIDIS (1991), i.e. it includes introduced, diadromous and some euryha- The current ichthyological picture of line species. In total, the checklist contains Greece 161 species, of which 14 are of unclarified taxonomic status and are given provisional The last published annotated checklist or generic names. The great increase in of the freshwater fish of Greece contained species numbers (56 species, 53%) since 105 species, including introduced, diadro- 1991 has resulted mainly from taxonomic mous and euryhaline species with a regular re-evaluations of existing taxa, rather than presence in freshwaters, and also five from the genuine discovery of new species. species of doubtful occurrence (ECONO- In fact, most of the new species contained MIDIS, 1991). Since then the list of in the checklist were known as biological species has expanded considerably as sev- entities in 1991 but were recognized as eral species have been recently described subspecies or were lumped with their (e.g. KOTTELAT, 2004; KOTTELAT & closely related species. BARBIERI, 2004; ECONOMIDIS, 2005; The apparent tendency towards KOTTELAT & ECONOMIDIS, 2006; species splitting is in accordance with the BOGUTSKAYA & ILIADOU, 2006; currently prevailing trend towards adop- STOUMBOUDI et al., 2006; KOTTE- tion of the Phylogenetic Species Concept LAT, 2007; KOTTELAT et al., 2007). (PSC) over the older Biological Species Some more taxa were named and desig- Concept (BSC). KOTTELAT (1997) has nated as species by KOTTELAT & strongly advocated the use of the PSC FREYHOFF (2007a) mostly on the basis (considered as equivalent to the Evolu- of morphological data. Molecular data tionary Species Concept) for species have contributed to some taxonomic clari- recognition and employed this concept in a fication, especially when morphological first revision of the systematics and

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http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 09/03/2019 12:43:28 | nomenclature of the European freshwater PSC might have on the users of fish data. fish. This revision resulted in a great We shall begin with the definition of increase in the number of species in species and the criteria used to delineate Europe with well over a hundred more species boundaries under the two species species than previously recognised. concepts, namely the BSC and the PSC. Recently KOTTELAT & FREYHOFF We shall also discuss the strengths and (2007a) produced a handbook that further weaknesses of the two concepts and the revises the systematics of European fresh- potential implications of the change in water fish. This second revision has radi- species concept on the uses (and users) of cally changed the ichthyofaunal list of fish data. Next, we shall provide an Greece by introducing new names for overview of the distributional status of the species and genera and raising many pop- freshwater fish of Greece, particularly with ulations and subspecies to species rank. In respect to regional patterns of richness and fact, 29 species from Greek freshwaters endemism. Last, we shall examine some (18%) are ‘new’ since the ECONOMIDIS policy-relevant implications of the fish (1991) checklist was published. With the dataset presented in this paper and we demise of the subspecies unit, many sub- shall explore areas of research and man- specific taxa have been lost while others agement priority. have been awarded species status. Overall, only 41% (67 species) of the species names Species concepts - distinctiveness criteria given in the present list are identical to and utility relative to the needs of users those used by ECONOMIDIS (1991). Judging from trends in recent taxo- How to define ‘species’ is one of the nomic publications, the new species con- most fundamental and controversial issues cept is accepted by several taxonomists in biology (KULLANDER, 1999; BAR- working with Greek freshwater fish and is TON, 2001; MAYDEN, 2002; REYDON, likely to become the dominant concept 2005). More than 20 species concepts have guiding fish systematics in the future been put forward (MAYDEN, 2002) and (MILLER, 1998). The changes of species considerable debate still exists about their names and the emergence of new species theoretical basis, applicability, and conse- may create problems to users of the fish quences for studies of ecology and biodi- data who do not have a background in sys- versity (MAYR, 1996; HENDRY et al., tematics. We have attempted to resolve 2002; ISAAC et al., 2005; AGAPOW & some of the anticipated problems by quot- SLUYS, 2005). We confine ourselves to a ing in the checklist of freshwater species brief presentation of the two prevailing (in Appendix I) the previous species species concepts, the BSC and the PSC, names and indicating new additions. Given focusing on topics of relevance to biodiver- that both the checklist and the distribu- sity conservation and watershed manage- tional compilation are based on the taxon- ment from a fish-based perspective. More omy proposed by KOTTELAT & detailed presentations and arguments in FREYHOFF (2007a), we consider it favour or against their conceptual and important to clarify the meaning of species methodological basis can be found in the under the two species concepts and to aforementioned publications and also in explore what impact the acceptance of the KOTTELAT (1997), TURNER (1999),

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http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 09/03/2019 12:43:28 | CRANDALL et al. (2000), RUFFING et changes that produce reproductive incom- al. (2002) and AGAPOW et al. (2004). patibility. Eventually, genetically distinct Both the BSC and the PSC seek to taxa may look very similar and, contrarily, partition the natural variability observed in large morphological differences may exist biological communities into distinguish- between very closely-related taxa. The able components (species) but differ over BSC has also been criticised for its difficul- the partitioning criteria and the characters ty in dealing with introgressive hybridiza- used to delineate species boundaries. The tion and its inability to cope with asexual BSC emphasizes reproductive compatibili- reproduction (TURNER, 1999). ty among individuals within and among The PSC emphasises membership in a populations and defines a species as ‘a unique genealogical (phylogenetic) lineage group of interbreeding natural popula- and defines a species as ‘the smallest diag- tions which is reproductively isolated from nosable cluster of individual organisms other such groups’ (MAYR, 1940). This within which there is a parental pattern of definition is straightforward and provides ancestry and descent’ (CRACRAFT, one definitive criterion for assessing dis- 1983). Under this concept, criteria for tinctness of species – inability to inter- species distinctiveness are monophyly (all breed. Different species maintain their members of a group are descended from a genetic integrity because gene flow single common ancestor) and autapomor- between species is prevented by reproduc- phy (presence of genetically-based charac- tive isolation. One of the major criticisms teristics shared by all members of the group of this concept is that interbreeding capac- and not found in other groups). Subspecies ity can only be assessed in sympatry do not exist according to the PSC. In com- (MAYDEN & WOOD, 1995). In organ- parison to the BSC, the criteria employed isms with disjunct distributions the inter- to delineate species are more functional breeding criterion becomes non-opera- and testable – for example, the problemat- tional, since there is no practical way to ic notion of having to demonstrate repro- test whether individuals belonging to ductive isolation is avoided, while the exis- allopatric populations would be able to tence of phenetic dissimilarity is not a pre- mate and produce viable and fertile off- requisite for considering genetically dis- spring. Due to the inability of assessing tinct taxa as separate species. Major advan- reproductive isolation in allopatric taxa, tages of the PSC are its potential to handle the distinctiveness of species is usually asexual organisms and to delineate and inferred indirectly, e.g. from observation classify allopatric taxa. Main grounds for of morphology and distribution. Tradition- criticism are the difficulty in demonstrating ally, species have been defined by morpho- monophyly, the tendency of PSC applica- logical traits under the assumption that tions to produce excessive ‘splitting’ of morphological variation reflects genetic species, the recognition of species irrespec- variation which, when sufficiently high, tive of the degree of phenotypic or genetic may cause reproductive isolation. The con- divergence between evolutionary lines, and ceptual problem inherent in this assump- the absence of standardised approaches for tion is that morphological changes pro- selecting traits and/or determining levels of duced by variation and selection do not trait discreteness needed for diagnosing necessary correlate with the genetic species-level differences. In fact, two

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http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 09/03/2019 12:43:28 | groups would qualify for the status of sepa- populations, especially when: (a) taxa from rate species simply on the basis of differ- contrasting environments show trait differences in any single character, morphologi- ences that correlate with environmental cal or genetic, provided that all individuals differences; (b) the comparisons involve a within a group share one evolutionary line- limited number of individuals and few age. Questions may therefore arise as to characters; and (c) the examined traits which traits are appropriate for establish- show slight differences and/or overlapping ing species, whether species can be delimit- values. Again, it is left to competent taxon- ed on the basis of genetic differences alone, omists to judge if the kind, amount and and how much difference in the selected direction of observed differences in trait(s) is enough for distinguishing species. species traits permit discrimination in Another problem associated with the species-level identifications. PSC is how to deal with genetic introgres- Overall, application of the BSC sion, given the tendency of freshwater fish encounters methodological difficulties, to hybridise even at the generic level particularly with regard to demonstrating (SMITH, 1993). For instance, how will a reproductive isolation in allopatric popula- taxon formerly characterised as a distinct tions. The PSC is equally vulnerable to var- species be defined and named if extensive ious methodological difficulties, such as introgressive hybridisation has occurred sample data variability and reliability on due to repeated stockings? In a strict statistical grounds, and problems in assess- application of the PSC, hybrid populations ing phylogenetic lineages. It is also open to cannot be classified as belonging to any criticism for the objectivity with which the species because they are polyphyletic at species-discrimination criteria can be cho- the species level. In practical applications sen or applied. Taxonomies based on the of the PSC, it is left to the taxonomists to BSC contain few and often ambiguously evaluate evidence for the monophyly of defined species. However, these taxono- lineages and to decide whether or not the mies tend to maintain their stability over phyletic integrity of the taxon has been dis- time, because persistent and multiple- rupted by stocking beyond acceptable lim- character differences must be demonstrat- its. A last issue of concern relates to phe- ed before altering the taxonomic status of notypic plasticity. Many fish display taxa. By contrast, taxonomies based on the remarkable morphological variation, and PSC usually contain more species and are it is rarely clear what portion of this varia- sensitive to revisions and changes, because tion is based on inheritable genetic varia- the species are more loosely and variously tion and what portion reflects local envi- defined. ronmental features. Although plasticity LEE & WOLSAN (2002) attempted to may posses adaptive value and be under reconcile the apparently contradictory some genetic control (BAMBER & views concerning the conceptual frame- HENDERSON, 1988; JENNINGS & work and operational applicability of the BEVERTON, 1991; SWAIN et al., 1991), two concepts, arguing that the BSC refers an extensive expression of plasticity may explicitly to synchronic species, while the mislead taxonomic identifications. Cau- PSC refers explicitly to diachronic lineages. tion is therefore warranted when interpret- Therefore a biological species (synchronic ing the results of comparisons of allopatric species) and a phylogenetic species

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http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 09/03/2019 12:43:28 | (diachronic lineage) are ontologically dis- rather than changing each time a taxo- tinct entities. The authors proposed that it nomic study is undertaken. might be appropriate to use the term species solely for a synchronic, integrated Biodiversity conservation. The freshwa- assemblage of organisms, as defined by the ter fish fauna of Greece is characterized by BSC, and to apply the term lineage for a a high level of endemism, which leads to a diachronic, non-integrated assemblage concomitant need for a conservation focus. which is the historical product of past evo- Given the high number of species in need lution. Nonetheless, most authors perceive of protection and the relatively poor species as both synchronic and diachronic resources available for biodiversity conser- entities. Inevitably, the conflict over the vation, biologists and environmental man- merits and practicability of the BSC and agers are faced with the problem of decid- the PSC will continue to persist. In Greece, ing which taxa warrant special protection a trend towards acceptance of the PSC is using criteria such as the magnitude of the evident in almost all recent publications. extinction threat, ecological value or role in Below we shall discuss the consequences of the ecosystem and biological distinctive- this trend for some applied biological disci- ness. Assessing ‘biological distinctness’ is a plines. After all, an important factor to be challenging issue with biodiversity conser- considered in systematics is how useful a vation. Several authors have argued that species concept is relative to the needs of the PSC may promote conservation efforts the users of fish data. better than other concepts (e.g. KOTTELAT, 1997; GOLDSTEIN et al., Taxonomic standardisation and species 2000). On the one hand, the PSC allows the lists. Many users of fish data (e.g. fisheries recognition of genetically distinct popula- managers and administrative officers tions as evolutionarily significant units involved in commercial catch statistics or regardless of morphological similarity or regional development programmes) are the ability to interbreed. On the other not interested in taxonomy per se but on hand, the taxonomic identification of vul- the applications of taxonomy. For these nerable populations makes them a clear users, one of the more serious implications target for conservation effort on political of adopting the PSC is that the fish species grounds. Indeed, policy-makers recognize lists are large, differ among basin areas and give more value to ‘distinct’ named and must change constantly. An additional species than to lower taxonomic categories. difficulty arises when the species contained For example, the Aphanius population in the lists have been defined under differ- inhabiting the spring of Kato Almiri (NE ent species concepts and with a mixture of Peloponnese) is more likely to be strictly criteria. This may also be a cause of insta- protected than other Aphanius popula- bility since any re-evaluation of taxa tions, simply because it has been given the defined and named under the BSC may name A. almiriensis. Opponents of this view result in the definition of new species and consider that adoption of the PSC may names under the PSC. Such taxonomic have little effect on conservation. Reasons changes may have a disturbing effect on invoked include the species-centred waste most users of fish data, who would like to of limited resources and the difficulty of see the species list remaining constant deciding what should be conserved, given

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http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 09/03/2019 12:43:28 | the proliferation of species under this con- nised as separate species under the PSC cept; and, also the huge bureaucratic exhibit a similar range of ecological char- processes triggered to update conservation acteristics and tolerances to a variety of policies (e.g. GARNETT & CHRISTIDIS, anthropogenenic disturbances (ECONO- 2007). Taking position, we recognise that MOU et al., 2007). For example, all Squal- the PSC has various operational limita- ius species inhabiting Greek freshwaters in tions, such as the problem of handling Western Greece have similar ecological hybridisations, and also that taxonomic requirements with respect to rheophily classifications are unstable. However, we and habitat use. Likewise, all Salmo take into account that (a) many freshwater species of Greece exhibit similar require- fish populations in Greece have a long his- ments with respect to habitat, thermal tol- tory of isolation, often dating back to the erance and oxygen demands. Therefore, in Miocene, and thus may represent unique ichthyological research aimed at ecological genetic units, deserving immediate man- quality assessments, high-level classifica- agement consideration; (b) only ‘species’ tions based on the BSC are more mean- have high prominence under the national ingful than the less inclusive fine-level clas- and the EU conservation legislations, and sifications based on the PSC. (c) many sites harbouring threatened fish In conclusion, it appears no species populations or species (some until recently concept can fulfil all of the different recognized only as subspecies) have not requirements posed by the various disci- been incorporated in officially designated plines and applications. Each has its protected-areas (e.g. under the NATURA strengths and weaknesses and may work 2000 framework). On these grounds, we better in some situations, while the other consider that the PSC may better assist in may apply better in other situations. Given efforts for conserving Greek freshwater the different demands of potential users of fish, despite its various limitations. fish data, a possible solution is to use standardised lists of species incorporating finer Ecological quality assessments. Today and coarser taxonomies suitable for differ- one of the most active areas of ichthyolog- ent applications, so that studies based on ical research in Greece is the use of ichthy- such lists can at least be consistent. Never- ological indicators for the assessment of theless, caution is needed in applications the ecological status of freshwater water- of the PSC concept in species descriptions; bodies, in accordance with the provisions otherwise the taxonomy will become un- of the EU Water Framework Directive manageable and vulnerable to disruptive 2000/60. The principle behind the applica- changes. tion of fish-based bioassessment methodologies is that freshwater fish assemblages Distribution pattern of native freshwa- reflect structural and functional aspects of ter fish aquatic ecosystems. Therefore, the important issue for applying such methodologies Ancient arrivals, travel routes and barriers is not the genetic discreteness of taxa, but to dispersal rather their ecological properties and ability to diagnose ecological degradation. Two major and perhaps non-exclusive Many closely related taxa that are recog- explanations have been proposed for the

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http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 09/03/2019 12:43:28 | arrival and dispersion of freshwater fish 2004; BOBORI & ECONOMIDIS, 2006; in the Balkans. One postulates that Euro- SKOULIKIDIS et al., 2008): (a) the grad- Siberian and Palaearctic species reached ual uplift of the Alps and the Balkan the area during Oligocene and Miocene Mountains from late Oligocene to the times through river captures (ECO- end of the Miocene isolated the Balkan NOMIDIS & B N RESCU, 1991). A drainages preventing faunal exchanges second wave of arrivals of central Euro- with the rest of Europe; (b) the rise of the pean and Danubian species that intruded Pindos mountain range created a north- into the area from the Danube and the west-southeast barrier for fish range Black Sea during Pliocene or Pleistocene expansions, while the rise of the Mount times is also postulated by the above Othrys cut the connections of the rivers of authors. The alternative hypothesis holds central-eastern Greece from those of that colonisation of freshwater species northern Greece; (c) at the around the circum-Mediterranean may Pliocene–Pleistocene boundary, a com- have occurred during a short period of munication of the NW Aegean drainages the late Miocene when the Mediter- with the Danube R. was temporarily ranean dried up completely and then was established through a river-capture partially refilled with freshwater from the involving the Morava R. and the Axios Paratethys (BIANCO, 1990). It seems R.; (d) at about the same time, a similar that no single explanation can account for communication of the Adriatic drainages the diversity of the Balkan ichthyofauna, with the Danube R. was established and of the Greek ichthyofauna in particu- through a river capture involving the lar (B N RESCU, 2004). Different Ohrid-Drim-Skadar system in the area of species may have arrived in different Kosovo; (e) also in Plio-Pleistocene times and through different pathways, times, intrusion of Black Sea waters (then and may have experienced various a freshwater lake) into the Mediter- degrees fragmentation and isolation ranean through the former Aegeopota- (ZARDOYA et al., 1999). Isolation, com- mos R. permitted dispersal of Black Sea bined with complex climatic events, pro- freshwater fish to the NE Aegean moted speciation and produced a great drainages; and (e) sea-level regressions at variety of endemic taxa. During periods the glacial maxima of the Pleistocene had of intense tectonism and marine regres- a homogenising effect on fish assem- sion secondary contacts of previously iso- blages allowing dispersal among neigh- lated populations may have occurred, bouring river basins. resulting to hybridization and genetic During the Pleistocene glaciations the introgression (e.g. DURAND et al., Greek rivers remained free of ice, serving 2003). The following vicariant and disper- as refugia for the preservation of ancestral sal events have been proposed to account elements of the European ichthyofauna, for the structural diversity and high which were eradicated from most other degree of endemicity of the Greek ichthy- parts of Europe. In the post-glacial times, ofauna (see ECONOMIDIS, 1974; the Greek rivers did not contribute to the ECONOMIDIS & B N RESCU, recolonization of the European rivers with 1991; ZARDOYA et al., 1999; freshwater fish; consequently, the fish fau- DURAND et al., 2003; B N RESCU, nas of the Greek rivers have retained their

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http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 09/03/2019 12:43:28 | unique endemic forms which thus repre- larity among regions (Table 9). Species sent distinct taxononomic entities different richness and endemicity levels also differ from the ones in the rest of Europe. among biogeographic regions. The western, central and southern parts of Greece Regional fish assemblages and endemicity (Ionian and Attiko-Beotia regions) hold patterns an old and long-isolated ichthyofauna, and present low species richness but a The present-day fish composition has high degree of endemicity. The northern been determined by a combination of vic- and eastern parts of the country present ariant and dispersion events and faunal higher richness and lower endemism, relaxation by extinction episodes. The most probably because these parts are in uplift of the Pindic Cordillera in the Mid- great proximity to the dispersal areas of dle Miocene (DERMITZAKIS & PAPA- the Danube and the Black Sea. Species NIKOLAOU, 1981) acted as a major fac- depauperation in East Peloponnese and tor for the faunal divergence of the west- the Aegean Islands makes faunistic com- ern and eastern Greece. Consequently, parisons difficult and particularly chal- two major aquatic biogeographical divi- lenging. Nonetheless, the data indicate a sions are unanimously recognised, defined faunistic distinctness of the Aegean region with different names by various authors, that most likely has its explanation in and referred to here as the Aegean and the independent origins of species. So far it Ionian divisions. However, opinions differ has been difficult to determine the faunis- over the number of minor divisions and tic relationships of the East Peloponnese their boundaries (STEPHANIDIS, 1939; with the other regions because only few, BIANCO, 1986, 1990; ECONOMIDIS & poorly studied fish are present in this B N RESCU, 1991; MAURAKIS & region. A general difficulty in performing ECONOMIDIS, 2001; MAURAKIS et al., biologically relevant comparisons among 2001). ECONOMIDIS & B N regions is that many areas in central and RESCU (1991) distinguished four main south-eastern Greece and in the Aegean ichthyogeographic divisions (regions) in Islands are in a bioclimatic semi-arid the Balkans of which three encompass the zone, where few species have survived Greek territory: the Ponto-Aegean (con- prolonged drought episodes or recent taining the subdivisions Thracian-East human water abstraction impacts. Macedonia and Macedonia-Thessaly), the The North Aegean basins harbour Attiko-Beotia, and the South Adriatic-Ion- many species with Black Sea and Danu- ian. B N RESCU (2004) retained this bian affinities (B N RESCU, 2004). ichthyogeographic scheme with some Despite many common faunistic elements slight modifications, e.g. he charted the over the entire North Aegean region (e.g. East Peloponnese as a separate entity. Scardinius erythrophalmus, Rutilus rutilus, The fish distributional data presented Silurus glanis) the Thracian rivers (east- in the present paper corroborate the wards of the Strymon R.) and the Mace- above biogeographical separation of donian-Thessalian rivers show ichthyologi- Greece indicating the presence of charac- cal distinctiveness that would justify their teristic endemics in each region (Appen- placement in different ichthyogeographic dix I) and a low degree of faunistic simi- regions. As pointed out by ECONOMI-

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http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 09/03/2019 12:43:28 | Table 9 Faunistic similarities among biogeographical regions (fish species with joint presence in more than two regions).

Fish species North Attiko- Ionian South East Aegean Aegean Beotia Adriatic Peloponnese Islands Alburnoides bipunctatus Alosa fallax Anguilla anguilla Aphanius fasciatus Barbus sperchiensis Chondrostoma vardarense Gasterosteus gymnourus Knipowitschia caucasica Pelasgus stymphalicus Petromyzon marinus Rhodeus amarus Salaria fluviatilis Salmo farioides Scardinius erythrophthalmus Squalius vardarensis

DIS & B N RESCU (1991) most fish contact with the former Paratethys Sea in occurring in the Thracian rivers are inhab- Miocene times (e.g. SONNENFELD, itants of still or slow-flowing waters and 2005). All Macedonian-Thessalian rivers may have reached the area chiefly from the show remarkable faunistic similarities, as Black Sea during its freshwater phase. expected, given that these rivers were con- Nonetheless, dispersal opportunities have nected during the last glacial maximum existed at least since the Miocene (LYKOUSIS et al., 2005). (SAKIN & YALTIRAK, 2005). Barbus The Ionian region is considered as one strumicae and Squalius orpheus are two of of the most isolated zoogeographic units in the most characteristic endemics of the Europe, since it is blocked from the rest of Thracian ichthyofauna. the Balkans by mountain ranges. This The fish inhabiting the Macedonian region contains unique endemics that are and Thessalian rivers show distant affini- often confined to one or few drainages ties with Danubian fish and may have (TSIGENOPOULOS & KARAKOUSIS, entered the area via the Axios R. in Plio- 1996; BARBIERI et al., 2002; Pleistocene times, as previously discussed. KETMAIER et al., 2003; MILLER et al., Indeed, several species endemic to this 2004a; ECONOMOU et al., 2004c). area (e.g. Cobitis vardarensis, Squalius var- Species of some genera (e.g. Squalius, darensis, Pachychilon macedonicum, Scardinius and Barbus) show deep genetic Rhodeus meridionalis and Zingel balcani- divergence from their counterparts in the cus) have sister group relationships with Balkans and the rest of Europe and often Danubian fish species. However, the first have a basal or almost basal position in arrivals might have occurred quite earlier, phylogenetic reconstructions (DOADRIO as it seems that the area had hydrological & CARMONA, 1998; KETMAIER et al.,

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http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 09/03/2019 12:43:28 | 1998, 2003; DURAND et al., 1999a, 1999b; instance, the present-day distribution of TSIGENOPOULOS & BERREBI, 2000; Squalius keadicus is restricted to Laconia TSIGENOPOULOS et al., 2002). The dis- (Evrotas and Vassilopotamos Rivers). tinctness and ancient origin of the Ionian However, there is evidence from genetic ichthyofauna is further indicated by (a) the studies that the historic range of this presence of endemic genera (Tropidophox- species was wider and included rivers of inellus and Economidichthys) (BIANCO et south-west Peloponnese from which it was al., 1987; STEPHANIDIS, 1974) and (b) extirpated by introgression with new the absence of widespread European gen- Squalius invaders (DURAND et al., 2000). era that are typically present in other The Attiko-Beotia region is a diverse Balkan regions (Chondrostoma, Barbatula, area which seems to be a true ‘genetic Gobio, Alburnus, Alburnoides, Phoxinus, crossroad’, as species have presumably Cottus and Rhodeus). Interestingly, some emigrated in both from the north, west and Ionian species show much closer relation- east, however they have been isolated long ships to Iberian species than to species enough to show differentiation and specia- inhabiting other Balkan regions (DO- tion. The rivers of this region share few ADRIO & CARMONA, 1998; ZAR- only species with the North Aegean and DOYA et al., 1999; PERDICES et al., the Ionian rivers (Table 9) and are inhab- 1996; SANJUR et al., 2003). Geological ited by a depauperate freshwater fish events that have contributed to isolation fauna that includes distinctive endemics. and speciation in the Ionian region The fluvio-lacustrines Luciobarbus grae- include: the early isolation of the southern cus, Scardinius graecus and Rutilus ylikien- part of Peloponnese by mountain barriers sis are confined in the Kifissos R. system and deep seas; the separation of the Pelo- and are probably remnants of the ichthy- ponnese by the opening of the Corinthian ofauna of the ancient lake Kopais, now Gulf during the early part of the Late drained. Several species of the Attiko- Pliocene; the progressive uplift of the Ion- Beotia region (of the genera Luciobarbus, ian islands throughout the Pliocene; and Rutilus, Pelasgus, Telestes and Scardinius) the entrance of seawater in the Patraikos have sister species in the Ionian region, Gulf during the Holocene. The existence which may reflect the past hydrological of paleo-lakes in Epirus, Acarnania, Arca- connection of the Sperchios basin with the dia and Laconia allowed the maintenance Amvrakikos Gulf in Miocene times. of old fish lineages. The confluence of the Today’s ichthyofauna is only a relict of the Epirus rivers, and the similar confluences past. An unclarified cyprinid, identified as of the Acarnanian rivers, permitted faunal a Squalius taxon, geographically isolated exchanges in the Pleistocene. from other Squalius taxa, inhabited the The breadth of fish diversity in the Beotian Assopos (STEPHANIDIS, 1974). Ionian region is easily underestimated if This species presumably disappeared one counts only the number of species before it was scientifically studied and sim- present. A larger and yet poorly explored ilar extinctions may have occurred in rivers amount of diversity exists below the of Attiki and Euboea, which have been species level, and is represented by unique severely impacted by human activities. phenotypes and genetic profiles often Although the knowledge of the showing a north-south clination. For Aegean island’s fish fauna is still incom-

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http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 09/03/2019 12:43:28 | plete, the available evidence suggests that Species richness and species – area rela- the eastern islands show faunal affinities tionships with Asia Minor. A distinctive endemic in this region is Ladigesocypris ghigii, inhabit- High species richness generally corre- ing streams of Rhodes Island, which has lates negatively with the degree of lost its connection with Anatolia in the endemicity, except in the Aegean Islands Pliocene (DERMITZAKIS, 1990). By and the East Peloponnese, where fish fau- contrast, other eastern Aegean islands, nal depauperation does not permit mean- including Lesvos and Samos, remained ingful comparisons. This is particularly connected to Anatolia until Pleistocene true when the fish faunas of the Ionian and times, and fish colonisations from the the North Aegean regions are compared. mainland, especially during marine Lower richness in the Ionian region (Table regressions, were possible. The ichthy- 7a; Fig. 4) may have a historical explana- ofauna of the East Peloponnese is tion: the Ionian fish faunas have probably extremely depauperate and not yet prop- remained isolated since the Miocene, and erly studied. In the 1990s an Aphanius therefore may have been subjected to population was discovered in the Kato extinction processes for a longer time than Almiri spring (ECONOMOU et al., 1997). the North Aegean fish faunas, which had This population was provisionally Pliocene or more recent contacts with the assigned to A. fasciatus, though the Danubian and Black Sea faunas. However, authors noted its morphological and this explanation should be considered with behavioural distinctiveness from the latter caution because the low species richness in species. Later, this taxon was described as the Ionian hydrographic basins may be due A. almiriensis (KOTTELAT et al., 2007). to ecological or physiographic factors. Our Whereas in the 1990s this species was data show an increase of species richness moderately abundant, recent investiga- with increasing basin size (see Fig. 3), con- tions have failed to show its persistence in firming long-standing generalisations that the site. more species exist in larger basins, either It is safe to conclude that the remark- because such basins contain a wider array able diversity of the fish assemblages of habitat configurations, or because the among biogeographic regions has a histor- probability of extinction is more likely in ical explanation (vicariance and isolation). small basins (REYJOL et al., 2006). Given It is interesting that many cases of shared the positive correlation between species species presences indicated in Table 9 con- richness and basin size, a logical explana- cern secondary freshwater or peripheral tion of the lower richness of the Ionian fish with the ability to utilise the marine basins is that their size is much smaller route for their dispersal (Aphanius fascia- than the size of the North Aegean basins. tus, Knipowitschia caucasica, Salaria fluvi- However, local ecological conditions and atilis, Anguilla anguilla, Alosa falax, Gas- landscape features often disrupt the rich- terosteus gymnurus and Petromyzon mari- ness-basin area correlation. For instance, nus). This is especially the case in the the spring-fed Louros R. hosts a larger Aegean Island region, where all species number of species (14 species) than the shared with other regions are peripheral or adjacent and much larger nivo-pluvial secondary freshwater species. Arachthos R. (11 species). Presumably,

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http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 09/03/2019 12:43:28 | hydrological stability and the presence of scheme and an important reason is the more extensive floodplain habitats in the information deficit on the organisms to be former river have resulted in a greater used as biotic quality elements (ECO- availability of habitat types and/or higher NOMOU et al., 2006). As of late 2007, ability of species to persist in the long Greece had not yet delineated river basin term. areas or constituent waterbodies, so even That both explanations may hold is not the basic geographic framework for eco- controversial. Richness may depend on logical quality assessments is still missing. both demographic processes (colonisation The river basin area distributions provided and extinction events) and ecological or here can contribute to bioassessment tool physiographic factors. When basins of sim- development in various ways, e.g. by ilar size are compared, those of the Ionian enabling the characterisation of historical region contain fewer species than basins of reference conditions and the selection of the North Aegean region, suggesting that appropriate metrics. They also reveal two historical factors and demographic kinds of difficulties in building robust fish- processes had an important structuring based bioassessment indices. The first kind effect on regional fish assemblages. Within relates to the faunistic idiosyncrasies and regions, however, physiographic factors, heterogeneity that characterised the such as basin size and relief, may better Greek hydrographic basins (low basin explain the observed richness patterns. For species richness, high degree of endemici- instance, most rivers of North Pelopon- ty, and varied basin-specific taxa assem- nese are small and have high slope, lacking blages). Such aspects of the ichthyofaunal floodplains; these rivers host only one or assemblages are characteristic of the high- two fish species that are tolerant to flashy ly heterogeneous Mediterranean-climate and erosive stream conditions. The depau- environmental conditions (GASITH & peration of the fish faunas of East Pelo- RESH, 1999; FERREIRA et al., 2007) ponnese and the Aegean Islands can simi- and, among others, restrict the number of larly be attributed to small basin size. potential metrics or prevent the applica- Their isolated fish communities are partic- tion of a common metric system over wide ularly vulnerable to repeated drying areas. The second kind of difficulties aris- episodes, and many historical extirpations es from taxonomic uncertainties, insuffi- may have taken place. cient knowledge of the species’ distributions, and the wide tolerance of many Policy relevant implications of the survey native species to varying environmental conditions. The adoption of the phyloge- River basin area management and WFD netic species concept (PSC) has exacerbat- application ed the problems of reference conditions and metric development because the As previously expressed, ecosystem application of a finer taxonomy generates health assessment and monitoring is one an ‘apparent’ increase of the biological aspect of aquatic conservation where fish heterogeneity among hydrographic basins. play an important role as biological indica- However, bearing in mind that phyloge- tors. Greece is lagging very much behind in netically closely related species are more the application of the WFD assessment likely to be ecologically similar, a solution

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http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 09/03/2019 12:43:28 | to these problems is to identify ecological- recent name changes, we are faced with a ly-equivalent species that respond to eco- remarkable number of localized endemic logical degradation in a similar manner. species as well as of taxa of unclear taxon- Base-line research projects for fish assem- omy. Our basin survey identifies a large blage community ecology is vital for estab- number of range-restricted species that lishing and standardising lists of ecologi- need conservation attention. For some cally-similar species. species complete distributional data are provided that can assist in conservation re- Species conservation assessments. For a number of other species, however, the survey indicates gaps Species entities are basic units in bio- in the knowledge of their distribution diversity research and conservation appli- and/or problems with their taxonomic sta- cations. Experience has shown that the tus. Distributional surveys and taxonomic more ‘distinct’ a species is, in relation to work are essential for planning conserva- other species, the greater the priority is tion-orientated research. given to its protection (MAITLAND, 2004). At the Mediterranean scale the Habitat conservation number of threatened species is very large and encompasses about 56% of the While it is widely accepted that saving endemic fish; this is one of the highest pro- the sites where vulnerable species live is a portional assessments of threatened very important aspect for their conserva- species worldwide (see SMITH & tion, current legislation does not include DARWELL, 2006; PETER, 2006). The satisfactory provisions for sites and aquat- distributional ranges of species are a key ic habitat types. For example, the Habitats issue for the assessment of their vulnera- Directive does not provide adequate bility and the design of protective meas- descriptions or classifications of the array ures. In Greece, little work has been done of aquatic habitat types which exist in to systematically document the distribu- inland freshwaters, since mostly terrestrial tions of endemic species; most studies pro- habitat types are listed and targeted for vide anecdotal accounts of species ranges conservation (DIMOPOULOS et al., and are not quantitative with respect to 2005). This is unfortunate, because it is abundance or density (ECONOMIDIS, well known that freshwater habitats belong 1995). Problems of this kind have impeded to the world’s most threatened ecosystems conservation efforts. As a consequence, (SMITH & DARWELL, 2006). These conservation priorities are usually based important weaknesses in current legisla- on fish assemblage data available at the tive coverage or enforcement of protection national level (i.e. check-lists) or regional of aquatic habitats and their biota have level (i.e. protected-areas), rather than on resulted in the ‘exclusion’ of many small detailed basin area-based fish compila- and highly vulnerable sites from the exist- tions. Therefore, it is not surprising that ing protected-area network of Greece. many endangered fish species are still not Our basin-based compilation of fish distri- given appropriate legal protection, e.g. butions indicates the occurrence of basins they are not present in Annex II of the containing potentially important evolu- Habitats Directive. Besides, with the tionary units. However, the compilation

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http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 09/03/2019 12:43:28 | does not detail the exact limits of the geo- least as much importance as economic graphic distribution of each unit within and social factors. basins. Further analyses and screening of site-specific data will be required to ensure Biogeography the preservation of independent genetic pools and to highlight unmet conservation Since freshwater fish are restricted to gaps on a nation-wide scale. pathways offered by hydrogeographic systems, their distribution largely reflects his- Fishery management torical patterns of drainage connections (e.g. VARGAS et al., 1998; REYJOL et Inland waters fisheries are facing sig- al., 2006). Moreover, fish do have better nificant problems due to mismanagement, documented distributional information water quality problems and the effects of than do most other freshwater-obligate invasive alien fish species (CRIVELLI et organisms in Greece (i.e. invertebrates, al., 1997; LEONARDOS et al., 2007). amphibians); therefore, they are capable Destructive fishing practices lead to dam- of supporting biogeographical analyses aged fish habitat and less fish, and species (LEGAKIS, 2004). The influence of his- introductions and translocations cause toric drivers determining fish distribution genetic pollution, representing a major is especially important in the species reason of the degradation of native gene extinction-invasion process. Many biogeo- pools. Misinformed fishermen and fish- graphic applications can be developed eries managers are definitely the largest when a complete inventory of fish species cause of alien fish species spread in occurring in the hydrographic basins of Greece (ECONOMIDIS et al., 2000a). In Greece will be created (for instance, planning and enforcing fisheries manage- assessing the relative importance of his- ment policies, it is important that reliable torical factors and physiographic or data on the composition of fish assem- hydrological characteristics in determin- blages and the native ranges of species are ing basin area fish assemblages). We available. The present dataset provides acknowledge that our database still needs baseline ichthyological information that verification and completion; nevertheless may help to track the spread of alien it is a first step in developing a nation- species and to report vital ichthyological wide basin-based inventory that can be data in a standaridised manner. However, used for conservation-relevant biogeo- commercial and sport fishing have major graphical research. economic and political implications that also need to be taken into account in the Research and management priorities watershed management plans. While it is clear that holistic approaches covering Below we summarize five imperative social, economic, environmental and tech- actions with respect to conservation-rele- nical aspects should be used to promote vant ichthyological research in Greece: fishery management, biodiversity issues should not be sacrificed for the sake of 1. Fish distributional surveys. Our distribu- development; the conservation value of tional data clearly suggest that conserva- species and habitats should be given at tion and management programmes should

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http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 09/03/2019 12:43:28 | refer to the geographic ranges of each tax- R. and various Knipowitschia populations onomic unit to ensure the preservation of in western and central-eastern Greece). independent genetic pools. Distributional Particular attention is needed in the data are also critical for the implementa- boundary areas between species where tion of the WFD (establishment of refer- genetically distinct units can occur. For ence conditions, metric selection, site- example, the progress of recent Squalius based index development), the Habitats peloponnensis populations and the colo- Directive (site protection and monitoring, nization of new aquatic systems might have populations reporting, conservation man- taken place in successive waves. Perhaps in agement) and basic environmental impact some systems of southern Peloponnese assessment. Likewise, fisheries manage- (e.g. in the SW Messinia streams or in the ment should be seen as a conservation montane plateau of the Lousios R.) the issue that needs planning and enforcement new invaders found old local Squalius on the basis of distributional and informa- stocks and either eliminated them or tion assemblage structure. Ichthyological mated with them. Similar hybridisation research is especially needed in small zones may exist in the boundaries of water features such as springs, wetlands species belonging to the genera Salmo and and coastal lagoons, as well as in deep sec- Pelasgus. tions of large rivers, which have not been adequately sampled. An important initia- 3. Aquatic habitat inventory. Habitat loss is tive to integrate the aforementioned needs the most important conservation problem would involve a coordinated atlas project for fish species, and this is especially acute for freshwater fish in Greece. in seasonally-semi-arid environments where many small watersheds are vulnera- 2. Fish taxonomy - genetic research. Taxon- ble to human pressures. Greece and the omy and conservation must come togeth- western Balkans have one of the largest er; and taxonomists who are motivated by concentrations of range-restricted species; conservation action must strive to produce many species are restricted to one or two reliable standardized taxonomic units. river basin areas – some are confined to Obviously, genetic research is critical in certain river segments or special habitat taxonomy. A community of taxonomists types (spring-fed wetlands, lakes, ponds must develop and a forum should be creat- etc). Coordinated efforts are needed to ed in order to help establish the validity of document aquatic habitats in an inventory taxa (CRIVELLI & MAITLAND, 1995). and to create wide-ranging campaigns for Genetic variation in any species being con- their preservation and restoration. As stat- served or managed is very important to ed by CRIVELLI & MAITLAND (1995), monitor. Without this basic research it is the conservation of freshwater habitats is impossible to effectively manage species, more important than that of individual populations or communities of fish. Some species; but to conserve these effectively isolated populations are poorly studied, we need: a) a list of all aquatic habitat and yet they may represent cryptic endem- areas, b) priority ranking and conservation ic species that may be worth protecting as evaluation of the habitat areas, c) conser- significant evolutionary units (e.g. vation management plans for sites and the Alburnoides bipunctatus in the Sperchios fish species accommodated within them.

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http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 09/03/2019 12:43:28 | 4. Alien species control. Alien species are aquatic biota. It is therefore important to widely considered as the second most promote legal protection schemes and spe- important threat to aquatic biodiversity cial management initiatives to keep aquat- after habitat loss. Awareness and enforce- ic habitats in existence (APERGHIS & ment of the control of the spread of intro- GAETHLICH, 2006). Carefully sited produced species is critical for conservation. tected-areas are needed in order to cover We have provided evidence that 25 alien linear aquatic features or focus on relative- species have been introduced to Greece of ly small sites and threatened population which some are spreading fast, seriously refuges (e.g. MOYLE & YOSHIYAMA, impacting the natural biota. Translocation 1994). Micro-reserves may be effective as a of fish among basins is an equally serious short-term direct protective measure. problem, since they are usually performed without any concern for the evolutionary Conclusion history of the species (for trout species: see APOSTOLIDIS et al., 1999). The impact Hydrographic basin areas are of high of translocations on genetic diversity may relevance to current water policy and con- exceed the impact of alien species intro- servation. Such areas have well defined ductions due to the high possibility of boundaries (watershed limits) and are introgressive hybridization between popu- becoming important for effective aquatic lations or closely related species. In the ecosystem monitoring, assessment, report- light of the recorded human effect on the ing and management. In this work, a basin distribution of the different species, it is area survey method was employed to com- desirable that both the donor populations pile the best-available distributional and the indigenous populations in the assemblage data for freshwater fish in recipient areas would be genetically Greece. Our work is a preliminary but screened before any introduction. Public wide-ranging attempt that may help to awareness of this problem is extremely identify: (a) unmet needs in our under- important. A research priority is to identi- standing of freshwater species’ distribu- fy areas hosting unaffected remnant tions in Greece, (b) problems with the indigenous fish stocks that can be pre- species’ recent taxonomic changes, and (c) served and used as a population source for basic gaps in science-based conservation rehabilitation projects. work on threatened species. Indeed, one of the major obstacles in effectively assess- 5. River basin management plans. A multi- ing the conservation status of fish, their tiered approach to biota and habitat pro- habitat needs, and the anthropogenic pres- tection must be incorporated in river basin sures they may face, is the large gap in management plans. Integrated and holistic knowledge of their geographical distribu- planning is needed to co-ordinate water tions. resource exploitation, conservation and A coordinated effort is needed to pro- restoration in basins; however, at this mote field ichthyology as a scientific enter- scale, biodiversity is not often given appro- prise that is directly useful to nature con- priate consideration. Indeed, as already servation. There is still poor baseline stated, traditional protected-areas often knowledge of Greece’s inland water aquat- disregard fish and other elements of the ic ecosystems. The number of different

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http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 09/03/2019 12:43:28 | aquatic habitat areas in Greece is large; as graphic compilation with references, dis- geographic entities, aquatic habitat sites cussions or fish data. Many colleagues certainly number in the thousands. We helped in fieldwork; we are indebted to U. have very little information for many of the Dussling, K. Blasel, G. Dal ge, R. Bjork- smaller more isolated ones, and the quali- land, P.J. Miller, B. Zimmerman, M. Kot- ty of the information for many parts of the telat and W. Beaumont for assistance in larger river basin areas is also poor. wide-ranging surveys. At HCMR we espe- Reviews from other inland water assess- cially thank colleagues Y. Chatzinikolaou, ments from the Mediterranean have simi- V. Tachos, E. Kalogianni, C. Daoulas, T. larly depicted the problem of important Psarras, N. Koutsikos and D. Kommatas, gaps in the inventory of ecosystems and who participated in field surveys, and N. their aquatic biota (ALVAREZ- Skoulikidis, A. Zenetos, P. Drakopoulou COBELAS et al., 2005), so this is not a and E. Moussoulis, who contributed in problem unique to Greece. However, the many other ways. We express our thanks ichthyofauna of Greece is especially rich in to E. Economou, who coordinated data- many range-restricted species and needs quality control and detailed reference immediate attention. Particular problems cross-checking. We appreciate the graphic concern conservation-relevant aspects of work done by A. Vidalis. Thanks are also its taxonomy, biology and ecology. Impor- due to E. Tzovara for her time and tant problems also exist with information patience in editing this manuscript. Final- management and dissemination. This ly, we are grateful to W. Beaumont for work also underlines the urgent need for constructive comments and editing and to building site-based inventories of fish B. Zimmerman and Z. Kaczkowski for assemblages in this country. reviewing drafts of this paper. Obviously, any errors during this compilation and its Acknowledgements presentation are the responsibility of the authors or are based on their opinion at This work is a product of extensive col- the time of writing. This work was partial- laboration with many scientists and natu- ly funded through a project of the Hellenic ralists who have shared data and reviewed Ministry of Development and we are fish species lists with us during the last five grateful to M. Ghini for her assistance in years. The authors have greatly benefited this cooperation. Lastly, the Department from the landmark works of P.S. Econo- of Environment and Natural Resources midis; constructive advice and discussions Management at the University of Ioannina with him are highly appreciated. Notable supported electrofishing field survey work assistance was provided by D. Bobori, D. involving the last author. Mdrak, A. Flloko, A. Apostolou, A. Legakis, P. Dimopoulos, Th. Naziridis, K. References Poirazidis, Y. Kazoglou, Ch. Papaioannou, Y. Leonardos, Y. Roussopoulos, C.G. ABELL, R., M. THIEME, E. DI- Papaconstantinou, A. Crivelli, G. Catsado- NERSTEIN, & OLSON D., 2002. A rakis, A. Apostolidis, G. Tingilis, Y. sourcebook for conducting biological Reyjol, M.T. Ferreira and A. Vlamis- assessments and developing biodiver- Gardikas who contributed to the biblio- sity visions for ecoregion conservation.

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Accepted in 2007

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http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 09/03/2019 12:43:28 | (continued) 111 NENNO OTHER NENNO 1 (ENWE) 010 OTHER ENNO ENNO Alburnoides bipunctatus prespensis Alburnus alburnus thessalicus, Alburnus alburnus macedonicus Chalcalburnus belvica Eudontomyzon hellenicus Alosa (Caspialosa) macedonica Alosa (Caspialosa) caspia vistonica Alosa fallax nilotica NO CHANGE 1 ENNO Pallas, 1771 NO CHANGE 0 OTHER Undescribed Brandt, 1869 NO CHANGE (1) ALN (Bloch, 1782) NO CHANGE 1 OTHER Linnaeus, 1758 NO CHANGE 0 OTHER Linnaeus, 1758 NO CHANGE 0 OTHER Linnaeus, 1758 NO CHANGE (1) ALN Karaman, 1924 Bonaparte, 1836 NO CHANGE 0 OTHER Bonaparte, 1845 NEW 1 ENBAL (Linnaeus, 1758) (Linnaeus, 1758) NO CHANGE 1 OTHER (Linnaeus, 1758) NO CHANGE(Linnaeus, 1758) NO CHANGE 0 (ALN) 0 OTHER (Karaman, 1924) (Walbaum, 1792) NEW 1 ALN Economidis, 1982 (La Cepède, 1803) (Vinciguerra, 1921) Economidis & Sinis, 1986 Brandt & Ratzeburg, 1833 NO CHANGE 0 ALN Vladykov, Renaud, Kott & Appendix I. Annotated Checklist of Freshwater Fishes of Greece. 2 1 5 Louros 3 sp. Name Authority Previous Nomenclature Freshwater Endemic scoranza 4 cf. Huso huso Acipenser stellatus Alburnus alburnus Alburnus belvica Alburnus Acipenser sturio Petromyzon marinus Acipenseridae Acipenser baeri Acipenser gueldenstaedtii Acipenser naccarii Acipenser ruthenus Alosa macedonica Alosa vistonica Cyprinidae Abramis brama Alburnoides bipunctatus Alburnoides prespensis Polyodontidae Polyodon spathula Anguillidae Anguilla anguilla Clupeidae Alosa fallax Petromyzonidae Eudontomyzon hellenicus Eudontomyzon 10 8 19 20 21 9 3 4 5 6 7 14 15 16 17 18 11 12 13 12 3 4 56 1 2

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http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 09/03/2019 12:43:28 | (continued) 1 ENBAL 1 ENBAL 1 OTHER 1 ENNO 1 ENBAL 111 ENBAL ENNO ENNO 11 ENNO ENBAL 11 OTHER ENBAL 1 NENNO 11 (ENWE) ENBAL Barbus barbus macedonicus, Barbus barbus thessalicus Barbus cyclolepis cyclolepis Barbus peloponnesius petenyi Gobio gobio feraensis Gobio gobio bulgaricus, Gobio gobio balcanicus Alburnus alburnus thessalicus Chalcalburnus chalcoides macedonicus Chalcalburnus chalcoides macedonicus Barbus cyclolepis sperchiensis, Barbus cyclolepis cholorematicus Barbus cyclolepis strumicae Carassius auratus gibelio Chondrostoma vardarensis Alburnus alburnus macedonicus Barbus peloponnesius rebeli Barbus peloponnesius peloponnesius b & ’ Appendix I (continued) Koller, 1926 In K&F2007 NEW 1 (ENNO) Heckel, 1837 Berrebi, 2002 (Bloch, 1782) Drensky, 1926 Linnaeus, 1758 NO CHANGE 1 OTHER Karaman, 1928 Karaman, 1955 Karaman, 1924Karaman, 1928 NO CHANGE 1 NENNO Karaman, 1928 Karaman, 1936 NEW 1 ENBAL Karaman, 1971Karaman, 1924 NEW NO CHANGE 1 1 NENNO OTHER (Linnaeus, 1758) NO CHANGE 1 OTHER (Linnaeus, 1758)(Linnaeus, 1758) NEW NO CHANGE 1 1 OTHER ALN Stephanidis, 1950 NO CHANGE 1 ENCE Stephanidis, 1973 Stephanidis, 1950 Stephanidis, 1950 Valenciennes, 1842 (Valenciennes, 1844) NO CHANGE 1 ALN (Valenciennes, 1844) NO CHANGE 1 ALN Freyhof & Kottelat, 2007 Freyhof & Kottelat, 2007 Kotli k, Tsigenopoulos, Ra 9 14 7 10 6 8 11 12 13 Name Authority Previous Nomenclature Freshwater Endemic skadarensis sp. Volvi cf. Barbus peloponnesius Barbus macedonicus Barbus cyclolepis Barbus euboicus Hypophthalmichthys molitrix Gobio Alburnus vistonicus Alburnus volviticus Aspius aspius Barbus strumicae Carassius auratus Barbus balcanicus Carassius carassius Carassius gibelio Alburnus thessalicus Chondrostoma vardarense Chondrostoma prespensis Ctenopharyngodon idella Cyprinus carpio Alburnus macedonicus Alburnus Gobio feraeensis Barbus prespensis Barbus rebeli Barbus pergamonensis Barbus sperchiensis Gobio bulgaricus 32 31 29 30 48 46 25 26 27 37 38 28 39 40 24 42 41 43 44 12 3 4 56 22 23 47 34 35 33 36 45

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http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 09/03/2019 12:43:28 | (continued) 111 ENWE NENNO OTHER 111 OTHER 1 (ENNO) OTHER ENBAL 111 ENAEG 11 ENWE ENCE 1 ENBAL ENBAL 11 ENWE 11 ENCE 1 NENNO ENWE ENWE+ OTHER 1 ENBAL Rutilus ylikiensis Rutilus ohridanus prespensis Rutilus rutilus mariza, Rutilus rutilus dojranensis, Rutilus rutilus vegoriticus Phoxinus phoxinus Phoxinus phoxinus Rhodeus sericeus amarus Rhodeus sericeus amarus Pachychilon pictus Barbus albanicus Barbus graecus Rutilus macedonicus Paraphoxinus epiroticus epiroticus Pseudophoxinus stymphalicus marathonicus Paraphoxinus epiroticus prespensis Pseudophoxinus stymphalicus stymphalicus Pseudophoxinus stymphalicus thesproticus Leuciscus borysthenicus Ladigesocypris ghigii ghigii Gobio uranoscopus elimeius Appendix I (continued) In K&F2007 NEW 1 ENCE (Bloch, 1782) Kottelat, 2007 (Heckel, 1843) NO CHANGE 1 OTHER (Kessler, 1859) Karaman, 1924 (Linnaeus, 1758) (Linnaeus, 1758) (Karaman, 1924) (Karaman, 1924) Economidis, 1991 NO CHANGE 1 ENCE (Boulenger, 1896) NEW 1 OTHER (Basilewsky, 1855) NO CHANGE 1 ALN Economidis, 1973) (Gianferrari, 1927) (Richardson, 1845) NO CHANGE 1 ALN (Stephanidis, 1939) (Vinciguerra, 1921) (Steindachner, 1870) (Steindachner, 1896) (Steindachner, 1892) (Steindachner, 1896) (Valenciennes, 1844) (Heckel & Kner, 1858) (Kattoulas, Stephanidis & (Kottelat & Barbieri, 2004) NEW 1 ENWE Bogutskaya & Iliadou, 2006 (Temminck & Schlegel, 1846) NO CHANGE 1 ALN 16 15 17 Name Authority Previous Nomenclature Freshwater Endemic phoxinus cf. sp. Sperchios Rutilus panosi Rutilus prespensis Rutilus rutilus Rutilus Rutilus ylikiensis Phoxinus strymonicus Pseudorasbora parva Rhodeus amarus Rhodeus meridionalis Romanogobio elimeius Phoxinus Luciobarbus albanicus Leucaspius delineatus Luciobarbus graecus Pachychilon macedonicum Pachychilon pictum Parabramis pekinensis Pelasgus laconicus Pelasgus prespensis Pelasgus stymphalicus Pelasgus thesproticus Petroleuciscus borysthenicus Petroleuciscus smyrnaeus Pelasgus epiroticus Pelasgus marathonicus Hypophthalmichthys nobilis Ladigesocypris ghigii 71 72 73 74 75 66 67 68 69 70 65 52 51 53 54 55 56 58 60 61 62 63 64 57 59 49 12 3 4 56 50

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http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 09/03/2019 12:43:28 | (continued) 1 ENBAL 1111 ENWE ENBAL ENCE ENWE+ 11 ENWE ENWE 1 ENBAL NEW 1 OTHER Leuciscus cephalus vardarensis Leuciscus cephalus albus, Leuciscus "svallize" Leuciscus cephalus vardarensis Pseudophoxinus beoticus Phoxinellus pleurobipunctatus Leuciscus cephalus peloponnensis Leuciscus keadicus Leuciscus cephalus macedonicus Appendix I (continued) In K&F2007In K&F2007 NEW 1 ENCE (Fowler, 1977) NEW 1 NENNO (Heckel, 1837) NO CHANGE 1 ENBAL Karaman, 1924Karaman, 1928 NO CHANGE NEW 1 NENNO 1 ENBAL Karaman, 1928 & Nalbant, 1998 (Linnaeus, 1758) NO CHANGE 1 OTHER (Linnaeus, 1758) NO CHANGE 1 OTHER Stephanidis, 1937 NO CHANGE 1 ENCE Economidis, 1991 NO CHANGE 1 ENWE (Richardson, 1856) NEW 1 OTHER (Stephanidis, 1939) (Stephanidis, 1939) (Stephanidis, 1939) NO CHANGE 1 ENWE (Stephanidis, 1971) (Stephanidis, 1971) NEW(Stephanidis, 1939) NEW 1 1 ENWE ENWE (Valenciennes, 1844) (Schmidt-Ries, 1943) NO CHANGE 1 ENWE Erk’akan, Atalay-Ekmekçi Economidis & Nalbant, 1997 NO CHANGE 1 ENNO Economidis & Nalbant, 1997 NO CHANGE 1 ENNO Economidis & Nalbant, 1997 NO CHANGE 1 ENWE Economidis & Nalbant, 1997 NO CHANGE 1 ENWE Kottelat & Economidis, 2006 19 21 20 23 22 Name Authority Previous Nomenclature Freshwater Endemic 18 cii cf. sp. Evia sp. Aoos In K&F2007 sp. Evinos Cobitis stephanidisi Cobitis punctilineata Cobitis meridionalis Cobitis ohridana Cobitis puncticulata Cobitis hellenica Vimba melanops Cobitidae Cobitis arachthosensis Squalius Squalius Squalius Tropidophoxinellus spartiaticus Squalius vardarensis Telestes beoticus Telestes pleurobipunctatus Tinca tinca Squalius Tropidophoxinellus hellenicus Squalius keadicus Squalius moreoticus Squalius peloponnensis Scardinius acarnanicus Scardinius erythrophthalmus Scardinius graecus Squalius pamvoticus Squalius orpheus Squalius prespensis 102 101 98 99 100 97 95 96 79 87 86 94 89 90 91 92 88 93 81 82 80 12 3 4 56 76 77 78 84 83 85

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http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 09/03/2019 12:43:28 | (continued) 111 ENBAL 1 OTHER ENBAL ENBAL 111 ENBAL 1 ENBAL ENBAL NENNO 1ALN 1 ENBAL Orthrias barbatulus Orthrias brandti bureschi Orthrias pindus Sabanejewia aurata balcanica Salmo trutta macrostigma, Salmo trutta dentex Salmo trutta macedonicus Salmo trutta pelagonicus Salmo trutta peristericus Coregonus lavaretus Salmo trutta dentex Appendix I (continued) Heckel, 1852 Garman, 1890 NO CHANGE 1 ENWE Linnaeus, 1758 NO CHANGELinnaeus, 1758 NO CHANGE 1 OTHER 1 OTHER Karaman, 1928 NO CHANGE 1 ENBAL Karaman, 1938 Karaman, 1938 Karaman, 1955 NO CHANGE 1 ENBAL (Drensky, 1928) (Burchell, 1822) NEW 1 ALN (Linnaeus, 1758) (Linnaeus, 1758) (Karaman, 1922) (Karaman, 1924) NEW 1 ALN (Karaman, 1938) (Karaman, 1924) (Walbaum, 1792)(Walbaum, 1792) NO CHANGE NO CHANGE 1 1 ALN ALN Stephanidis, 1974 NO CHANGE 1 ENWE (Rafinesque, 1818) NEW 1 ALN (Economidis, 2005) Stoumboudi, Kottelat & Barbieri, 2006 NEW 1 OTHER 24 27 26 Name Authority Previous Nomenclature Freshwater Endemic lavaretus 25 cf. Nemacheilidae Barbatula barbatula Oxynoemacheilus bureschi Oxynoemacheilus pindus Oxynoemacheilus theophilii Ictaluridae Ictalurus punctatus Silurus glanis Siluridae Silurus aristotelis Clariidae Clarias gariepinus Esocidae Esox lucius Coregonidae Coregonus Salmo cf. macedonicus Sabanejewia balcanica Cobitis vardarensis Salmo letnica Salmo pelagonicus Salmo peristericus Oncorhynchus mykiss Salmo dentex Salmonidae Oncorhynchus kisutch Cobitis trichonica Cobitis strumicae Salmo farioides 107 108 109 110 111 113 112 114 115 116 122 106 105 121 123 124 118 119 117 12104 3 4 56 103 120

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http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 09/03/2019 12:43:28 | (continued) 0 OTHER (1) ALN Gasterosteus aculeatus Gambusia affinis NEW 0 ENWE Appendix I (continued) Risso, 1827 NO CHANGE 0 OTHER Risso, 1810 NO CHANGE 0 OTHER Risso, 1827 NO CHANGE 0 OTHER Cuvier, 1829 In K&F2007 NEW 1 ENWE Girard, 1859 Lesuer, 1821 NEW 1 ALN (Risso, 1827)(Risso, 1810) NO CHANGE NO CHANGE 0 0 OTHER OTHER (Risso, 1810) NO CHANGE 0 OTHER (Kessler, 1859) NO CHANGE (1) OTHER Linnaeus, 1758 NEW 1 ALN Linnaeus, 1758 NO CHANGE 0 OTHER Linnaeus, 1758 NEW 1 ALN (Mitchill, 1814) NO CHANGE 1 ALN (Sauvage, 1880) NO CHANGE (1) ENWE+ (Linnaeus, 1758) NO CHANGE 0 OTHER Stoumboudi, 2007 Stephanidis, 1971 NO CHANGE 1 ENCE Kottelat, Barbieri & (Valenciennes, 1821) NO CHANGE 0 OTHER 31 32 28 29 30 Name Authority Previous Nomenclature Freshwater Endemic 33 latipinna trutta cf. cf. sp. Louros Moronidae Dicentrarchus labrax Syngnathidae Pungitius platygaster Pungitius hellenicus Poecilia Salmo Cyprinodontidae Aphanius almiriensis Aphanius fasciatus Gasterosteidae Gasterosteus gymnourus Salmo Salmo salar Syngnathus abaster Valenciidae Valencia letourneuxi Poeciliidae Gambusia holbrooki Liza saliens Mugil cephalus Atherinidae Atherina boyeri Salvelinus fontinalis Mugilidae Chelon labrosus Liza ramada Liza aurata 143 142 141 137 127 138 139 140 126 125 12 3 4 56 144 135 136 132 133 134 128 129 131 130

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http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 09/03/2019 12:43:28 | ts. s K&F 2007) with 1 OTHER 1 ENBAL 1 OTHER Stizostedion lucioperca Zingel streber balcanicus Proterorhinus marmoratus Appendix I (continued) (Berg, 1916) NO CHANGE (1) OTHER (Asso, 1801) NO CHANGE (0) OTHER Ahnelt, 1991 NEW (1) ENWE (Holly, 1929) NO CHANGE (1) ENWE+ (Pallas, 1814) NO CHANGE 0 OTHER Kottelat, 2004 NEW 1 ENWE (Heckel, 1837) Linnaeus, 1758 NO CHANGE 1 OTHER (Linnaeus, 1758) (Linnaeus, 1758) NEW (1) ALN (Linnaeus, 1758) NO CHANGE (1) ALN (Linnaeus, 1758) NO CHANGE 0 OTHER (Karaman, 1936) (La Cepède, 1802) NEW (1) ALN (Vinciguerra, 1921) NO CHANGE (1) ENNO (Ahnelt & Bianco 1990) NO CHANGE (1) ENWE Economidis & Miller, 1990 NO CHANGE 1 ENWE sp. Louros, an operational name provided by this study). All species with valid name as well as undescribed operational taxa names are numbered and considered as separate taxonomic uni 35 Valid scientific name or operational taxonomic unit names are given. Eudontomyzon 34 Species author and date are given. Undescribed taxa are given only if provided by KOTTELAT & FREYHOF (2007a) (hereby referred a Name Authority Previous Nomenclature Freshwater Endemic one exception ( Economidichthys trichonis Knipowitschia caucasica Salaria fluviatilis Gobiidae Economidichthys pygmaeus Zingel balcanicus Sander lucioperca Cichlidae Oreochromis niloticus Blenniidae Salaria economidisi Centrarchidae Lepomis gibbosus Percidae Perca fluviatilis Micropterus salmoides Proterorhinus semillunaris Zosterisessor ophiocephalus Pleuronectidae Platichthys flesus Knipowitschia goerneri Knipowitschia milleri Knipowitschia thessala 154 155 152 153 149 148 150 151 12 3 4 56 145 147 146 159 160 161 156 157 158 Column 3: Authority. LEGEND Column 1: Species annotation. Column 2: Scientific Name.

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http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 09/03/2019 12:43:28 | e CRITICAL COMMENTS ON CHECK- LIST

1 Eudontomyzon hellenicus refers only to the populations in the Strymon R. An r in waters with high unidentified Eudontomyzon species has also been recorded from the Almopeos

not included in ECONOMIDIS tributary of the Aliakmon R. but it ).

al boundaries of Greece. remains undescribed (ECONOMIDIS Beotia region are included). southern part of Albania.

Attiki and Euboea). & B N RESCU, 1991). cies, brackish water or marine species that 2 Eudontomyzon sp. Louros is not a valid iour of these fish in Greece a provisional des- provisional a Greece in fish these of iour ly in Asia Minor and the adjacent Greek islands. Greek adjacent the and Minor Asia in ly taxa described for the Greek freshwater ichthyofauna (not referred to by an operational name by K&F 2007). The Eudontomyzon in the Louros basin is given this provisional name here since its distribution in Greece is extremely localized and this taxon has been known to differ morphologically from E. hellenicus for several years now (ECONOMIDIS, 1995). K&F 2007 remark that the population from Louros "either belongs to E. stankokaramani or represents a distinct, unnamed species". 3 Acipenser stellatus is considered as extirpated in the Aegean basins (K&F 2007), documentation of its past presence is evident in older publications (PAPACONSTANTINOU, 1988). Re- cent occurrences of the species in the Evros R. seem to be from stocking or escapes from Bulgaria (APOSTOLOU, pers. comm.; KOUTRAKIS & ECONOMIDIS, 2006). 4 Huso huso Subspecies replacement names and notation on changes with respect to ECONOMIDIS (1991) are as follows: NO CHANGE: No recent nam is now considered as an alien Fish known to be restricted to freshwaters in Greece are distinguished from species spending a part of their lives in the sea o species in Greece since all populations are from stocking and/or escapees from

level of endemic status at the national scale, i.e. restricted distributional range or endemic status relative to the territori fish farms (KOUTRAKIS & ECONOMIDIS, 2006). Unfortunately we cannot confirm the possible existence of natural populations of this sturgeon ENWE: Endemic to Western Greece and/or the Peloponnese. salinity. Typically freshwater species that usually do not readily enter seawater conditions are denoted with 1; Euryhaline spe behav the on exists uncertainty Where 0. with denoted are conditions freshwater in lives their of part spend or of tolerant are ignation is given in parentheses. NENNO: "Near-endemic" to Northern Greece; global populations restricted to Greece’s transboundary lakes (i.e. Prespa or Doirani ENNO: Endemic to Northern Greece; including Thessaly, but not the Pagasiticos Gulf (additionally two species that reach Attiko- ENBAL: Endemic to the Balkans (i.e. south of the Danube R.). ENAEG: Endemic to the Aegean Islands. on population localized a having species and species, widespread includes this patterns; distribution other with Species OTHER: ALN: Alien species; there is no substantiated evidence that the species is native within the Greek territory. Where doubts exist a provisional status is given in parentheses. ENWE+: "Near-endemic" to Western Greece; i.e. restricted to Greece, but a small fraction of global population extends into the ENCE: Endemic to Central Eastern Greece, i.e. the Attiko-Beotia region (south of Pagasiticos Gulf, including Fthiotis, Beotia, changes (i.e. species names retained as given by ECONOMIDIS, 1991); NEW: New additions to the (1991) Greek under the current freshwater taxon name. fish checklist, (or even its transient occurrence) in the northern Aegean in the past; although the species may possibly have existed up Column 6: Endemicity: Column 4: Previous Nomenclature. Column 5: "Freshwater residence".

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http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 09/03/2019 12:43:28 | until the 19th or early 20th centuries. A extreme southern part of the country few unconfirmed records in the Aegean (MARKOVA et al., 2007). are reported in earlier accounts 10 Barbus sperchiensis has isolated popula- (ONDRIAS, 1971; ECONOMIDIS, tions in the Sperchios valley and in 1973; PAPACONSTANTINOU, 1988) Northern Euboea (formerly considered but these have later been regarded as Barbus cyclolepis sperchiensis), the Paga- extremely doubtful (KOUTRAKIS & siticos Gulf’s Cholorema basin (former- ECONOMIDIS, 2006). ly considered B. cyclolepis choloremati- 5 The existence of Alburnus scoranza is cus) and in Thessaly (formerly consid- undocumented in Greece. A species ered Barbus cyclolepis strumicae) regarded as Alburnus alburnus was (ECONOMIDS & BOGUTSKAYA, detected within the Greek part of the 2003). This unusual distribution strad- Aoos (ECONOMOU et al., 2007) and is dles a biogeographic boundary and tentatively given the provisional opera- includes insular populations on Euboea tional name A. cf. scoranza here, since Island. the only Alburnus species in the immedi- 11 Carassius auratus is frequently confused ate vicinity and within the Southern in the literature with Carassius gibelio, Adriatic biogeographic region is A. sco- which was formerly treated as "a wild ranza (K&F 2007). feral form of C. auratus" (K&F 2007). 6 Alburnus sp. Volvi is an unnamed taxon The taxonomic entity of Carassius aura- very similar to Alburnus alburnus. Speci- tus (the goldfish) is not present as an mens only from L. Volvi and L. Kerkini independent taxonomic unit in (Strymon basin) were collected in a first ECONOMIDIS (1991) although it is provisional description (FREYHOF & present in BOBORI & ECONOMIDIS KOTTELAT, 2007a). (2006). 7 Alburnus thessalicus belongs to the 12 The status of Carassius gibelio with Alburnus alburnus complex with four respect to its being native in Greece is taxa in Greece (A. thessalicus, A. mace- still unresolved and controversial; we donicus, A. sp. Volvi, A. alburnus) (K&F adopt its status "as probably native" only 2007). These taxa share many character- in the Strymon and the Evros following istics and may be difficult to identify in ECONOMIDIS (1991). There is consid- the field. erable doubt as to this species native sta- 8 Barbus balcanicus populations in the tus in southeast Europe, although popu- lower Axios were formerly considered as lations have long been naturalized. both Barbus peloponnesius petenyi and 13 The status of Cyprinus carpio with Barbus cyclolepis; difficulty of identifica- respect to its being native to Greece is tion of these fish on morphological unclear; natural populations have prob- grounds is noted in K&F 2007, but the ably been established and are natural- species can be distinguished using ized for centuries. We adopt this species molecular markers (KOTLI’K et al., status as "native" in Thessaly, Macedo- 2002). nia, and Thrace according to 9 Barbus peloponnesius is not recorded in ECONOMIDIS (1991). Albania following K&F 2007, but there 14 The existence of Gobio skadarensis is is recent evidence of its existence in the undocumented in Greece. The species

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http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 09/03/2019 12:43:28 | found inhabiting the Aoos (STOUMBOUDI et al., 2006) although (ECONOMOU et al., 2007) is tentative- K&F 2007 refers to this population as ly given the operational name Gobio cf. Squalius cf. cii. Unidentified Leucisinae skadarensis in this list since the only called "Leuciscus cephalus" (BIANCO, known Gobio sp. in the immediate vicin- 1990) also exist on Samos. ity and in the South Adriatic biogeo- 19 The identity of the Squalius in the Ionian graphic region is Gobio skadarensis, pre- river basins is controversial and unre- viously referred to as Gobio gobio alban- solved. Formerly these chubs were given icus ( ANDA et al., 2005). as Leuciscus cephalus but it is certain that 15 Phoxinus phoxinus was until recently the the chubs west of the Pindos are differ- only species in this genus in Greece. ent species. In the distributional compi- K&F 2007 present a new Phoxinus lation we refer to nearly all "Ionian species in the Strymon (Phoxinus stry- chubs" by the provisional operational monicus), but no classification is provid- name Squalius cf. peloponnensis ed for the Phoxinus populations of Ali- although we do give the specific three akmon, Loudias, Nestos, Filiouris and taxa proposed by K&F 2007 in the spe- Evros; although they state that "Phoxi- cific recorded basins. Despite consider- nus populations in the Loudias and Fil- able amount of genetic work iouris drainages are possibly conspecif- (IMSIRIDOU et al., 1997, 1998, 2000; ic". The aforementioned authors do not DOADRIO & CARMONA, 1998, give Phoxinus phoxinus as inhabiting any 2003; SANJUR et al., 2003; ZARDOYA area in the southern Balkan river basins et al., 1999; DURAND et al., 1999b, or in Greece. Pending an accurate 2000), further taxonomic research description of the unclassified popula- involving populations from northern and tions we propose the fish not in the type south-western Peloponnese as well as locality river (Strymon) be provisionally from the Epirus rivers is needed to termed Phoxinus cf. phoxinus. resolve the problem of Squalius species 16 Romanogobio elimeius may possibly distinction in the Ionian basins. include a very similar-looking rheophilic 20 Squalius moreoticus described as Leucis- gudgeon called Romanogobio kessleri cus cephalus moreoticus by banarescui (ECONOMIDIS, 1991); STEPHANIDIS (1939) was originally K&F 2007 were "unable to dististinguish considered as being confined to Lake R. banarescui and R. stankovi from R. Stymphalia. K&F 2007 give a wider dis- elimeius on the basis of the available lit- tribution for this taxon (i.e. including erature and material and therefore ten- Vouraikos R. in the northern Pelopon- tatively treat them as synonyms". nese). We tentatively limit the S. 17 Rutilus sp. Sperchios was observed in moreoticus distribution to the aforemen- the lower Sperchios in the late ‘90s but tioned lake but its current status is unde- was never described. Only two small- fined, as the originally described taxon sized specimens were collected in 1997 may be extirpated or even extinct. The (K&F 2007); this taxon needs immedi- lake completely dried-out in the early ate study. 1990s and the present population hosted 18 Squalius cii was first proposed as a ten- in the lake may have been introduced tative name for the Squalius of Lesbos from another basin – as purported by

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http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 09/03/2019 12:43:28 | local fisherman. Greece needs confirmation. 21 Squalius pamvoticus was described by 26 Salmo farioides is now formally consid- STEPHANIDIS (1939) as Leuciscus ered as the dominant trout of western cambeda var. pamvoticus presumably Greece, a different taxon from popula- confined to Lake Pamvotis. K&F 2007 tions east of the Pindos. The population give a wider distribution for this taxon, of the Alfios (Peloponnese) are isolated including nearly all the rivers of Epirus and in need of conservation-relevant (except Aoos). Care is needed in the taxonomic confirmation (K&F2007). classification of populations in the rivers 27 Salmo macedonicus is not recorded as a of Epirus; and pending further taxo- species inhabiting the Greek territory in nomic evidence we treat the Epirus river K&F 2007. These authors do not give taxa as Squalius cf. peloponnensis distributional information or the identi- (STEPHANIDIS, 1939 considered ty of trout in the Strymon, Nestos or these populations as Leuciscus cephalus Evros; however, in previous works the peloponnensis). The taxonomy and dis- taxon S. trutta macedonicus has been tribution of Squalius pamvoticus needs described in these areas (KARAMAN, immediate attention. 1927). We use the tentative name of S. 22 Squalius sp. Evia was previously consid- cf. macedonicus to refer to the fish ered as a distinct form of Leuciscus inhabiting the aforementioned basins. cephalus vardarensis. This taxon name is Confirmation of this taxon’s presence provisionally given for specimens from and its distribution in Greece is needed. the Manikiotikos R. on Euboea island. 28 Salmo sp. Louros is one of the most sur- The classification of Squalius popula- prising new "taxa" described from the tions in the streams of northern Euboea Louros R. DELLING (2003) refers to remains unresolved (although we tenta- this fish as Salmo louroensis a name not tively place the fish from the Kireas R. accepted by K&F 2007. under this taxon name in the distribu- 29 Varieties of farmed Salmo trutta from tional accounts). Evia should be spelled European hatcheries have been intro- Euboea, the anglicised classical rendi- duced into Greek waters particularly tion often used in the scientific bibliog- near fish-farming units. raphy. 30 The systematic status of Atherina boyeri 23 Squalius sp. Evinos was proposed as a populations has been recently ques- tentative taxonomic unit by K&F 2007 tioned. Molecular data (KLOSSA- for Squalius populations inhabiting KILIA et al., 2007) reveal deep genetic Mornos, Evinos and Acheloos. This divergence between marine populations taxon is morphologically very similar to of Atherina boyeri and those living in some populations of Squalius pelopon- lagoons and lakes, possibly indicating nensis. the existence of cryptic or sibling 24 Coregonus cf. lavaretus requires identifi- species. cation due to recent taxonomic changes 31 Gambusia holbrooki has replaced for- in K&F 2007. mer claims of Gambusia affinis in 25 Salmo dentex is mentioned as inhabiting Greece. K&F 2007 state that there is no Aoos (and possibility Alfios) in confirmed record of G. affinis in K&F2007. This species’ existence in Europe.

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http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 09/03/2019 12:43:28 | 32 Poecilia cf. latipinna has tentatively been territory but records are from very near identified as the alien aquarium escapee the border in FYR Macedonia. This which is abundant in Vouliagmeni L., species is difficult to collect and it is Attiki (KOUTSIKOS pers. comm.); for- assumed that it is almost definitely pres- merly this species was described as Poe- ent in Greek waters, probably as far as cilia sphenops (CHINTIROGLOU et Axioupolis, as stated by K&F2007. The al., 1996). species is also included within the fresh- 33 The marine family Syngnanthidae is water fish faunal list of BOBORI & poorly documented in freshwaters in ECONOMIDIS (2006). Greece. BOBORI & ECONOMIDIS 35 The Knipowitschia population in the (2006) provide five species of this family lower Evinos R. was formally referred to in their freshwater fish fauna list. Pend- as Knipowitschia panizzae (AHNELT & ing new information that confirms that BIANCO, 1991) but this is uncon- these species actually reside for long firmed; perhaps an unidentified taxon periods in freshwaters we retain only (or taxa) of Knipowitschia exists in sev- Sygnathus abaster in this list, as did eral areas of western Greece (MILLER ECONOMIDIS (1991). et al., 2004b). 34 To our knowledge Zingel balcanicus has not yet been collected within the Greek

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